CONTENTS

CLARKE, P. A.

Myth as History? The Ngurunderi Dreaming of the Lower Murray, South Australia.

CRAIG, B

Following the tracks of Edgar Waite in New Guinea for the Pacific Arts Symposium

in Adelaide.

GOWLETT-HOLMES, K. L. & ZEIDLER, W.

Mollusc type specimens in the South Australian Museum. 6. Additions and corrections

to Part 1 (Cephalopoda and Scaphopoda) and Part 3 (Polyplacophora).

HARTMANN-SCHRODER, G. & PARKER, S. A.

Four new species of the family Opheliidae (Polychaeta) from southern Australia.

HUTCHINSON, M. N. & RAWLINSON, P. A.

The water skinks (Lacertilia: Eulamprus) of Victoria and South Australia.

JONES, P. G.

‘A special inducement’: The establishment of the Records of the South Australian Museum.

JONES, P. G.

Obituary — Norman Barnett Tindale

VAN VONDEL, B. J.

Revision of the Haliplidae (Coleoptera) of the Australian region and the Moluccas.

WASSON, K.

The kamptozoan Pedicellina whiteleggii Johnston & Walker, 1917 and other pedicellinids

in Australia and New Zealand.

WATTS, C. H. S.

Revision of the Australasian genera Agraphydrus Regimbart, Chasmogenus

Sharp and Helochares Mulsant (Coleoptera: Hydrophilidae).

WILLIAMS, G. C.

Revision of the Pennatulacean genus Sarcoptilus (Coelenterata: Octocorallia), with

descriptions of three new species from southern Australia.

YENGOYAN, A.

Obituary — Joseph B. Birdsell.

Volume 28(1) was published on 30 October 1995.

Volume 28(2) was published on 29 December 1995.

ISSN 0376-2750

PAGES

143-157

33-52

103-111

185-207

53-59

159-176

61-101

131-141

113-130

13-32

177-183

FOUR NEW SPECIES OF THE FAMILY OPHELIIDAE (POLYCHAETA) FROM.

SOUTHERN AUSTRALIA

G. HARTMANN-SCHRODER & S. A. PARKER*

HARTMANN-SCHRODER, G. & PARKER, S. A. 1995. Four new species of the family

Opheliidae (Polychaeta) from southern Australia. Records of the South Australian Museum

28(1): 1-12.

Recent examination of specimens of Euzonus, Ophelia, Ophelina and Travisia (Polychaeta:

Opheliidae) in the South Australian Museum revealed a previously undescribed species in each

genus: Euzonus zeidleri sp. nov., Ophelia bulbibranchiata sp. nov., Ophelina longicirrata sp.

nov. and Travisia oksae sp. nov. The previous report of Travisia forbesii Johnston, 1840 from

Victoria is referred to T. oksae sp. nov. T. oksae sp. nov. is known from South Australia and

Victoria, the other three from South Australia only. Euzonus zeidleri sp. nov. represents the

first record of its genus for Australia

G. Hartmann-Schréder, Zoologisches Institut und Zoologisches Museum, Martin-Luther-King-

Platz 3, 20146 Hamburg, Germany, and S. A. Parker, South Australian Museum, North Terrace,

Adelaide, South Australia 5000. Manuscript received 24 May 1993.

The family Opheliidae is widely distributed in

the seas of all climatic zones from the polar

regions to the tropics, and from shallow waters to

abyssal depths. Most members of the family live

as deposit-feeding burrowers in various types of

sediment.

Six genera and 15 named species of opheliids

have so far been reported from Australia (mainly

from southern Australia and New South Wales):

Armandia Filippi, 1861: A. maculata (Webster,

1884), A. intermedia Fauvel, 1902, A.

secundariopapillata Hartmann-Schréder, 1984, A.

bilobata Hartmann-Schroder, 1986; Ophelia

Savigny, 1818: O. ashworthi Fauvel, 1917, O.

dannevigi Benham, 1916, O. elongata Hutchings

& Murray, 1984, O. multibranchia Hutchings &

Murray, 1984, Lobochesis Hutchings & Murray,

1984: L. bibrancha Hutchings & Murray, 1984,

L. longiseta Hutchings & Murray, 1984; Ophelina

Orsted, 1843: O. breviata (Ehlers, 1913)!, O.

gigantea (Rullier, 1965); Polyophthalmus

Quatrefages, 1850: P. pictus (Dujardin, 1839);

Travisia Johnston, 1840: T. lithophila Kinberg,

1866, T. olens Ehlers, 1897 (Day & Hutchings

1979, Hutchings 1982, Hutchings & Murray 1984,

Hartmann-Schréder 1980, 1984, 1985, 1986). A

further species, Travisia forbesii Johnston, 1840,

has been reported from Victoria (Poore et. al.

1975), but the specimens proved, upon

examination, to have been misidentified (see

below under T. oksae sp. nov.). A seventh genus,

Euzonus Grube, 1866, has to date been reported in

* Shane Parker died on 21 November 1992.

' Listed as Ophelia breviata by Day & Hutchings (1979:129)

Australasia only from New Zealand (E. otagoensis

Probert, 1976).

MATERIALS AND METHODS

The present study is based largely on material

in the South Australian Museum, viz. specimens

of Ophelia and Ophelina obtained at Pearson

Island in June 1973, a large series of Travisia

from upper Spencer Gulf collected by E. Oks of

the South Australian Fisheries Department in

1985-1987, and a large series of Euzonus

collected at Reevesby Island in the Sir Joseph

Banks Group in 1985 and 1986 by the junior

author. This new material was augmented by a

loan of Travisia from the NMV. Comparative

material of previously described species included

the four (of six) syntypes of Ophelia ashworthi in

the SAM, and from the ZMH the syntypes of

Euzonus furciferus (Ehlers, 1897), and the

specimens of 7. forbesii. The subfamilial and

generic classification follows Hartmann—Schroder

(1971).

Measurements are in millimetres, made with an

eyepiece graticule. Drawings were executed with

the aid of a camera lucida on a Zeiss microscope.

Abbreviations of institutions mentioned are: AM,

Australian Museum, Sydney; BMNH, Natural

History Museum, London; NMV, Museum of

Victoria, Melbourne; NTM, Northern Territory

Museum, Darwin; SAM, South Australian

Museum, Adelaide; USNM, National Museum of

Natural History, Washington DC; ZMH,

Zoological Museum, Hamburg. Material is depos-

ited in the AM, NTM, SAM, USNM and ZMH.

2, G. HARTMANN-SCHRODER & S. A. PARKER

SYSTEMATICS

Family OPHELIIDAE Malmgren, 1867

Subfamily OPHELIINAE Malmgren, 1867

Genus Euzonus Grube, 1866

Euzonus zeidleri sp. nov.

(Figs 1-5)

Types (all collected on Reevesby Island, South

Australia, by S. A. Parker, measurements for

paratypes are of length of longest specimen in

each sample).

Holotype: Haystack Bay, 20.1.1986, SAM

E2145; length 28 mm, width (at thorax) 3.5 mm.

Paratypes (276 specimens): Haystack Bay,

21.14.1985, SAM 2146(36), 23 mm; Haystack Bay,

311.1985, SAM E2147(58), 22 mm; McCoy Bay,

221.1985, SAM E2148(36), 31 mm; McCoy Bay,

311.1985, SAM E2149(2), 19 mm; Haystack Bay,

20.1.1986, SAM E2150(73)/ZMH: P20816(6)/

USNM 169135 (4)/AM W21745 (4)/ NIM

W6421-424(4), 27 mm, Haystack Bay, 22.1.1986,

SAM E2151(14), 35 mm, Haystack Bay,

30.1.1986, SAM E2152(9), 28 mm; Haystack Bay,

1.11.1985, SAM E2153(26), 24 mm:

Other material examined

Euzonus furciferus (Ehlers, 1897): syntypes,

Punta Arenas, Strait of Magellan, ZMH:

V4869(7); Itanhan, Brazil, ZMH: P15144(1);

Santos, Brazil, ZMH: P15421(4); Zapallar, Chile,

ZMH: P15124(1).

Diagnosis

A Euzonus with bifurcate branchiae (character

of subgenus Thoracophelia Ehlers, 1897 sensu

Hartman 1956), with body formula 12a + 20b +

6a (i.e. branchiae occuring on setigers 13-32); all

branchiae bifurcate; branches of branchiae

subequal, superior one bearing 1—2 small pinnae;

setae on segments 1-36; pygidium relatively large,

broad, lacking elongated terminal (ventral) cirrus,

6-9 lateral cirri on each side.

Description

Body fusiform, 38 segments, 36 setigers, total

length 9-35 mm; divided into three distinct

regions: cephalic (prostomium and two setigers),

thoracic (eight setigers) and abdominal (26

setigers and two asetigerous segments). All

segments annulated. Prostomium with small

apical point; eyes absent; nuchal pits present (Fig.

1). Thoracic region inflated, delimited from

cephalic region by pronounced constriction,

delimited from abdomen by thick, smooth lateral

glandular ridges. Abdominal region comprising

two prebranchial, 20 branchiate and six

postbranchial segments (body formula thus 12a +

20b + 6a, see Tebble 1952). Last two asetigerous

abdominal segments with close-set longitudinal

furrows. Pygidium wide at base, broadly rounded

at tip, not tapering to an elongated ventral cirrus;

dorsal anal cirri disposed in a V over pygidium,

6-9 on each side (Fig. 5). Abdominal region

bearing deep longitudinal midventral and lateral

grooves, former continuing to pygidium, latter

becoming shallower beyond last branchiate setiger

(32nd).

Branchiae on setigers 13-32, bifurcate,

branches subequal, superior branch bearing 1-2

small pinnae distally on dorsal surface (Fig. 2-4).

Parapodia barely or not visible, lobes immersed

in surrounding tissue, from which bundles of setae

appear to arise directly. Notosetae and neurosetae

simple, capillary, 6-25 and 6-30 setae per bundle

respectively, on segments 1-36. Neurosetae

shorter than notosetae of same setiger, except on

setigers 34-36, where they increase to a subequal

length; also, setae of setdgers 34-36 markedly

longer than those of preceding setigers, tending to

curve and spread conspicuously (Fig.5).

Colour of individuals in life bright red,

especially at anterior end.

Comparisons with other species

Eleven species have been described to date

(Rozbaczylo & Zamorano 1970, Probert 1976). Of

these, six fall in the subgenus Thoracophelia

Ehlers, 1897, sensu Hartman 1956, having the

branchiae bifurcate rather than trifurcate or

pectinate: E. furciferus (Ehlers, 1897, type species

of Thoracophelia), E. mucronata (Treadwell,

1914), E. williamsi (Hartman, 1938), E.

profundus Hartman, 1967, E. otagoensis Probert,

1976 and E. heterocirrus Rozbaczylo &

Zamorano, 1970.

Within this group, E. zeidleri sp. nov. is most

similar to E. furciferus of South America, the only

member of the subgenus with a body formula of

12a + 20b + 6a, i.e. twelve abranchiate anterior

segments, 20 branchiate segments and 6 posterior

abranchiate segments. The closest species

geographically, E. otagoensis of New Zealand,

differs in having a body formula of 14a + 18b +

6a. From E. furciferus, E. zeidleri differs in

possessing 1—2 small pinnae on the superior

branch of the branchiae (absent in E. furciferus),

FOUR NEW SPECIES OF OPHELIIDAE 3

geeee 1eKky SoU

2mm 5

FIGURES 1-5. Euzonus zeidleri sp. nov. 1, anterior end, lateral view; 2, segments from middle of body, lateral view

(setae omitted); 3, median body region showing branchiae; 4, branchiae, detail; 5, posterior end, lateral view (all of

paratype ZMH: P—20816).

4 G. HARTMANN-SCHRODER & S. A. PARKER

the last two segments asetigerous (last one

asetigerous in E. furciferus), last 3-4 segments

not telescoped together as in E. furciferus,

pygidium relatively large, broad, bluntly rounded

at tip, not small and tapering to elongated ventral

cirrus as in E. furciferus, and 6—9 dorsal cirri on

each side vs 4 on each side in E. furciferus.

Etymology

We name this species in honour of our

colleague Wolfgang Zeidler, South Australian

Museum, who led the 1985 and 1986 collecting

trips to the Sir Joseph Banks Group.

Distribution and ecology

Euzonus zeidleri is known only from Haystack

Bay and McCoy Bay, two quartz-grain surf-

beaches on the eastern, weather side of Reevesby

Island, Sir Joseph Banks Group, South Australia,

where it was collected in damp sand of the lower

and middle intertidal, within a spade’s depth of

the surface. Both beaches squeaked underfoot, and

thus belong to the type known as singing beaches

(for a discussion of this phenomenon, apparently

produced by the shearing of the well-sorted, well

rounded quartzgrains under pressure, see the

account of Squeaky Beach, Victoria by Beasely

1972).

Subsequent to the discovery of E. zeidleri, S.A

P. sought the species in other exposed sandy

beaches, e.g. on Younghusband Peninsula and at

Cape Jaffa, Boatswain’s Point and Robe, south-

eastern South Australia. None of these contained

E. zeidleri, and none squeaked underfoot; all were

of silicate rather than quartzgrains. In answer to

our enquiry, Dr P. K. Probert (in litt. 11.11.1992)

confirmed our suspicion that the type locality of

Euzonus otagoensis Probert, 1976 (Allan’s Beach,

Otago Peninsula, New Zealand) was a singing

beach. It thus seems possible that the occurence of

E. zeidleri and E. otagoensis (and perhaps other

species of the genus) coincides with that of

singing beaches, with their characteristic well-

sorted, well-worn quartzgrains.

E. zeidleri was more abundant at Haystack Bay

than at McCoy Bay; at the latter locality the

quartzgrains averaged larger but were equally

rounded. The only other macroscopic species

observed in these beaches were the cosmopolitan

marine acanthodriline oligochaete Pontodrilus

litoralis (Grube, 1855; first South Australian

record) and a minute enchytraeid oligochaete;

these were collected at Haystack Bay, from less

damp sand of the upper subtidal.

Genus Ophelia Savigny, 1818

Ophelia bulbibranchiata sp. nov.

(Figs 6-9)

Types

Holotype: Pearson Island, Investigator Group,

South Australia, 26.vi.1973 (?coll.), SAM E1604,

30 setigers, length 30 mm, width 4 mm.

Paratypes: Same data as holotype: SAM

FIGURES 6-9. Ophelia bulbibranchiata sp. nov. 6, anterior end, lateral view; 7, posterior end, lateral view; 8, first

branchia from anterior segment; 9, parapodium from middle of body.

FOUR NEW SPECIES OF OPHELIIDAE 5

E1605(3), 30 setigers, length 2-27 mm; ZMH:

P20667(1), 30 setigers, length 30 mm.

Other material examined

Ophelia ashworthi Fauvel, 1917; ‘St Vincent

and Spencer Gulfs’, South Australia, SAM E306

(four syntypes).

Diagnosis

An Ophelia 24-33 mm long with 30 setigers

(including 10 prebranchial and 5 postbranchial),

15 pairs of simple, bulbous-based branchiae, 7—9

small and two large anal papillae; posterior

dorsolateral ridges absent.

Description

Body club shaped, divided into anterior region

of nine setigers and a posterior region of 21

setigers; a faint constriction between setigers 2

and 3. Prostomium small, conical, with small

nuchal organs basally. No eyes visible. All

segments annulate, those of anterior region

somewhat areolate (Fig. 6); last five to six

segments decreasing in size. Lateral furrows and a

ventral furrow from setiger 11. Posterior segments

lack dorsolateral ridges. Branchial fenestrations

absent. Pygidium cylindrical with longitudinal

folds or furrows, bearing seven to nine small

dorsal and lateral anal papillae and two large

ventral papillae (Fig. 7). First bundle of setae very

small, easily overlooked, in region of mouth (Fig.

6). Subsequent parapodia with short, broadly

rounded, postsetal lobes (Fig. 9). Setae capillary,

notapodial slightly longer than neuropodial but

nowhere obviously long. Fifteen pairs of branchiae

from setiger 11 to setiger 25. Branchiae short, with

bulbous bases, latter largest in middle branchiae

(Fig.8-9). Nephridial pores not visible.

Comparisons with other species

O. bulbibranchiata sp. nov. is unique among

the known species of the genus in possessing

bulbous bases to the branchiae.

Species of Ophelia with a similar number of

setigers to O. bulbibranchiata (30) are O.

multibranchia Hutchings & Murray, 1984 (27),

O. peresi Bellan & Picard, 1965 (29), O. celtica

Amoureux & Dauvin, 1981 (29), O. elongata

Hutchings & Murray, 1984 (30), O. bipartita

Monro, 1936 (31), O. dannevigi Benham, 1916

(32) and O. ashworthi Fauvel, 1917 (32). Of

these, O. peresi, O. celtica and O. bipartita have

not been recorded in Australia. For differences see

Table I.

Etymology

The epithet bulbibranchiata (L.), refers to the

bulbous shape of the basal part of the branchiae,

unique in the genus.

TABLE 1. Comparison of Ophelia bulbibranchiata sp. nov. with other species of the genus with similar number of

setigers

Number of sizein number of number of pairs shape of anal posterior

setigers mm prebranchial _postbranchial _ of branchiae papillae dorsolateral

segments segments branchiae ridges

O. multibranchiata 27 4-6 7 3 17 simple 10 small absent?

ones

O. peresi' 2) 1~12 10 4 15 simple 12 small absent

ones

O. celtica ' 29 30-45 10 3 16 simple 12 small absent

+ 2 large ones

O. elongata 30 4-7 8 6 16 simple 10 small present

ones

O. bipartita ! 33) 63 9 5 17 simple 16 small present

+ 2 large ones

O. dannevigi 32 20 10 3 19 simple 16 small absent?

+ 2 large ones

O. ashworthi 32 ? 10 yy 20 bifurcate several small absent?

+ | large one

O. bulbibranchiata 30 24-33 10 5 15 bulbous 7-9 small present

base + 2 large ones

‘no records from Australia

6 G. HARTMANN-SCHRODER & S. A. PARKER

Distribution and ecology

Known only from the type series obtained at

Pearson Island, eastern Great Australian Bight in

1973. No notes on depth or habitat accompany the

specimens.

Subfamily OPHELININAE Hartmann-Schréder,

1971

Genus Ophelina Orsted, 1843

Ophelina longicirrata sp. nov.

(Figs 10-12)

Types

Holotype: Pearson Island, Investigator Group,

South Australia, 26. vi. 1973 (?coll), SAM E1606,

4] setigers, length 52 mm, width 3.0 mm.

Other material examined

Ophelina breviata (Ehlers, 1913): holotype,

FIGURES 10-12. Ophelina longicirrata sp. nov. 10,

anterior end, lateral view; 11, posterior end, lateral view

(cuticulum lifted away from underlying tissue through

fixation); 12, parapodia and branchiae of segments 34

and 35.

ZMH: V8583, Kaiser-Wilhelm-II-Land,

Antarctica, P190559(10), P19728(3), Antarctica;

ZMH V9548(2), Bass Strait, ZMH: V11871(1),

Port Lockray, New South Wales (Augener 1927:

DSF aes 2051S, 15020518).

Diagnosis

A large Ophelina (52 mm long), with 41

setigers, long cirriform presetal lobes on anterior

parapodia, long ventral cirri, anal tube cylindrical.

Description

Body fusiform, 41 setigers. Prostomium conical,

longer than wide at base, with oblong palpode

(Fig. 10). Proboscis a folded bag. No eyes visible.

Nuchal organs slightly protruded, nuchal slits

probably horseshoe-shaped. Segments strongly

annulate. Anal tube cylindrical, annullate, only

half width of last segments and as long as 5-6 last

segments together. Anal opening ventroterminal,

with 11 lanceolate anal papillae and a ventral

cushion that may be base of a lost unpaired cirrus

(Fig. 11). Presetal lobes of anterior parapodia long,

cirriform, half as long as the branchiae on setiger

2, decreasing in size to setiger 12. From setiger

26, presetal lobes rounded-conical, with digitate

extension of same length as ventral cirrus (Fig.

10-11). Ventral cirrus of setiger 1 small, filiform,

increasing in size to setiger 14, thence of same

length as presetal lobe (Fig. 11-12). Setae

nowhere obviously long. Branchiae long, cirriform

to filiform, overlapping on dorsum, absent only

from setiger 1.

Comparisons with other species

Species of Ophelina with a cylindrical anal tube

and cirriform presetal lobes are O. hachaensis

Augener, 1934 and O. fauveli (Caullery, 1945),

neither of which has been reported from Australia.

O. longicirrata sp. nov. is much larger than O.

hachaensis, with fewer setigers (52 mm, 41

setigers, vs 9 mm, 48 setigers), longer ventral

cirri’, and anal tube equal to the last 5-6 segments

(vs the last 3-4 in O. hachaensis). O. fauveli has

only 31 setigers and a length of 20 mm. The anal

tube of both species is much shorter than that of

O. longicirrata.

Besides O. longicirrata, two other species of

Ophelina have been reported from Australia, O.

breviata (Ehlers, 1913) of Bass Strait, New South

Wales and Antarctica and O. gigantea (Rullier,

1965) of Queensland. O. breviata differs from O.

' Augener (1934) misinterpreted the long presetal lobe of

O.hachaensis as a ventral cirrus.

FOUR NEW SPECIES OF OPHELIIDAE 7

longicirrata by its smaller size (28 setigers, 29

mm length). In O. breviata the branchiae are

absent from the first and the last four segments,

the posterior four segments are very short, and the

presetal lobes are rounded rather than filiform. O.

gigantea differs from the new species (and from

all the other species discussed) in having the anal

tube not cylindrical but spoon-shaped, open

ventrally for the whole of its length. It is also

larger (62-64 mm), with more setigers (65-68).

Etymology

The epithet longicirrata (L.) refers to the long

cirriform ventral cirri.

Distribution and ecology

Known only from the unique holotype obtained

at Pearson Island, eastern Great Australian Bight

in 1973. No notes on depth or habitat accompany

the specimen.

Subfamily TRAVISIINAE Hartmann-Schroder,

1971

Genus Travisia Johnston, 1840

Travisia oksae sp. nov.

(Figs 13-19)

Travisia forbesi: Poore, Rainer, Spies & Ward

(non Johnston), 1975: 29, 56 (Port Phillip Bay,

Victoria); Day & Hutchings (non Johnston), 1979:

129 (pars, Victoria).

Types

Holotype: Station 2, 32°35'04"S, 137°46'08"E,

upper Spencer Gulf, South Australia 6 m, medium

sand, coll. E. Oks, S. Aust. Fisheries Dept.

x1.1985, SAM E2701, 27 segments, 24 setigers,

length 15 mm, width 5.5 mm (moderately

contracted).

Paratypes: (52 specimens, all collected by E.

Oks, upper Spencer Gulf, 1985-1987): Station 2,

SAM E2702 (4, in same sample as holotype) (27,

25, 20.5), SAM E2703(2) (27, 24, 12), SAM

E2704(1) (26, 24, 8), SAM E2705(1) (26, 26, 5);

Station 3, SAM E2706(1) (26, 25, 5), SAM

E2707(4) (26, 24, 7), SAM E2708(6) (23, 20, 3),

SAM E2718(1) (27, 24, 11); Station 4, SAM

E2709(1) (26, 24, 4); Station 5, SAM E2710(3)

(25, 25, 4.5); Station 6, SAM E2711(2) (26, 25,

9.5), SAM E2712(7)/USNM 169136 (1)/AM

W21746 (1)/(26, 24, 9.5); Station 7, SAM

E2713(1) (27, 25, 24); Station 8, SAM E2714(2)

(Qi, 2, ND), ZAMIR PADD) Ci, 2D, IDE

Station 9, SAM E2715(1) (26, 22, 4.9), SAM

E2716(7)/NTM W6425(1) (27, 24, 16.5), SAM

E2717(1) (26, 23, 6), ZMH: P20669(3) (27, 24,

155) (figures in parentheses after each sample

refer to the number of segments, number of

setigers and total length of the largest individual

in the sample).

Other material examined

T. oksae sp. nov.: Victoria: Port Phillip

Environmental and Benthic survey: Stn 974,

13.x.1971, NMV F60027(2), 60028(5), 60029(3)

(Poore et al. 1975, under T. forbesi) Western Port

Environmental Study: Stn 1704, NMV F60035(2),

Stn 1722, NMV F60033(5), 60037(2), 60043(3),

Stn 1723, NMV F60030(1), 60040(8), Stn 1724,

NMV F60036(1), 60039(3), 60044(1), Stn 1727,

NMV F60041(1), Stn 1731, NMV F60042(1), Stn

1733, NMV F60031(1), Stn 1735, NMV

F60034(2), 60038(3) (ix. 1973, i. 1974).

T. olens Ehlers, 1897: syntypes, Punta Arenas,

Chile, ZMH: V4865(7), 4866(4), 4807(20),

4868(1); Punta Arenas, ZMH: V11937(1), ZMH:

PE1031(6); South Georgia, ZMH: V11877(1);

Antarctica, ZMH: P19085(9); Exmouth Gulf,

Western Australia, ZMH: P16892(1) (Hartmann-

Schroder 1980: 74).

T. forbesi Johnston, 1840: North Sea, ZMH:

V9190(5); Bay of Kiel, Baltic Sea, ZMH:

P19241(22); Bohuslan, Sweden, ZMH: V5355(4);

Tromso, Norway, ZMH: V7576(4), ZMH:

PE1021(1); Spitzbergen, ZMH: PE1023(1); Jan

Mayen, ZMH: V7527(1); Franz-Joseph-Land,

ZMH: PE1025(4); Murman Coast, ZMH:

V1439(2).

Diagnosis

A Travisia with 23-27 segments and 20-25

setigers, notopodial lobes from segment 15,

neuropodial lobes from segment 16, and setae

smooth, bilimbate.

Description

Body fusiform, 23-27 segments, 20-25

setigers, length 3-24 mm. Prostomium small

conical, pointed, with small nuchal organs at base.

First two segments faintly biannulate dorsally,

lacking annulations ventrally. Segment 3

triannulate dorsally, biannulate ventrally (Figs 13-

15). Segments 4-19 triannulate dorsally and

ventrally, the annulations more and more restricted

to the median part of dorsum and venter towards

the posterior end of body (Figs 16-18). Segments

20-22 dorsally and segments 22-24 ventrally

biannulate again; last five segments dorsally and

last three segments ventrally lacking annulations.

8 G. HARTMANN-SCHRODER & S. A. PARKER

FIGURES 13-19. Travisia oksae sp. nov. 13, anterior end, lateral view; 14, anterior end, dorsal view; 15, anterior

end, ventral view; 16, posterior end, lateral view; 17, posterior end, dorsal view; 18, posterior end, ventral view; 19,

seta.

FOUR NEW SPECIES OF OPHELHDAE 9

Posterior margin of the last nine segments with

more or less obvious crenulations dorsally (Figs

16-17). Segment 27 very small and short. Anus

terminal, encircled by seven blunt lobes (Figs 16-

18). Entire surface of body (except the branchiae)

covered with small pustules. Lateral parapodial

region of body slightly swollen from segment 13

to end of body, expanding from parapodial region

to near mid-dorsal and mid-ventral region in more

posterior segments (Figs 16-18).

Fourteen anterior—most parapodia without lobes

(Figs 13-15). Segment 15 with pair of minute

notopodial lobes, above the bundle of notosetae;

notopodial lobes increasing to ovoid lobes and still

present on small last segment. Neuropodial lobes

below neurosetae start on segment 16, similar in

shape and size to notopodial lobes, but missing on

segments 26 and 27 (Figs 16-18).

Bundles of notosetae and neurosetae from

segment 1 to segment 24; last three segments

usually asetigerous (though setae occur up to last

segment in at least one specimen, in sample SAM

E2710). Notosetae and neurosetae may be of

different length within a bundle, otherwise similar,

smooth and bilimbate (Fig. 19).

Branchiae simple, cirriform, present from

segment 2 to 23 (Figs 13-15, 18). Lateral organ a

small pit between bundles of notosetae and

neurosetae. Nephridial pores present on segments

3-14, very conspicuous on segments 7—14 (Figs

13, 15).

Comparisons with other species

Species of Travisia with a similar number of

setigers to T. oksae sp. nov. (22-29) are T.

forbesii Johnston, 1840, T. olens Ehlers, 1897, T.

antarctica Hartman, 1967, T. brevis Moore, 1906,

T. chiloensis Kiikenthal, 1887, T. kerguelenensis

McIntosh, 1885 and T. doellojuradoi Rioja, 1944.

Of these, only T. olens has been authentically

reported from Australia (see above).

The species appearing most similar to 7. oksae

is the Arctic-boreal T. forbesii (23-26 segments,

22-25 setigers). It is identical to the former in the

annulations of the segments, in the origins and

shape of the notopodial and neuropodial lobes and

the presence of nephridial pores on segment 3-14.

It differs in being on average larger (11-32, vs 3-

14 mm) and in having the setae minutely hirsute

(vs smooth and bilimbate in-T. oksae).

Also similar to T. oksae is T. olens from New

Zealand, Western Australia and South Africa,

which differs in being larger (up to 72 mm) with a

greater average number of segments and setigers

(27-32 segments, 23-29 setigers, vs 23-27 and

20-25), in having the neuropodial lobes present

(though small) from segment 1, becoming larger

from about segments 12-15 (vs present from

segment 16 only in 7. oksae), and in having the

setae minutely hispid. The reports of 7. forbesii

from South Africa and New Zealand by Ehlers

(1904, 1907, 1908) were referred to T. olens by

Augener (1922). By their descriptions, Day’s

TABLE 2. Details of stations at which Travisia oksae sp. nov. was collected.

Survey Station Lat.(S) Long (E) Depth(m) Quartzgrain size

1 2 32°35'04" 137°46'08" 6 medium

3 B2ee inl wi 137°46'00" 12 medium

4 32°40'00" 137°45'28" 13 medium

S) 32°42'20" 137°47'26" 15 medium/coarse

6 32°45'00" 137°50'00" 16 coarse

7 32°47'18" 137°49'12" 15 coarse

8 32°47'18" 137°50'00" 11 coarse

9 32°50'00" 137°49'00" 17 coarse

2 974 38°16.3' 144°44.7' 5

3 1704 38°16.12' 145°24.52' 12

2D SIMO) 145°15.45' 9

1723 KOU 145°14.86' 14

1724 38°18.56' 145°14.72' 18

72a BS lelis 145°15.93' 9

1731 SSDS £33) 145°19.28' 8

7/3335 38°23.09' AS OFS le 10

73S) 38°21.60' 145°30.59' 9)

1. Upper Spencer Gulf Benthic Survey (South Australia)

2. Port Phillip Environmental and Benthic Survey (Victoria)

3. Western Port Bay Environmental Study (Victoria)

10 G. HARTMANN-SCHRODER & S. A. PARKER

(1961, 1967) records of 7. forbesii from South

Africa are also referable to T. olens.

In T. antarctica (28 segments, 24 setigers),

notopodial and neuropodial lobes are not

mentioned, nor are there any figures by which one

might ascertain whether they are present; the setae

are described as ‘long, slender and capillary’; the

annulations of the segments are different (the first

16-17 segments are triannulate, followed by six

biannulate segments, whereas in 7. oksae

segments 4-19 are triannulate followed by three

biannulate segments); and each segment is crossed

by circlets of papillae, “resembling closely strung

beads”. T. brevis (26-27 segments, 24-25

setigers) differs by its hirsute setae and by the

nephridial pores being situated on segments 7—25.

T. chiloensis (27 segments, 24 setigers) has two

pairs of notopodial and neuropodial lobes starting

at segment 20 or 21. In T. kerguelenensis (23-27

segments, 21-23 setigers) segments 4-18 are

biannulate; there are no triannulate segments; the

posterior margins of the posterior 5—6 segments

are strongly crenulated or lobed. T. doellojuradoi

differs from 7. kerguelenensis only by its deep

purple colour when alive (black in alcohol).

Apart from T. olens and T. oksae, the only other

Travisia recorded from Australia is T. lithophila

Kinberg, 1866, known by two specimens (47, 48

mm) from New South Wales (Kinberg 1866,

Hutchings & Murray 1984). This species differs

from all the above in its greater number of setigers

(44, 53); its notopodial and neuropodial lobes

begin on setiger 12, and as in T. olens its setae are

finely hispid.

Etymology

Named after Ene-mai Oks, collector of the type

series.

Distribution and ecology

Travisia oksae is so far known from three areas

of sheltered coastal waters in southern Australia:

upper Spencer Gulf, South Australia (eight

stations from 32°35'S to 32°50'S, in medium to

coarse sand at 5—17 m), Port Phillip Bay, Victoria

(a single station, 974, just inside entrance, in sand

at 5 m) and Western Port Bay, Victoria (eight

stations, all in sand).

ACKNOWLEDGMENTS

We thank Ene-mai Oks, South Australian Department

of Environment and Planning, for depositing at the South

Australian Museum the extensive series of polychaetes

collected during the Upper Spencer Gulf Benthic Survey

of 1985-1987, Mr Timothy Stranks, Museum of

Victoria, for the loan of additonal material of Travisia

oksae sp. noy., and Dr Keith Probert, University of

Otago, for information on Allan’s Beach, type locality of

Euzonus otagoensis Probert.

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REVISION OF THE PENNATULACEAN GENUS SARCOPTILUS (COELENTERATA:

OCTOCORALLIA), WITH DESCRIPTIONS OF THREE NEW SPECIES FROM

SOUTHERN AUSTRALIA

GARY C. WILLIAMS

WILLIAMS, G. C. 1995. Revision of the pennatulacean genus Sarcoptilus (Coelenterata:

Octocorallia), with descriptions of three new species from southern Australia. Records of the

South Australian Museum 28(1): 13-32.

The pennatulacean genus Sarcoptilus Gray, 1848 from southern Australia and New Zealand

is revised. Three previously named species assignable to the genus are reassessed and two of

these are described from recently collected material. One of these three taxa is here recognized

as valid (S. grandis Gray, 1848), one is relegated to synonymy (S. roseum Broch, 1910), while

the validity of the third is questionable at present (S. bollonsi Benham, 1906). In addition, three

new species are described (Sarcoptilus nullispiculatus, S. rigidus, and S. shaneparkeri),

making a total of five species known—four considered valid taxa from southern Australia and

one of questionable validity from New Zealand. A dichotomous key to the species is included as

well as a complete list of all pennatulacean species presently known to occur in southern

Australia.

Gary C. Williams, California Academy of Sciences, Department of Invertebrate Zoology and

Geology, Golden Gate Park, San Francisco, California 94118. Manuscript received 18

November 1993.

During the 1980’s and early 1990’s, a relatively

large collection of pennatulacean coelenterates

was collected from southern Australia. From this

work as well as the literature, nineteen species of

sea pens are here recorded as occurring in

southern Australia and New Zealand (see Table

1). Five species of Sarcoptilus (family

Pteroeididae) are here treated from southern

Australia and New Zealand. Sarcoptilus grandis

Gray, 1848 is a relatively well-known member of

the southern Australian fauna. The other two

described species assignable to the genus have up

until now been poorly known (Sarcophyllum

bollonsi [Benham, 1906] and S. roseum [Broch,

1910]). The latter is here considered synonymous

with S. grandis, while the former is of

questionable validity. In addition, three new

species are also described. Southern Australia is

here defined as the continental shelf region of

southern Western Australia, South Australia,

Victoria, Tasmania, and southern New South

Wales (i.e. the southern coastal waters extending

from Perth to Newcastle). The genus seems to be

geographically restricted to between 31° and 48°

south latitude in Australia and possibly New

Zealand.

Sarcoptilus Gray, 1848 is differentiated from

related genera by having siphonozooids restricted

to a conspicuous swollen pad at the intersection of

the dorsal margin of each polyp leaf with the

dorsal edge of the rachis, the absence of

spiculiferous rays in the polyp leaves, a well

developed axis extending throughout most of the

length of the colony, and the presence of

mesozooids on the distal/ventral portion of the

rachis. Sarcophyllum Kolliker, 1869 is considered

synonymous with Sarcoptilus by Williams (in

press).

Gray (1848) and Benham (1906) described new

species of sea pens referable to the genus

Sarcoptilus from Australia and New Zealand,

respectively. Broch (1910) and Briggs (1915)

identified and described several pennatulacean

species from southern Australian coastal waters.

Utinomi and Shepherd (1982) briefly reviewed the

shallow-water sea pens of southern Australia.

Their work represents the only previously

published survey of the regional pennatulacean

fauna.

To date, material representing the following

pennatulacean taxa have been collected from the

coastal waters of Western Australia, South

Australia, Victoria, Tasmania, New South Wales,

and New Zealand, and are housed in the

collections of several institutions (see below):

Sarcoptilus, Gyrophyllum and Pteroeides

(Pteroeididae); Pennatula (Pennatulidae);

Sclerobelemnon (Kophobelemnidae); Funiculina

G. C. WILLIAMS

TABLE 1. Pennatulaceans from southern Australia and New Zealand.

Species Distribution References

& Depth (m)

Anthoptilum grandiflorum SA/VIC/NSW/TAS present work

(392-1157)

Funiculina quadrangularis

Gyrophyllum sibogae TAS (520)

Halipteris willemoesi TAS (520)

Pennatula fimbriata possibly

Australia

Pennatula phosphorea SA/TAS(436-636)

Pteroeides elegans

SA/TAS(520-597)

NSW (40-110)

Pteroeides hymenocaulum WA (11-12)

Pteroeides multiradiatum SA (40-50)

Pteroeides sp. NZ (0-5)

Ptilosarcus sinuosus possibly

(probably synonymous Australia

with Pennatula fimbriata)

Sarcoptilus bollonsi NZ (73)

Sarcoptilus grandis WA/SA/V/NSW

(10-146)

Sarcoptilus nullispiculatus SA/V/NSW,

(18)

Sarcoptilus rigidus WA (depth?)

Sarcoptilus shaneparkeri WA/SA (6-18)

Sclerobelemnon schmeltzi

Umbellula sp.

Virgularia gracillima

Virgularia gustaviana

Virgularia mirabilis

NSW (40-110)

NZ (449-4066)

NZ (18-20)

WA/SA (depth?)

SA/VIC(depth?)

present work

von Kolliker, 1872

as Leioptilus

grayi;, Batie, 1972

present work

Briggs, 1915

Broch, 1910

Utinomi & Shepherd, 1982

present work

Batie, 1972

Benham, 1906 & 1907

Gray, 1860; Briggs, 1915,

Utinomi & Shepherd, 1982;

present work

Utinomi & Shepherd, 1982;

present work

Utinomi & Shepherd, 1982;

present work

Briggs, 1915

present work

von KOlliker, 1880; Dendy, 1896;

Benham, 1907.

Utinomi & Shepherd, 1982;

present work

Utinomi & Shepherd, 1982

(Funiculinidae); Anthoptilum (Anthoptilidae);

Halipteris and Virgularia (Virgulariidae); and

Umbellula (Umbellulidae). The material, much of

it recently collected, originates mainly from Perth,

Albany, Great Australian Bight, Spencer Gulf,

Gulf St Vincent, several localities off Victoria,

New South Wales, and Tasmania, as well as the

South Island of New Zealand.

The material (including types) used in this study

is deposited at the South Australian Museum,

that of Bayer, Grasshoff, and Verseveldt (1983).

All figures in the present work are by the

author.

Key To Tue Species OF SARCOPTILUS

1 — Sclerites absent altogether

Likseearaelas Lae S. nullispiculatus sp. nov

— Sclerites present, mostly relatively densely

Adelaide (SAM); Western Australian Museum, 5 a ee Leeiti : Js nee aaa

P : . — Basal region of peduncle without robust o -

Srl OG >. New oA pala oelenae HikemsCl enite Seer eneee S. shaneparkeri sp. nov.

Institute (NZOI), Wellington; and the California

Academy of Sciences—Department of Invertebrate

Zoology and Geology, San Francisco (CASIZG).

Terminology used in the present work conforms to

— Basal region of peduncle with numerous robust

otolith-like sclerites in the interior, 0.2-1.2 mm

NC) Viethen das asap fintbca se ehadcoatancadooanbuTscrice 4

3 — Colonies stiff and rigid. Polyp leaves thick,

REVISION OF THE SEA PEN GENUS SARCOPTILUS 15

swollen, and turgid, fan-shaped to rectangular or

trapezoidal, <6 mm in length. Total colony length

<Al, OSM. §. cceeenee ren: os S. rigidus sp. nov.

— Colonies flexible to limp. Polyp leaves flattened

or thin, not turgid, semi-circular to kidney-

shaped, up to 45 mm in length. Total colony

length up to 350 mm........ S. grandis Gray, 1848

SYSTEMATIC ACCOUNT

Family PTEROEIDIDAE Kélliker, 1880

Sarcoptilus Gray, 1848

Sarcoptilus Gray, 1848: 45 (in part). Gray, 1860:

23 (in part).

Sarcophyllum K6lliker, 1869: 224. Leuckart,

1872: 280.

Kiikenthal & Broch, 1911: 441. Kiikenthal, 1915:

INT,

Pteroeides Balss, 1910: 60 (in part).

Diagnosis

Colonies feather-like and stout. Bilateral

symmetry throughout length of rachis. Axis

extends throughout entire length of colony, and

round in cross secton. Polyp leaves present and

conspicuous, rounded on margins, mostly kidney-

shaped or fan-shaped. Polyp leaves without rays.

Autozooids congested on distal margin of polyp

leaves. Anthocodiae small, retractile into their

basal protuberances. Siphonozooids restricted to

swollen pads at base of each polyp leaf where the

polyp leaves join rachis. Proximal surfaces of

polyp leaves devoid of siphonozooids. Mesozooids

usually present on distal ventral portion of rachis

in a single longitudinal row or scattered. Sclerites

smooth, not three-flanged. Spindles or rods of

polyp leaves do not form rays; long needles absent.

Rods or flattened rods present in the rachis. Small

ovoid or biscuit-shaped plates or rods may occur

in peduncle.

A genus of five species: southern Australia (four

valid species) and New Zealand (one species of

questionable validity).

TABLE 2. Table of comparative characters for the genus Sarcoptilus

Species Colour Peduncular Maximum Polyp Polyp

(in Sclerites Length Leaf Leaves

alcohol) (mm) Shape Per Side

S. bollonsi pale large 155 sickle 30

(based on reddish calcareous or fin-

the original bodies in shaped

description) the interior

S. grandis cream- ovoid 350 semi- 32-36

white finger- circular,

to pale biscuits, kidney-

yellow rods or shaped, or

or rose spindles, crescent-

& otolith- shaped

like forms

S. nulli- apricot absent 74 semi- 18-30

spiculatus to pale circular,

orange or fin-

shaped

S. rigidus cream- robust ii 7/ fan-shaped, 22-26

white spindles rectangular

& otolith- to

like forms trapezoidal

S. shane- blue- stout rods 120- triangular 22-38

parkeri grey & minute 180 or fin-

ovals shaped to

semi-circular

16 G. C. WILLIAMS

Sarcoptilus bollonsi (Benham, 1906) new comb.

Sarcophyllum bollousi Benham, 1906: 66

(misspelling—originally named for Captain

Bollons of the Government steamship ‘Hinemoa’).

Sarcophyllum bollonsi Benham, 1907: 193.

Remarks

I have not been able to locate the type specimen

for this species and I do not know of any

specimens that can be identified as Sarcoptilus

bollonsi. The species is known only from the

descriptions of Benham (1906, 1907), and

apparently has not been collected since. From

Benham’s descriptions, it is very difficult to

distinguish this species from Sarcoptilus grandis.

In my opinion, the descriptions and figures are not

detailed enough to distinguish the species. I

therefore have not included it in the key to the

species of Sarcoptilus.

At least two specimens of a species of

pteroeidid sea pen closely resembling Pteroeides

dofleini (Balss, 1909) (see d’Hondt, 1984: 18)

have recently been collected from near the type

locality of Sarcoptilus bollonsi from 0-5 metres

depth by the New Zealand Oceanographic

Institute (NZOI 0840 and NZOI Q97). These

specimens superficially resemble members of the

genus Sarcoptilus. It is possible that the two taxa

may be confused.

It is necessary to examine type material of

Sarcoptilus bollonsi in order to make a sound

assessment of its taxonomic status.

Distribution

New Zealand (southwestern coast of the South

Island). Recorded only from the type locality at

Doubtful Sound in 73 metres of water.

Sarcoptilus grandis Gray, 1848

(Figs 1-5)

Sarcoptilus grandis Gray, 1848: 45. 1860: 23.

1870: 25. Utinomi & Shepherd, 1982: 209.

Sarcophyllum australe K6lliker, 1870: 229. 1872:

186. Hickson, 1890: 140. Thomson & Mackinnon,

1911: 694.

Sarcophyllum grande Ko6lliker, 1880: 2. Balss,

1910: 60. Kiikenthal & Broch, 1911: 441. Briggs,

1915: 93. Kiikenthal, 1915: 118.

Sarcophyllum roseum Broch, 1910:

Kiikenthal, 1915: 118, 120. syn. nov.

hy.

FIGURE 1. Sarcoptilus grandis. Underwater photograph of several living sea pens. Photograph: Fred Bavendam.

REVISION OF THE SEA PEN GENUS SARCOPTILUS 17

FIGURE 2. Sarcoptilus grandis. Photographs of a single

specimen, 170 mm in length (SAM H11919). A, Dorsal

view; B, Ventral view.

Material

South Australia: SAM—10931 (H195), Great

Australian Bight, Ceduna District (32°24'S,

133°30'E), 49 metres depth, 23 August 1973, P.

Symonds (collector), 2 specimens. SAM—H11919

(H771), Great Australian Bight, near Ceduna

(BLISS, M33 DME), BO Oetolboysie WOW, IP:

Symonds (collector), 1 specimen. SAM—H11920

(H774), Spencer Gulf: 15 km S of Cowell, Stn

10X (34°0'S, 136°56’30"E), 28 September 1981,

(N.A. Carrick (collector), 1 specimen. SAM-—

H11922 (H785), Gulf St Vincent, Adelaide

District: Hallett’s Cove, 1 specimen. CASIZG-—

088073, Spencer Gulf, ca 1.5 Nm NW of Point

Riley (33°52'S, 137°35'E), 20-25 metres depth,

FV Kara George (prawn trawler), 14-15

December 1988, K. L. Gowlett-Holmes and P.

Briggs (collectors), 5 specimens. CASIZG—

088074, Gulf St Vincent, Adelaide, 4 miles SW of

end of Outer Harbour, 12 metres depth, 27 June

1965, P.D. Mitchell (collector), 2 specimens.

CASIZG-—091432, Gulf St Vincent, Port

Noarlunga Beach, 20 January 1964, Mr. Castleton

(collector), 1 specimen. Western Australia:

WAM-65-59, King George Sound, just outside

entrance to Oyster Harbour, 4.6 metres depth, 7

January 1959, 1 specimen. WAM-68-59, Albany,

Emu Point Channel, 4.6 metres depth, 15 January

1959, 1 specimen. WAM-707-91, Western

Australia, Albany, Oyster Harbour ca 6 metres

depth, 17 July 1965, E. P. Hodgkin (collector), 1

specimen. WAM-517-88, Esperance, between

Sandy Hook Island and Cape LeGrande, 31-35

metres depth, 23-25 June 1986, A. Longbottom

on LFBE ‘Triumph’ (collector), 1 specimen.

WAM-467-59, Albany, Emu Point Channel, 4.6

metres depth, 15 January 1959, 1 specimen.

WAM-706-91, Albany, Middleton Beach,

beached after storm, 5—7 August 1984, V. Milne

(collector), 8 specimens. Tasmania: SAM—H-

13071, Port Davey, Bathurst Channel, S Point of

Sarah Channel, 10—12 metres depth, 3 April 1993;

W. Zeidler, K. L. Gowlett—-Holmes, F. A.

Bavendam (collectors), 4 specimens. SAM-—

H13072, Port Davey, Bathurst Channel, 6-10

metres depth; W. Zeidler, K. L. Gowlett-Holmes,

F. A. Bavendam (collectors), 2 specimens.

Description

Specimens examined range in length from 115—

350 mm. Additional material in South Australian

Museum collection measured by Shane Parker

(late Curator of Lower Marine Invertebrates)

ranges between 165 and 325 mm in length. Rachis

comprises 50-72% of total colony length. Polyp

leaves semi-circular or kidney-shaped (usually 25—

50 mm in length and up to 30 mm in width),

relatively thin (mostly 2-4 mm thick), and number

approximately 32-36 per side. Autozooids are

small, conical in shape, and congested in several

rows (usually 10-15) that form a band along the

distal margin of each polyp leaf, extending to

approximately 5 mm down each side of a

particular leaf. Each autozooid approximately 1.0

mm in length and 0.6 mm in width.

Siphonozooids restricted to swollen pad at border

between base of each polyp leaf and rachis. Each

pad ovoid to elliptical/oblong in shape and

approximately 7 mm long by 4 mm wide. Each

siphonozooid minute (approximately 0.07 mm in

diameter). Siphonozooids highly congested and

cover surface of each pad. In addition, mesozooids

present on distal/ventral region of rachis. These

circular in shape, approximately 0.6 mm in

diameter and are either congested in several

indistinct longitudinal rows or disposed more

sparsely in a single or double longitudinal row.

Retracted mesozooids have minute slit-like

aperture, 0.12 mm in length. Sclerites of polyp

G. C. WILLIAMS

FIGURE 3. Sarcoptilus grandis. A, Upper surface of a single polyp leaf (39 mm in length), with basal siphonozooid

pad; B, A single autozooid, maximum width 0.7 mm; C, View from above of a single mesozooid, 0.6 mm in

diameter; D, Polyp leaf sclerites, 1.15 mm. 0.48 mm, 1.25 mm; E, Sclerites from the surface of the peduncle, 0.19

mm, 0.17 mm, 0.07 mm, 0.12 mm; F, Sclerites from the interior of the upper and middle portions of the peduncle,

0.28 mm, 0.23 mm , 0.21 mm, 0.17 mm, 0.55 mm; G, Sclerites of the interior of the lower portion of the peduncle,

1.1 mm, 0.87 mm; H, Map showing distribution of the species: * = type locality, @ = other collecting stations; arrow

shows type locality of Sarcophyllum roseum.

36,

B, 0.91 mm

1mm

I, 0.85 mm.

G, 0.66 mm; H, 1.10 mm

1S)

PL,

joa]

ica]

ie)

ea}

E, 0.82 mm; F, 1.23 mm

1.05 mm; D, 1.02 m

FIGURE 4. Sarcoptilus grandis. Scanning electron micrographs of polyp leaf sclerites. A, 1

20 G. C. WILLIAMS

FIGURE 5. Sarcoptilus grandis. Scanning electron micrographs of peduncular sclerites. A-C, Sclerites from the

interior of the upper and middle portion of the peduncle. A, 0.57 mm; B, 0.76mm; C, 0.53 mm. D-H, Sclerites from

the interior of the basal portion of the peduncle; D, 0.85 mm; E, 0.74 mm; F, 1.05 mm; G, 0.77 mm; H, 0.58 mm.

REVISION OF THE SEA PEN GENUS SARCOPTILUS All

leaves smooth slender rods or spindles (0.38-1.35

mm in length). Many of these slightly clavate,

being enlarged at one end and tapering to a

pointed opposite end. Polyp leaf sclerites

distributed relatively densely in portion containing

autozooids, and very sparse or absent on proximal

faces of polyp leaves. Sclerites of surface of polyp

leaves often form short lines between groups of

autozooids in radiating fashion. Autozooids

contain sclerites that either run parallel to

longitudinal plain of each autozooid or converge

to form inverted ‘V’ bordering each autozooid.

Sclerites of peduncle are of three distinct types—

small ovoid fingerbiscuits (0.06-0.19 mm in

length) common throughout the surface, stout rods

and spindles (0.17—0.76 mm in length) in interior

of upper and middle portions of peduncle, and

large robust otolith-like forms (0.20-1.1 mm in

length) in interior of lower portion near base.

Colour in life off-white, orange, or rose; cream or

tannish-white preserved in alcohol.

Distribution

Southern Australia (Vicinity of Albany,

Western Australia to at least as far as the Manning

River/Cape Hawke region, New South Wales);

4.6-146 metres in depth. Utinomi and Shepherd

(1982: 211) state the range as being from the

Great Australian Bight to southern Queensland.

This is by far the most common and widespread

species in the genus. Collecting stations for the

species come from Briggs (1915), Gray (1860,

1870), Kolliker (1870, 1872, 1880), Hickson

(1890), Thomson and MacKinnon (1911), as well

as from a large number of recently acquired lots in

the collections of the Western Australian Museum,

the South Australian Museum, and the California

Academy of Sciences. The maximum depth record

is reported by Briggs (1915: 94) from King Island,

Bass Strait. The type locality was not recorded in

the original description by Gray (1848: 45) but

only later by Gray (1860: 23 and 1870: 25) as

Sydney, Australia.

Discusssion

I have unfortunately not been successful in

locating the type specimen of Sarcophyllum

roseum Broch, 1910 for comparison. However,

after having examined a large number of

specimens of Sarcoptilus from a wide geographic

scope, I here conclude that Sarcophyllum roseum

should be considered as a junior synonym of

Sarcoptilus grandis. In my opinion, the minor

morphological variance observed in Sarcophyllum

roseum, which was used by Broch to distinguish

his species from Gray’s, can certainly be

accommodated by the range of variation in

Sarcoptilus grandis. Included in the specimens

examined for the present study are several from

Broch’s type locality (Albany, Western Australia),

which agree well in virtually all respects with his

original description.

The four characters used by Broch (1910: 121)

to distinguish the species are as follows: (1) rachis

polyps—forming a long row or plate in S. grandis

vs a simple or double row in S. roseum; (2) the

autozooid region of the polyp leaves—in which

this polyp zone is wide with numerous sclerites on

both sides of the leaf in S. grandis vs narrow with

sclerites only on the under surface of the leaf in S.

roseum, (3) the sterile surface of the polyp leaf—

with numerous sclerites in S. grandis vs no

sclerites in S. roseum; and (4) length of sclerites

from the surface of the rachis and peduncle—

rachis sclerites <0.7 mm long and peduncle

sclerites <0.4 mm long in S. grandis vs <0.5 mm

and <0.2 mm respectively in S. roseum. These are

all variable characters that show gradations

between specimens and hence cannot be used to

justify the differentiation of two species. An

example that contradicts Broch’s distinction is a

specimen from King George Sound (WAM — 65—

59), which has the rachis mesozooids disposed in

a plate as in S. grandis, but is without sclerites in

the sterile portion of the polyp leaf as in S.

roseum. 1 conclude that Sarcophyllum roseum and

Sarcoptilus grandis are morphologically

indistinguishable and therefore conspecific.

Apparently a substantial amount of variation is

present in the size of the peduncular sclerites.

Kiikenthal (1915: 119) recorded sclerites from the

interior as large as 4.5 mm in length, and those

from the surface up to 0.4 mm long.

Sarcoptilus nullispiculatus sp. nov.

(Figs 6, 7A,B)

?Sarcoptilus sp. Utinomi & Shepherd, 1982: 211.

Material

Holotype. SAM—H10929A, South Australia:

Gulf St Vincent, Adelaide District, Port Stanvac,

18 metres depth, D. Cooper (collector), 1 whole

specimen.

Paratype. SAM-H10929B, same data as

holotype, 1 whole specimen.

Other material studied. SAM—H10929C, same

data as holotype, 1 whole specimen.

oy G. C. WILLIAMS

FIGURE 6. Sarcoptilus nullispiculatus. A, Lower surface of a single polyp leaf (6 mm in length), with siphonozooid

pad; B, View from above of a single mesozooid, 0.45 mm in length; C, Distal portion of a single autozooid, 0.6 mm

in width; D, Map showing distribution of the species; * = type locality, @ = localities reported by Utinomi and

Shepherd (1982) for Sarcoptilus sp.

Diagnosis

Colonies under 75 mm in length, slender and

limp. Rachis and peduncle roughly equal in

length. Polyp leaves 18-30 per side, fin-shaped to

semi-circular, <7 mm in length. Autozooids in 1-

2 rows, 25-36 per leaf. Siphonozooids restricted

to circular pads at dorsal base of each leaf. Single

longitudinal row of up to 20 mesozooids present

on ventral/distal surface of rachis. Sclerites absent

altogether. Colour orange or pale-salmon in

alcohol.

Description

The three specimens examined range in length

from 58-74 mm. Holotype is 68 mm long while

paratype is 74 mm in length. Colonies are slender

and flaccid. Rachis comprises 47-58% of total

colony length. Dorsal side of rachis has a medial

longitudinal groove for its entire length. Polyp

leaves number 18-30 per side, fin-shaped to semi-

circular, each leaf up to 7 mm in length.

Autozooids disposed in one or two rows along

distal margin of each leaf, and number

approximately 25-36 autozooids per leaf. Each

autozooid 0.4-0.6 mm in diameter. Most

autozooids have a single non-spiculated, flexible,

nipple-like protuberance projecting distally from

their apices. Siphonozooids contained on circular

to ovoid pads, 1.5 mm in diameter, on the dorsal

base of each polyp leaf adjacent to lateral margin

of rachis. Each siphonozooid approximately 0.07

mm in diameter. In addition, a single medial row

of 10-20 mesozooids present on distal/ventral

portion of rachis, each mesozooid 0.20-0.45 mm

REVISION OF THE SEA PEN GENUS SARCOPTILUS 28

FIGURE 7. Photographs of holotypes. A,B. Sarcoptilus nullispiculatus. A, Ventral view; B, Dorsal view. C,D.

Sarcoptilus rigidus, C, Ventral view; D, Dorsal view.

in diameter. Sclerites apparently absent altogether

from all parts of colonies. Colour throughout

apricot-orange to pale orange in alcohol.

Etymology

The specific epithet is derived from the Latin,

nullus = not any or none, and spiculum = a point

or dart; in reference to the lack of sclerites in this

species.

Distribution

Southern Australia (Gulf St Vincent, South

24 G. C. WILLIAMS

Australia, and possibly Pt Fairy, Victoria to Jervis

Bay, New South Wales—as reported by Utinomi

& Shepherd, 1982); 18 metres in depth. Type

locality is Gulf St Vincent, South Australia.

Discussion

This species is differentiated from other

members of the genus by a complete lack of

sclerites. It is likely that Sarcoptilus sp. of

Utinomi and Shepherd (1982) is conspecific with

this species. Even though they do not mention the

lack of sclerites in this species, their description

agrees in all other aspects with the present

material.

Sarcoptilus rigidus sp. nov.

(Figs 7C,D, 8, 9)

Material

Holotype. WAM-—363-31, Western Australia:

vicinity of Perth: Cottesloe Beach, 1 whole

specimen.

Paratype. WAM-364-31, same data as

holotype.

Other material studied. WAM-714-91,

Western Australia: Albany: Middleton Beach,

beach drift, 10 August 1991, S. Bolton (collector),

1 specimen.

Diagnosis

Colonies stiff and rigid, dart-shaped. Polyp

leaves thick and turgid, fan-shaped to rectangular

or trapezoidal; <6 mm in length; 22-26 leaves per

side of rachis. Autozooids in single row of 25—30

per polyp leaf. Several mesozooids scattered on

ventral distal extremity of rachis. Siphonozooid

pad conspicuous and kidney-shaped. Peduncular

sclerites: robust spindles in upper part (0.25—0.42

mm long) and robust, ovoid, otolith-like bodies

(0.3-1.2 mm long) in lower part. Colour in alcohol

uniformly cream-white or bicoloured with rachis

grey and peduncle cream-white.

Description

Specimens examined dart-shaped, markedly

stiff and rigid, 65-117 mm in length. Holotype is

80 mm long and paratype 65 mm in length.

Peduncle makes up approximately 38-43% of

total length. Axis extends throughout entire length

of each specimen. Polyp leaves thick and turgid,

fan-shaped to rectangular or trapezoidal with

rounded corners, 3-6 mm in length. 22-26 polyp

leaves per side. Autozooids completely retractile

and disposed in single row at distal margin of

each polyp leaf, generally 25-30 per polyp leaf.

Each retracted autozooid has slit-like aperture,

perpendicular to plane of polyp leaf. This aperture

surrounded by slightly ovoid ring-like basal

swelling without teeth. Siphonozooids contained

on swollen, more-or-less elliptical to kidney-

shaped pads at dorsal base of each polyp leaf,

adjacent to lateral margin of rachis. Each

siphonozooid pad approximately 1.5—3.0 mm in

length, while an individual siphonozooid is

approximately 0.1 mm in diameter. In addition,

several mesozooids scattered on distal ventral

extremity of rachis, each approximately 0.3—-0.5

mm in diameter. Upper and lower surfaces of

polyp leaves densely or sparsely ornamented with

sclerites that are mostly longitudinally disposed in

parallel. Sclerites of polyp leaves slender elongate

rods and spindles, 0.3-1.3 mm in length. Many of

these faintly longitudinally grooved, some rounded

on ends while others angled and obliquely truncate

at ends. Sclerites of rachis similar to those of polyp

leaves, sparsely distributed. Sclerites of upper

portion of peduncle smooth robust spindles, 0.25—

0.42 mm in length. These tapered to rounded on

ends and often slightly constricted in middle.

Lower part of peduncle contains ovoid otolith-like

bodies, 0.3-1.2 mm in length. Most of these

robust with rounded ends, and many somewhat

constricted in middle. Colonies vary in colour in

alcohol from bicoloured—rachis grey with

peduncle cream-white, to monochromatic—

entirely cream-white.

Etymology

The specific epithet is derived from the Latin

rigidus = stiff or rigid; in reference to the rigid

and turgid nature of the colonies of this species.

Distribution

Western Australia (Perth and Albany); shallow

sublittoral (depth range unknown). This species is

known from only three specimens collected at two

localities. The type locality is Perth, Western

Australia.

Discussion

This species differs markedly from other

members of the genus by its remarkably rigid form

and conspicuously turgid polyp leaves. The only

other species containing similar otolith-like

sclerites in the peduncle is Sarcoptilus grandis.

The density of spiculation on the polyp leaves

varies greatly from very dense (and covering the

entirety of both surfaces) in the holotype and

paratype, to very sparse in the specimen from

REVISION OF THE SEA PEN GENUS SARCOPTILUS aS

FIGURE 8. Sarcoptilus rigidus. A, Ventral view of a single polyp leaf (4 mm in length), with siphonozooid pad; B,

Polyp leaf sclerites, 0.95 mm, 1.30 mm, and 0.40 mm in length; C, Peduncular sclerites, 0.27 mm, 0.42 mm, 0.35

mm, 1.2 mm, 1.0 mm, and 0.5 mm in length; D, View from above of a single autozooid, 0.27 mm in length; E, Map

showing distribution of the species: * = type locality; @ = other locality.

Albany, in which only a few sclerites are scattered Scytalium sp. Utinomi & Shepherd 1982: 209.

in the distal margin of each leaf just below the

autozooids. Material

Holotype. SAM-H10923A, South Australia:

; : upper Spencer Gulf: 1.6 km E of Douglas Point,

Sarcoptilus shaneparkeri sp. nov. 17 metres depth, May 1988, N. Holmes

(Figs 10-13) (collector), 1 whole specimen.

26 G. C. WILLIAMS

FIGURE 9. Sarcoptilus rigidus. Scanning electron micrographs. A—C, Sclerites of the polyp leaves. A, 0.99 mm; B,

0.33 mm; C, 0.41 mm. D-G; Sclerites of the peduncle. D, 0.77 mm; E, 0.54 mm; F, 0.73 mm; G, 0.86 mm.

Paratype. SAM-H10923B, same data as metres depth, May 1988, N. Holmes (collector),

holotype, 1 whole specimen. 11 specimens. SAM-H10924, upper Spencer

Other material studied. South Australia: Gulf: 1.6 km E of Douglas Point (Kinhill Survey

SAM-H10922, upper Spencer Gulf, 1.6 km E of Station 3A), 17 metres depth, May 1988, N.

Douglas Point (Kinhill Survey Station 3A), 17 Holmes (collector), | specimen. SAM—H10926,

REVISION OF THE SEA PEN GENUS SARCOPTILUS 27

FIGURE 10. Sarcoptilus shaneparkeri. Photographs of

holotype. A, Ventral view; B, Dorsal view.

upper Spencer Gulf: 1600 m E of Douglas Point;

27 June 1988, N. Holmes (collector), 3 specimens.

SAM-H11842, Spencer Gulf, 22 km S of Port

Augusta, 10 metres depth, 3 February 1983, R.

Henderson (collector), 1 specimen. SAM-—

H10918, Spencer Gulf between Douglas Point and

Mambray Creek, 15-18 metres in depth, June

1987, N. Holmes (collector), 1 specimen. SAM—

H10920, Spencer Gulf between Douglas Point and

Mambray Creek, 15-18 metres depth, June 1987,

N. Holmes, 2 specimens. SAM—H10921, upper

Spencer Gulf, 1.6 km E of Douglas Point, 15

metres depth, 5 May 1988, N. Holmes, 1

specimen. SAM—H10917, upper Spencer Gulf, S

of Redcliff Point, 15 metres depth, 9-11 April

1980, 2 specimens. SAM—H10916, upper Spencer

Gulf, Port Germein Bay, 6 metres depth, February

1980, N. Holmes (collector), 1 specimen. SAM-—

H10919, upper Spencer Gulf near Redcliff, 15

metres depth, 17 November 1980, S.A. Shepherd

(collector), 1 specimen. SAM-—H 10927, Gulf St

Vincent, Adelaide District, Seacliff, 15 metres

depth, 12 May 1990, N. Holmes (collector), 4

specimens. SAM-—H10925, upper Spencer Gulf,

1.6 km E of Douglas Point (Kinhill Survey Station

3A), 17 metres depth, May 1988, N. Holmers

(collector), 1 specimen. SAM—H10928, Kangaroo

Island, Penneshaw, 14 metres depth, 3 August

1971, J. Kroezen (collector), 2 specimens.

Western Australia: WAM-—709-91, North Mole,

Fremantle Harbour, 25 November 1983, S. Slack

Smith/L. Marsh/J. Watson/C. Bryce (collectors), 5

specimens. WAM-—707-91B, Albany, Oyster

Harbour, ca 6 metres depth, 17 July 1965, EP.

Hodgkin (collector), 1 specimen.

Diagnosis

Maximum length 180 mm. Polyp leaves thin,

triangular or semi-circular to kidney-shaped, 22—

38 pairs of polyp leaves. Polyp zone of each leaf

with autozooids in 1 or 2 rows, at least autozooids

20 per row. Retracted autozooids with single basal

tooth composed of converging sclerites. Several

mesozooids scattered or in a single longitudinal

row on ventral distal extremity of rachis.

Peduncular sclerites mostly elongate biscuit-

shaped rods, 0.11-0.17 mm long. Colour mostly

bicoloured: rachis dark blue-grey or brownish-grey

with peduncle cream to orange. Monochromatic

orange form less common.

Description

Specimens examined range in length from 60—

180 mm. Holotype 108 mm in length while

paratype 100 mm long. Length of peduncle

constitutes 45-55% of total length of single

specimen. Rachis may contain deep longitudinal

medial groove along entire length of its dorsal

side. 22-38 pairs of polyp leaves, which are thin

and variable in shape: from deltoid/fin-shaped to

semicircular or kidney-shaped, mostly 8-13 mm

in length. Retractile autozooids arranged usually

in one or two rows along distal margin of each

polyp leaf, with 20-35 autozooids per row. Distal

tip of each autozooid has single tooth, which is

small, blunt to pointed, and triangular in shape.

This polyp tooth projects over top of each retracted

autozooid. Siphonozooids densely-crowded on

small roughly circular pad, situated on dorsal base

of each polyp leaf, adjacent to rachis. Each pad ca

1.7 mm in diameter, and single siphonozooid ca

0.1-0.2 mm in diameter. In addition, 10-28

mesozooids present on distal ventral portion of

rachis, usually in single longitudinal row or

sometimes randomly scattered. These vary from

0.23-0.50 mm in diameter. All three forms of

polyps contain scattered sclerites in polyp walls.

Sclerites of polyp leaves and surface of rachis are

mostly slender rods and spindles, 0.22—0.93 mm

in length. Those of polyp leaves sparsely to

densely-set in longitudinal rows between the

G. C. WILLIAMS

SENTERO TAN NAME EL

RTT a TET

FIGURE 11. Sarcoptilus shaneparkeri. A, Ventral view of a single polyp leaf (13 mm in length), with siphonozooid

pad; B, Sclerites of the polyp leaves and rachis, 0.33 mm, 0.48 mm, 0.58 mm, and 0.64 mm in length; C, Sclerites of

the peduncle, elongate forms are 0.19 mm, 0.18 mm, 0.17 mm, 0.15 mm, and 0.13 mm in length, minute ovals are

0.01—0.02 mm in length; D, Lateral view of a single mesozooid, 0.43 mm in length; E, Distal portion of a single

autozooid, 0.45 mm in width; F, Map showing distribution of the species: * = type locality, @ = other localities.

autozooids, as well as being sparsely-scattered

obliquely over face of each autozooid just below

peristome. Longitudinal rows extend over most of

surfaces of both sides of each polyp leaf and

radiate outward toward distal margin from base of

each leaf. Two adjacent longitudinal rows

converge at distal apex of each autozooid to form

single inverted V-shaped polyp tooth. Some of

sclerites forming tooth blunt and somewhat

flattened at distal end. Sclerites from surface of

rachis and peduncle arranged mostly

longitudinally and in parallel, with a few obliquely

REVISION OF THE SEA PEN GENUS SARCOPTILUS 29

AWB C am ~D E

/ |

_ yo -

ye a y

| i

FIGURE 12. Sarcoptilus shaneparkeri. Scanning electron micrographs of polyp leaf sclerites. A, 0.62 mm; B, 0.60

mm; C, 0.59 mm; D, 0.39 mm; E, 0.93 mm; F, 0.27 mm; G, 0.30 mm; H, 0.44 mm; I, 0.23 mm.

disposed. Sclerites of surface of peduncle of two

distinct kinds—stout smooth rods, sometimes

slightly constricted in middle, with rounded ends

(0.11-0.17 mm in length and 0.02-0.03 mm in

width), and numerous minute ovals (0.01-0.02

mm on length). Sclerites of interior of rachis and

peduncle very sparse or apparently lacking. Colour

in alcohol of rachis and polyp leaves varies from

dark bluish-grey to brownish-grey or violet-grey

(rarely orange), contrasting with peduncle which

is yellowish-white or pale-orange. Sclerites are

colourless.

Etymology

This species is named for Shane A. Parker, late

Curator of Lower Invertebrates at the South

Australian Museum. Shane originally suggested

that the southern Australian pennatulacean fauna

be studied. In addition, he supplied on loan much

of the essential material for study. Without his

interest, enthusiasm, and good humour, this

project would not have been possible.

Distribution

Southern Australia (Fremantle Harbour and

Albany, Western Australia; Spencer Gulf, Gulf St

Vincent, and Kangaroo Island, South Australia);

6-18 metres in depth. This species is apparently

locally common in shallow-water of protected

embayments, on silty bottom or in hollows of

coarse sand between seagrass beds. The type

locality is the upper Spencer Gulf, South Australia.

G. C. WILLIAMS

B, 0.15

Bae!

io)

S *

a. &

Susy

= iS

5 §

nas

Sas

San

ary

= 2

S &

ea =)

i) SG

REVISION OF THE SEA PEN GENUS SARCOPTILUS 31

Discussion

Utinomi and Shepherd (1982: 208-209)

incorrectly identified this species as an

undetermined species of Scytalium. Members of

the genus Scytalium have sclerites that are

exclusively oval-shaped plates and the

siphonozooids are arranged only on the rachis

between the polyp leaves. In Sarcoptilus, the

sclerites are spindles or rods and the

siphonozooids are mostly situated on circular pads

at the dorsal base of each polyp leaf.

There is a considerable amount of variability

observed in this species. Members of the South

Australian populations do not exceed 120 mm in

length, while those from Western Australia are

markedly larger—up to 180 mm in length. The

arrangement and amount of spiculation on the

polyp leaves varies considerably—from very

sparse to very dense. In some specimens, the

proximal portion of each polyp leaf is devoid of

sclerites. The amount of development of the

terminal polyp tooth is also variable—being

conspicuous in most members of the South

Australian populations, and less strongly

developed in others, particularly the specimens

from Western Australia. The amount of

spiculation present in the surface of the rachis and

peduncle is variable—relatively sparse in some to

dense in others.

Observations

This species is preyed upon by arminacean

nudibranch molluscs (presumably Armina sp.) (S.

A. Parker, pers. comm.). Many of the specimens

observed have the ventral margins of the polyp

leaves devoid of polyps—showing signs of being

partially eaten.

The following notes on colour, written by S.A.

Parker, were found with five specimen lots:

(SAM-H10922)—‘Colours shortly after

collection: stalks bright orange to pale orange,

rachis brownish olive to light orange-brown;

leaves dark olive-grey-brown; autozooids pale

orange to brownish-cream, siphonozooids pale

orange’; (SAM-H10923A Holotype and 10923B

Paratype)—‘Colours shortly after collection: stalk

pale apricot-buff, rachis & leaves dull leaden grey

with slight brownish tinge; autozooids very pale

greyish white to off-white. Siphonozooid pads pale

greyish white’; (SAM—H10927)—‘Lobes violet-

grey, stem creamy-orange’; (SAM—H10928)—

‘Colour in life: orange’; (SAM—H10925)—

‘Colours shortly after collection: stalk bright

orange, rachis leaves & siphonozooids paler;

autozooids whitish orange’. The bicoloured grey/

cream form is apparently the most common form

and is present throughout the range of the species.

The less common monochromatic orange form has

only been encountered in upper Spencer Gulf and

Kangaroo Island (South Australia) and Albany

(Western Australia).

CONCLUSIONS

Three species were previously assignable to the

pennatulacean genus Sarcoptilus: S. grandis as

the type species, with Sarcophyllum bollonsi and

Sarcophyllum roseum transferable to the genus.

This revision adds three new species (S.

nullispiculatus, S. rigidus, and S. shaneparkeri),

one synonymy (Sarcophyllum roseum as a junior

synonym of Sarcoptilus grandis), and one new

combination (Sarcoptilus bollonsi), establishing a

total of five species for the genus (four valid

species in southern Australia and one species of

questionable validity in New Zealand (S.

bollonsi)).

ACKNOWLEDGMENTS

I am grateful to Shane Parker, late Curator of Lower

Marine Invertebrates at the South Australian Museum in

Adelaide, for originally suggesting the study of the

southern Australian pennatulacean fauna and for kindly

lending the material used in this study.

I thank Loisette Marsh, Curator of Aquatic

Invertebrates, Western Australian Museum in Perth, and

Dennis Gordon and Paul Anderson of the New Zealand

Oceanographic Institute in Wellington for the loan of

material from their collections. I also thank Karen

Gowlett-Holmes, Wolfgang Zeidler, and Eric Matthews

(South Australian Museum) for their cooperation. I am

grateful to Darrell Ubick (CAS Scanning Electron

Microscope Unit), and Charlotte Fiorito and Caroline

Kopp (CAS Photography Department) for their

assistance in the production of the plates.

I am indebted to Fred Bavendam, professional

freelance photographer, for use of his underwater

photograph in Figure 1.

I sincerely thank Marie-José d’Hondt (National

Museum of Natural History, Paris) for her helpful ideas

concerning the manuscript.

3, G. C. WILLIAMS

REFERENCES

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BAYER, F. M., GRASSHOFF, M., & VERSEVELDT,

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BENHAM, W. B. 1907. On a new species of pennatulid

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BRIGGS, E. A. 1915. Report on the alcyonarians

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F.1.S. ‘Endeavour’ 1909-1914 3 (2): 59-94.

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genus of Pennatulidae. Proceedings of the Zoological

Society of London 1848: 45.

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with descriptions of some new species in the British

Museum. Annals and Magazine of Natural History

(3) 5: 20-25.

GRAY, J. E. 1870. ‘Catalogue of the sea-pens or

Pennatulariidae in the collection of the British

Museum’. British Museum: London.

HERKLOTS, J. A. 1858. Notices pour servir a |’ étude

des polypiers nageurs ou pennatulides. Bijdragen tot

de Dierkunde 7: \-31.

HICKSON, S. J. 1890. Preliminary report on a collection

of Alcyonaria from Port Phillip. Proceedings of the

Royal Society of Victoria, Melbourne, (new series) 2

(2): 136-140.

HICKSON, S. J. 1916. The Pennatulacea of the Siboga

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HONDT, M.-J. d’. 1984. Pteroeides (Octocorallia,

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Muséum national d’histoire naturelle (4) 6 (A, 1):

3-29.

KOLLIKER, R. A., von. 1869-72. Anatomisch-

Systematische Beschreibung der Alcyonarien. I. Die

Pennatuliden. Abhandlungen von der

Senckenbergischen naturforschenden Gesellschaft 7:

111-255 (1869); 487-602 (1870); 8: 85-275 (1872).

KOLLIKER, R. A., von. 1880. Report on the

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years 1873-1876. Report on the Scientific Results of

the Voyage of H.M.S. Challenger during the years

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von R. Friedlander: Berlin.

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Wissenschaftliche Ergebnisse der Deutschen Tiefsee-

Expedition auf dem Dampfer ‘Valdivia’ 1898-1899

13 (1) Lieferung 2: 113-576.

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recueillis par la Commission de dragages de I’ aviso le

‘Travailleur’ durant les campagnes 1880-1881.

Oeuvres posthumes de A.-F. Marion. Réunies par Paul

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expédition du Travailleur. Expédition scientifique

Travailleur et Talisman: 103-151. Masson: Paris.

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pennatulides recueillis par le Travailleur et le

Talisman, dans |’Océan Atlantique, au large du

Maroc. Bulletin du Muséum d@’histoire naturelle 11:

454-458.

STUDER, T. 1891. Note préliminaire sur les alcyonaires

provenant des campagnes du yacht ‘l’Hirondelle’

1886, 1887, 1888. Part 2. Alcyonacea and

Pennatulacea. Mémoires de la Société zoologique de

France 4: 86-95.

THOMSON, J. A. & HENDERSON, W. D. 1906. ‘An

account of the alcyonarians collected by the Royal

Indian Marine Survey Ship Investigator in the Indian

Ocean. Part I. The alcyonarians of the deep sea’.

Indian Museum: Calcutta.

THOMSON, J. A. & MACKINNON, D. L. 1911. The

alcyonarians of the ‘Thetis’ Expedition. Australian

Museum Memoirs 4: 661-695.

TIXIER-DURIVAULT, A. & d’HONDT, M.-J.

1973(1974). Nouvelles récoltes d’Octocoralliaires a

Madagascar. Téthys 5 (2-3): 251-266.

UTINOMI, H. & SHEPHERD, S. A. 1982. Seapens

(Order Pennatulacea). Jn Shepherd, S. A. & Thomas,

I. M. (eds), Marine invertebrates of southern

Australia, Part I. Handbooks Committee, South

Australian Government.

WILLIAMS, G. C. In Press. Living genera of sea pens —

illustrated key and synopses (Coelenterata:

Octocorallia; Pennatulacea). Zoological Journal of the

Linnean Society.

FOLLOWING THE TRACKS OF EDGAR WAITE IN NEW GUINEA

FOR THE PACIFIC ARTS SYMPOSIUM IN ADELAIDE

BARRY CRAIG

CRAIG, B. 1995. Following the tracks of Edgar Waite in New Guinea for the Pacific Arts

Symposium in Adelaide. Records of the South Australian Museum 28(1): 33-52.

The Fifth International Symposium of the Pacific Arts Association, held in Adelaide during

1993, provided an opportunity for the South Australian Museum to reforge links with New

Guinea and the islands of the Bismarck Archipelago, established through the visit there by

Edgar Waite as Museum Director 75 years earlier. This paper describes, in narrative style, the

author’s experiences during a fieldtrip made to collect relevant artefacts and data and to organise

dance performances for the Adelaide Symposium.

B. Craig, South Australian Museum, North Terrace, Adelaide, South Australia, 5000.

Manuscript received 20 April, 1994.

In 1918, Edgar Waite, the Director of the South

Australian Museum, undertook a collecting

expedition to New Guinea and the islands of the

Bismarck Archipelago (Hale 1956: 115-16).

Waite was accompanied by Augustus C. Davis

who previously had been a Protector of Natives in

New Ireland and was familiar with the area to be

covered by the expedition. The former German

territory had come under Australian military

control following on the declaration of war in

1914, and by 1918 it was only just reopening to

other interests. The Bismark Archipelago was not

well represented in the Museum’s collections and

until this opportunity arose, natural history and

ethnographic collections would have had to have

been purchased through traders such as Richard

and Phoebe Parkinson and, even earlier, through

‘Queen’ Emma and Thomas Farrell, as did the

Australian Museum in Sydney (Thomsett 1993:

3):

Waite was first and foremost a naturalist, not an

ethnographer (Jones 1992). As was common in

the late nineteenth and early twentieth centuries

though, museum collectors were generalists when

in the field and Waite collected a wide range of

natural science specimens and ethnographic

material. Although Waite did not stay in any one

place long enough to begin to obtain systematic

cultural information, he did make many

serendipitous observations. In a sense, everything

was new and unfamiliar, so everything was worth

a comment. His ethnographic data has to be

treated with caution nevertheless.

Today the visitor to the Pacific Gallery of the

South Australian Museum can see a representative

sample of the ethnographic material collected by

Waite and Davis, presented in a manner that must

differ little from the way it was first displayed in

1919 (see ‘New Museum Exhibits’, The Register,

September 13, 1919). The manner of display (Fig.

1) is only a slight improvement on the renaissance

‘Cabinets of Curiosities’. The major points of

interest are the carved figures and masks made for

the series of funerary rituals called malangan in

New Ireland, and a rare mask from the Sulka

people of Wide Bay, East New Britain (Fig. 15).

The South Australian Museum has never had a

specialist Curator for its Foreign Ethnology

collections. The Pacific collections (including

Melanesian material) comprise the largest

component of these. Adelaide has arguably the

second most significant collection of Pacific

material in the country. In 1992, the Anthropology

Division of the South Australian Museum offered

to host the Fifth International Symposium of the

Pacific Arts Association. It seemed to this author

that the Symposium could present an opportunity

for demonstrating what could be done by the

Museum to upgrade its Pacific Gallery and the

collections, and data about those collections, on

which the Gallery is based.

It was decided that the projects should be joint

ventures with the Papua New Guinea National

Museum in the case of the Sulka component and

with the New Ireland Provincial Government in

the case of the New Ireland component. Chris

Issac, Acting Director of the J. K. McCarthy

Museum in Goroka, a branch of the PNG National

Museum, who had been one of my staff when I

was Curator at the PNG National Museum, would

act as my colleague for the Sulka Project. He

himself is Sulka, from the village of Guma on the

34 B. CRAIG

FIGURE 1. New Ireland malangan collected by Edgar Waite in 1918, on display in the Pacific Gallery of the South

Australian Museum (Photo by Trevor Peters, SAM).

south-eastern shore of Wide Bay, East New

Britain. In 1982, he was part of the team from the

PNG National Museum which collected a

magnificent hemlaut mask, several susu and other

masks, and dance wands, clubs and shield from

the Sulka. This team consisted of myself, Chris

Issac and Rowena Hill' who was the Conservator

at the PNG National Museum at that time. Chris

' Rowena Hill documented the materials used in the construction of

the masks and dance wands and later carried out a detailed study of

materials and techniques relevant to the masked rituals of the Sulka

and Sulkanised Mengen of Wide Bay. This material has been

presented as a Masters thesis to the Department of Sociology and

Anthropology at the University of Queensland. She has also recorded,

and had translated, a corpus of songs relating to the dancing of the

masks (see Hill 1982; Craig 1993a).

Issac also worked with George Corbin of the City

University of New York in his ‘Salvage Art

History’ project among the Sulka in 1983 (Corbin

1990).

At the South Pacific Festival of Arts held in

Cook Islands in 1992, Susan Cochrane’ had met

Noah Lurang, New Ireland Provincial Cultural

Officer. She suggested he would be an ideal

person to involve in the New Ireland component

of the project. He was an experienced high school

teacher, was articulate about the malangan culture

of New Ireland and able to perform masked

dances associated with the malangan rites.

2. Co-ordinator of the Pacific Arts Symposium in Adelaide.

FOLLOWING THE TRACKS OF EDGAR WAITE

FIGURE 2. Map showing Sulka area, Wide Bay, East New Britain Province, Papua New Guinea.

35)

36 B. CRAIG

® Maragon

SIMBERI

Maragat

Anoos Rock

Tomalabat®

—_

TABAR ISLANDS-showing locations from which

Edgar Waite collected cultural materials

FIGURE 3. Map showing Tabar Islands, New Ireland Province, Papua New Guinea.

FOLLOWING THE TRACKS OF EDGAR WAITE 37

At first, it was hoped that funding would be

received early enough to commission a copy of the

South Australian Museum’s rare Sulka hemlaut

mask, obtained in 1919 and designed to be danced

by two men simultaneously. Funding was not

confirmed until November 1992, which did not

provide sufficient time for a copy to be

constructed. However, the performance of mask

dancing associated with the initiation of boys and

girls is common over the New Year period among

the Sulka, allowing an opportunity to record

performances and to purchase some of the masks

before they were destroyed. It is normal practice to

destroy the masks shortly after their performance.

In the case of New Ireland, we proposed to

obtain at least two tantanua masks,’ used in

malangan performances, to place opposite the

tantanua masks collected by Waite in 1918.

Malangan performances are rare, but as New

Irelanders no longer destroy the tantanua masks

afterwards, there would be a good chance of being

able to buy some.

The fieldwork was not intended merely as a

collecting exercise. It was intended to identify two

New Irelanders and two Sulka men who would be

prepared to travel to Adelaide for the Pacific Arts

Symposium and perform the masked dances. The

masks would then be set up opposite their much

older counterparts in the Pacific Gallery to

demonstrate the continuity of these masking

traditions and the re-establishment of the

relationship between the South Australian

Museum and these communities in Papua New

Guinea.

A secondary purpose evolved at the last

moment. A few days before I was due to depart

for Papua New Guinea, I was handed a photocopy

of Waite’s handwritten journal of his New Guinea

expedition. I resolved to follow Waite’s footsteps,

as much as possible in the short time I had‘,

clarifying any obscurities in his journal (Waite

1918) and obtaining photographs and data that

related to the locations visited by Waite.

Tue New IRELAND COMPONENT

Originally I intended to go to the Sulka area

first, and I arranged to arrive in Papua New

Guinea in mid-December. In the meantime, Chris

Issac established that the people in his village of

*. For illustrations of this type of mask, see Helfrich 1973: Abb.1—45.

4. Six weeks for the whole exercise, including two weeks in New

Ireland Province.

Guma, on the south-east shore of Wide Bay

(Fig.2), were planning masked performances for

early January. I therefore changed the itinerary and

went first to New Ireland. I had previously

informed Noah Lurang that I would be arriving in

New Ireland about 7th January and could not get

through on the telephone to inform him of the

change of plans. When I got to Kavieng, the

capital of New Ireland Province, I found that he

had taken the opportunity to return to his village

on the island of Tatau (one of the Tabar group of

islands) over the Christmas—New Year period.

It happened that Waite’s main area of activity

had been the north-west coast of New Ireland and

the Tabar group of islands to the north of New

Ireland (Fig. 3). Thus to find Noah I would be

following Waite’s travels of 75 years earlier. I had

the peculiar sensation of simultaneously existing

in two streams of time—being with Waite from

June to August 1918, and looking for Noah in

December 1992.

hay fo

A LI

YC toy

Te,

FIGURE 4. Youth demonstrating use of ‘friction drum’

(lunuat), photographed by Waite on 2nd July 1918 at

Lemusmus, south coast of New Ireland.

38 B. CRAIG

The first place I wanted to visit was

Lakurafanga. Waite had spent several days at

Lakurafanga Plantation, from 27th June till 5th

July 1918, as a guest of the manager Mr Ostram,

a Finn. With Ostram’s place as his base he

explored caves nearby to collect bats and narrowly

escaped death when he slipped into a small hole

in the roof of a deeper cave (Waite 1918: 21-22).

My informants suggested that the caves were most

likely those they call Ling-saksavak.

Waite walked across New Ireland to Lemusmus

on the south-west coast where Mr Hetreich, a

sailor and marine artist, had a small plantation.

Hetreich gave Waite a friction-drum (/unuat) for

the collections; these are still being carved and

used today and at Kavieng on the way out I was

FIGURE 5. Shark catching equipment held by Sialis of

Munun village, Simberi Island, Tabar Group, New

Ireland Province. The ‘propeller’ float is called kat and

the rattle is called sorkuoi (Photo by B. Craig, 27th

December 1992).

able to purchase one for the Museum. The /unet is

held between the legs, and the moistened palms of

the player’s hands are rubbed towards the body

over the ‘tongues’ of the drum to create a shrill

vibrating sound. These are played by men only, in

connection with funeral rituals, so it is hardly

surprising that when Waite tried to get a girl to

demonstrate its use for a photograph, she declined

(ibid.: 30). He was subsequently able to find a

youth who demonstrated its use (Fig. 4).

Back at Lakurafanga, Waite bought shark-

catching equipment. This consists of a rope noose

with a propeller-shaped float attached (Fig. 5).

The shark is ‘called’ by rattling a rattan ring of

coconut shells and the noose is slipped over the

shark’s head as it surfaces next to the caller’s

canoe. When the noose tightens and the shark

submerges, the propeller is dragged down through

the water and exhausts the shark, which is then

pulled to the canoe and clubbed to death. A film,

‘Shark Callers of Kontu’, was made in 1982 by

Dennis O’ Rourke showing how sharks are caught

this way.

During his time in this area, Waite collected,

purchased and was given a wide range of material,

including birds, insects, reptiles, and a meteorite

(gift of Rev. Peekel of the Catholic Mission at

Lemakot—ibid.: 25), as well as ethnographic

items (Fig. 6). Among the latter were many fine

figures and masks associated with malangan

funeral rituals. So great was his interest in these

carvings that he was soon nicknamed ‘Masta

bilong faiawud’ (master belong firewood) (Jones

1992; Craig 1994) because normally the

malangan carvings were burnt after the rituals

were completed.

I had been advised to stay with Lapaseng Meli

and his wife Tangalabo at Sali village, located on

the Boluminski Highway about sixty kilometres

from Kavieng. I explained to my hosts that I was

interested in seeing the site of Ostram’s house and

talking with anyone who might have been there

when Waite visited in 1918. Tangalabo brought

me to the house of a very old man named Pasa

Atunais (Fig. 7), of the village of Lakurafanga.

Pasa rose feebly from his bed for the interview; he

was about 90 years old and blind now but able to

recall that when he was a teenager a white man

travelled along Boluminski’s road collecting

things like snakes, caterpillars, butterflies and

artefacts. As it turned out, Pasa was the only

person alive today who could recall seeing Waite,

although a few others who were infants in 1918

had been told stories of the man’s collecting

activities.

FOLLOWING THE TRACKS OF EDGAR WAITE 39

FIGURE 6. Masks and spears collected and photographed by Waite on Sth July 1918 at Lunapai village, north coast

of New Ireland. Most of these items are currently on display in the Pacific Gallery, South Australian Museum.

I was taken to the site of Ostram’s house but all

that remained were two concrete house stumps

and a concrete water tank, bullet-scarred from

straffing during the Second World War.

From Lakurafanga, Waite and his assistant

Davis sailed to the Tabar Islands on a pinnace

owned by Van der Ghynste, manager of Kopo

Plantation at Maragat Bay on the island of Tatau.

They took ten hours to get there. I travelled by a

different route. I took leave of my gracious hosts

and continued along the coast to Konos, where I

stayed overnight. From Konos, the journey to

Tatau Island takes just over two hours by

outboard-powered, fibreglass ‘banana’ boat. The

sea between New Ireland and Tabar was relatively

calm; there are times when rough seas challenge

the nonchalance of the Tabar Islanders and even

they prefer to wait for calmer weather.

Van der Ghynste was nicknamed ‘Masta Sak-

sak’ and his pinnace was christened the ‘Sak-sak’.

Waite reports (ibid.: 22): “Wanderghinste grows a

beard. When the war broke out he said he would

not shave until it was over, so they call him sac

sac, which means grass (grown on the chin).’ In

fact, sak-sak means ‘sago’. The son of the Simberi

Islander who worked on Van der Ghynste’s boat

supplied another interpretation, explaining that

‘Masta Sak-sak was so named because, whenever

all other food supplies ran out, he had his

labourers go into the bush and prepare sak-sak

(sago) to feed himself and the plantation workers.’

Waite and Davis used the Kopo plantation

house of Van der Ghynste as a base to explore the

islands and add to the already large collections.

Waite’s journal records: ‘Obtained various

articles, at different villages, including an old

Malagan, estimated to be over 80 years old. For

this and a shark float I paid 2 Pounds, for a pig

net One Pound’ (ibid.: 43).

Upon arrival at the village of Tatau on the

north-west corner of Tatau Island, I was taken to

the house of Noah Lurang but he had gone off into

the bush. Melenga Lembiang attached himself to

me and suggested we go look for Noah. Some

hours later we met on a mountainside and I was

immediately sure that Noah was the perfect choice

for Provincial Cultural Officer and would provide

a most impressive performance at the Pacific Arts

Symposium. Over the next two weeks, we worked

together closely and he introduced me to Edward

Salle, the best carver in the ‘traditional’ (early

twentieth century) style. These were the two men

who would be invited to the conference in

Adelaide. I later bought a small figure, called

40 B. CRAIG

FIGURE 7. Pasa Atunais of Lakurafanga village, north

coast of New Ireland. Pasa remembered collecting

insects for Waite as a teenager in 1918 (Photo by B.

Craig, 21st December 1992).

walik (Fig. 8), from Edward and it is now on

display opposite the old figures collected by

Waite.

After a short while, Waite decided to visit

Simberi Island, the northern-most of the three

major islands of Tabar. Masta Sak-sak took him

and Davis across the passage between Simberi

and Tatau, the sea so rough that they almost

turned back. They landed at Simberi village and

walked north along the west coast past Willie

Pettersson’s plantation to his brother Carl

Pettersson’s plantation at Maragon Bay. We

followed Waite’s route and at Simberi village I

was shown a malangatsak figure with

outstretched arms carved by Tames (Thomas)

Litir, which I purchased the next day for the South

Australian Museum (Fig. 9). We left Simberi and

walked north. My guides were able to take me to

Willie Pettersson’s grave, marked by cordyline

plants in a grove of coconuts and wild ginger

undergrowth.

Carl Pettersson was known as Charles among

the Europeans in New Ireland, as Sali among the

Tabar Islanders, and as ‘The King of Tabar’ in

Swedish newspapers (Regius 1993). He had a

common law wife named Shindu from Tiripats on

the island of Tabar. When Waite arrived, she had

just had her sixth child by Sali. Waite

FIGURE 8. Figure called walik, carved by Edward Salle

of Tatau village, Tatau Island, Tabar Group, New Ireland

Province (Photo by B. Craig, 24th December 1992).

Purchased for South Australian Museum, Accession

number A.74145, and currently on display in the Pacific

Gallery of the Museum.

FOLLOWING THE TRACKS OF EDGAR WAITE 41

—

FIGURE 9. Figure called malangatsak, carved by Tames (Thomas) Litir of Simberi village, Simberi Island, Tabar

Group, New Ireland Province; Tames is at left (Photo by B. Craig, 26th December 1992). Purchased for South

Australian Museum, Accession number A.74141, and currently on display in the Pacific Gallery of the Museum.

fm (Oe

FIGURE 10. Charles Petterssen, his Tabar Island wife Shindu, and their six children, photographed in Swedish

national costume by Waite on 12th July 1918 at Petterssen’s plantation, Maragon, on the west coast of Simberi

Island, Tabar Group, New Ireland Province.

42 B. CRAIG

photographed the family with Pettersson and

Shindu in Swedish national dress (Fig. 10). I was

able to record brief histories of each of the children

and I met Uto, an elderly woman who had been

the wife of Sali’s third child, Hans (called Anis by

the villagers). I was shown the site of Sali’s house;

nothing but an old gnarled fruit tree remained

from those times, everything else being completely

obliterated by the Japanese, decay and forest

regrowth.

Waite and his host Sali took the opportunity to

investigate a turtle-breeding ground on tiny

Marwui Island off the west coast of Simberi. They

discovered a nest and counted 155 eggs in it.

Waite then set out to walk around the northern

half of Simberi Island to Pigibut where Meyer, a

German plantation manager, lived. Following

Waite’s trail, we were overtaken by darkness and

slept the night at Lava. Next day, at Munun, I met

Sialis, one of the last of the shark-callers on Tabar

Islands, and photographed the propeller-like float

and the coconut shell rattle (Fig. 5).

We walked on to Pigibut and there we saw

Meyer’s house—the outside walls had been

renewed but the concrete stumps, the timber frame

and floor were said to be all original. The house

had been obtained by Kennecott Mining Company

to provide a base for gold exploration and they

had added a large kitchen, mess hall, bunkhouses,

workshop and separate office to the old building.

Although this company had been gone for only a

couple of years, the motorbikes, tractors and other

vehicles were rusted through, looking as though

they had been there for decades.

Waite stayed overnight and reported (ibid.: 59):

‘G. M. gave me several things and natives brought

others which we purchased. I paid G. Meyer 13

shillings for Bird of Paradise coins: five, two and

one mark.’ A number of malangan carvings on

display at the South Australian Museum are

labelled ‘Pigibut’. Waite was supposed to be

picked up at Pigibut by Masta Sak-sak but the

seas were too rough. The boat retreated to a safe

anchorage at Katatar on the south coast and Waite

continued walking, thus circling the island.

Returning to Van der Ghynste’s plantation at

Kopo on Tatau Island, Waite decided to go by

outrigger canoe and on foot to Klett’s plantation at

Tomalabat, overlooking the narrow passage

between Tatau and Tabar to the south. On the

way, he collected marine animals among the

mangroves and visited ‘Anus [Anoos] Rock’.

Anoos Rock is not marked on the maps and I

assumed it was a mountain inland from Maragat

Bay. However, I was able not only to locate it on

the southern tip of Maragat Bay but also recorded

the story of an old man named Lome who had

been insulted by his wife and went into seclusion

on the coral limestone rock in protest. Every day

he was seen sitting in the sun on a rock wall he

had built and he ignored the pleading of the

villagers to return. He refused to eat, wasted away

and eventually died. His bones, and the wall he

built, are said to remain to this day.

I hired a boat to follow Waite’s path along the

west coast of Tatau. We called in at Maragat Bay

and visited the site of Kopo Plantation but there

was no sign that there had ever been a house there.

I was taken to Anoos Rock and shown the bones

of Lome.

Further south, we came ashore again and

slipped and slid along a very muddy track to get to

the old site of Marai village on a ridge-top. Waite

recorded: ‘At a village, Marai, on the summit, I

had a “cooler” [juice of the kulau or green

coconut], the boy climbing the coconut tree

without the usual ankle strap’ (ibid.: 60). The site

now is completely unrecognisable as having been

a village. Cement grave markers had been erected

there during the late 1970s but were already

overgrown.

We returned to our boat and continued south to

Tamalabat and I photographed the view south to

correspond to a panorama Waite had sketched in

his diary. We then left in the late afternoon with

insufficient time to return to Tatau village before

dark and insufficient fuel. We came ashore five

kilometres short of the village and poled the boat

in the shallow water inside the reef for two hours

to get there.

The night before I was due to return to New

Ireland, I arranged to record a series of songs

appropriate to the masks to be ‘danced’ in

Adelaide at the Symposium. Dancing a mask

without the rhythm of the garamut and the

appropriate songs would be an absurdity. After a

few trials, with the elderly Joel Pitsia as lead

singer, and a master drummer to beat the slit-

drum, the group provided an hour of fine singing.

They asked me to play the recording back to them

to ensure it was satisfactory.

Also a master weaver, Joel Pitsia specialised in

constructing the woven malangan called worwora

and in making the various masks called ngeis

which are not carved but constructed of fibrous

materials. I bought three of his small worwora

and it was agreed that Noah and Edward would

bring to the Symposium not only two tantanua

masks and two carved wanis masks (with the

intricately-carved ‘ear’ panels) but also a number

FOLLOWING THE TRACKS OF EDGAR WAITE 43

FIGURE 11 a, b. Malangan display structure (mi-rorou-

si-mi-matbu), with wowora disc malangan on right,

Tatau village, Tatau Island, Tabar Group, New Ireland

Province (Photo by B. Craig, 24th December 1992).

of ngeis, to be made by Pitsia.* Whereas tantanua

and wanis masks are usually represented in

collections of New Ireland material, the worwora

and ngeis are much rarer because they are

ephemeral, so I was pleased to be able to add

them to the Museum’s collections (Fig. 11a, b).

There had been rough weather for a week but

during this night the wind dropped, the sky cleared

and the voices of the group singing the songs for

the masks seemed to bring calm to the sea. Our

trip the next morning was uneventful, except that

we reached Konos on New Ireland with barely a

spoonful of fuel left in the tank. By then I had

come to trust the ability of my friends to cope with

their environment—not necessarily always to get

it right, but to deal with problems with humour

and resourcefulness.

They had taken me along the paths trod by

Waite 75 years before and provided information

that immensely enriched his brief diary entries. It

‘For illustrations of these types of masks, see Helfrich 1973: Taf.

XII, Abb. 47, 51, 58-64 (wanis); Taf. X, Abb. 114-19 (ngeis).

44 B. CRAIG

was a pleasure in April to show Noah Lurang and

Edward Salle first hand the material Waite

brought back from these islands and to return their

people’s hospitality to me. Edward was also able

to provide a great deal of information about the

carvings Waite had collected.

During the Symposium, Noah presented the

South Australian Museum with one of the wanis

masks (Fig. 12) as a gift from the clan leader, Joel

Pitsia; Noah gave two shell rattles and the two

slit-drum beaters used during the performance of

the wanis masks at the Symposium’s opening

FIGURE 12. Mask (wanis-si-mi-chur-bangbang)

brought to Adelaide for performance at the Opening

Ceremony of the 5th International Pacific Arts

Symposium in April 1993. This mask was carved by

Edward Salle of Tatau village, Tatau Island, Tabar

Group, New Ireland Province. It was presented as a gift

to the South Australian Museum by Noah Lurang, New

Ireland Provincial Cultural Officer, on behalf of Joel

Pitsia, Kuk clan leader, Tauvoi hamlet, Tatau village;

Accession number A.74146, It is currently on display in

the Pacific Gallery of the Museum (Photo by Trevor

Peters, SAM).

FIGURE 13. Sulka hemlaut mask, constructed by John

Telko and others, being danced at an initiation

ceremony, Guma village, Wide Bay, East New Britain

Province. Design on underside of ‘umbrella’ represents a

rainbow (akrei) (Photo by B. Craig, 9th January 1993).

ceremony. The other masks were purchased. From

the above account, it is clear what the South

Australian Museum got out of the exercise; and

those who attended the Symposium benefitted

immensely from the performance of the wanis and

from the explanation of the various types of masks

provided by Noah and Edward in a more academic

context later in the day. What Noah perceived to

be the benefits is best reported by quoting his letter

to the South Australian Museum’s Director, dated

29th April 1993:

It's been a wonderful time we had together over at

Adelaide and I wish to express how much I and the

other guys have enjoyed the Symposium. The most

satisfying thing about this event was the exposure

which has broadened our scope on international

events and issues, and of course the opportunity for

give and take [at the] gatherings. To me, this was a

great thing.

Since New Ireland has again made another

significant contribution to your collection, especially

with the gift of the wanis (WANIS-SI-MI-CHUR

FOLLOWING THE TRACKS OF EDGAR WAITE 45

BANGBANG) with two bundles of shells strung together

as rattles and two short lengths of cane to beat the

garamut, I would ask the museum to honour this

with respect and favourable consideration in the

future. I don’t mean that the gift was an investment,

no; but that maybe one day the Tatau people may

build a small culture house to exhibit their

malangans, the museum could become invaluable in

this way by assisting somehow.

To me the gift was an expression of good co-

operation and a better working relationship and of

course respect for one another. I hope this

understanding will prevail in years to come.

Noah’s letter is an expression of the importance

of reciprocity. Activities which are expressions of

Melanesian reciprocity may appear to be the same

as what Westerners call ‘investment’ but as Noah

has pointed out, there are differences. In our

culture, an investment is made to obtain a ‘return’.

Melanesian reciprocity is a cyclical rather than

linear phenomenon. It was initiated between the

South Australian Museum and New Ireland by

Waite but was in hiatus for some 75 years. Noah’s

hope is that the cycle has been reactivated and

will not be allowed to lie dormant again for such a

long time.

Tue SuLKA, New Britain COMPONENT

I flew from Kavieng to Rabaul to meet Chris

Issac and we purchased supplies for our stay in

his village. Guma is situated on the southern

shores of Wide Bay; the high mountains of

southern New Ireland are discernible on a clear

day from high ground behind the village. Waite

did not visit this area but obtained the spectacular

Sulka mask from Major H. Balfour Ogilvy who

was an officer of the military administration in

New Britain (Gash & Whittaker 1975: Plate 392).

The normal means for getting to Wide Bay and

points further south on the east coast of New

Britain is by small diesel-powered tugs which

function as passenger ferries, carry supplies in to

villages, missions and administrative centres, and

carry out copra and cacao.

The Sulka, and their culturally alike (but

linguisticly distinct) Mengen neighbours to the

south-east, are gardeners inhabiting a narrow

coastal fringe, fishing from small outrigger

canoes, hunting in the hinterland and raising pigs

for ceremonies such as those for which the

hemlaut and susu masks are danced. They obtain

FIGURE 14. Sulka hemlaut mask, under construction by John Telko and others, for an initiation ceremony at Guma

village, Wide Bay, East New Britain Province. Design on underside of ‘umbrella’ represents tips of fern leaves

(priau) (Photo by B. Craig, 7th January 1993). This mask was purchased for the South Australian Museum

(Accession number A.74139) and was danced by John Telko at the Opening Ceremony of the Sth International

Pacific Arts Symposium in April 1993. It is currently on display in the Pacific Gallery of the Museum.

46 B. CRAIG

some cash by harvesting copra and cacao from

their many small plantations. A variety of trees

produce nuts and fruits in abundance and the

chewing of betel nut is universal—more common

than smoking tobacco. Life is so pleasant in the

villages that the older men have to exert

considerable pressure to get the young men to go

to Rabaul to do occasional business.

The trip takes about 10 hours to Guma and we

arrived after dark. We were put ashore in a three-

metre dinghy and Chris’s family made us

welcome. The next day we were taken to a hut a

few hundred metres up behind the village where a

dozen young men were preparing two hemlaut

masks for performance in a couple of days’ time. I

photographed the completion of one of the masks,

which involved the delicate sewing of strips of

flattened pith to a rattan armature, and the

painting of the designs on the conical headpiece

and underside of the ‘umbrella’ form on top of the

masks. One of the designs, parallel bands of

colours spiralling towards the centre of the

‘umbrella’, is named akrei (rainbow); the other is

named priau (tip of fern leaf) (Figs. 13, 14). I

later discovered the name of the design on the

hemlaut in the South Australian Museum (Fig.

15): a hevotek kalogu (claws of a freshwater cray).

These interpretations are visual metaphors and

may be connected to clan totems but only a long-

term study would be likely to elucidate their full

cultural significance.

I realised I was going to have considerable

difficulty deciding which mask I would choose for

the performance in Adelaide—one was

sculpturally more interesting but the other had a

more interesting design on the underside of its

‘umbrella’. I was also concerned about how to

select the two men who would be invited to come

to Adelaide to dance the mask. It was a relief to

learn that the mask and the men had already been

chosen, on the basis of messages between Chris

and me and between Chris and his relatives in the

village. This clearly indicated that the project had

the understanding and enthusiastic support of the

men of the village. Similarly, the price asked for

the mask was kept at a modest amount, meant to

compensate those men who had assisted to make

the mask but who had not been chosen to go to

Adelaide.

There was to be an afternoon performance of

mask dancing at the village of Tagul, two hours

walk south-east along the coast from Guma. We

set out early in the afternoon, the sounds of the sea

on our left and of the wind in the coconuts on our

right. As we approached the village of Tagul

FIGURE 15. Sulka hemlaut mask of a rare type called

lopela, designed to be danced by two men

simultaneously. This mask was obtained by Waite in

1919 from Major H. Balfour Ogilvy, whom he had met

on his New Guinea Expedition in 1918. This mask

(Accession number A.7416) has been on display in the

Pacific Gallery of the South Australian Museum for

several decades. The design on the underside of the

‘umbrella’ represents claws of a freshwater crayfish

(hevotek kalogu). Exact provenance, and photographer,

are not known.

through a tunnel of trees, we could hear the

drumming and singing that accompanies the

performance of the hemlaut masks. We crossed a

creek, ascended sharply and there, coming toward

us from the large crowd gathered at the centre of

the village, was the first of three hemlaut being

danced that day: an apparently gigantic figure

shrouded in green palm leaves wearing a red

conical structure supporting a two-metre diameter

‘umbrella’ painted on the underside with curving

flower-like forms.

I followed the hemlaut into the preparation area

behind the village and ran into a line of eleven

susu masks getting ready to make their

appearance. A senior initiate rushed to head off

the maskers and berated them soundly for failing

FOLLOWING THE TRACKS OF EDGAR WAITE 47

to await their cue. His abuse in Mengen, assuring

an uncomfortable reward for a premature

performance, was punctuated with the English

emphatic: ‘Honest to God!’ In due course they

were given a signal and the masks moved in

solemn procession into the village (Fig. 16), their

shiny green palm-leaf body-shrouds swaying in

unison to the drumming of a group of singing

women.

The ceremonies in which hemlaut and susu are

danced are primarily for the initiation of boys and

girls, the boys being circumcised and the girls

having their noses pierced. The school holidays

over Christmas—New Year are therefore the period

in which the masks are most likely to be danced.

They are danced also for funerals, marriages

and other special events. In 1982 I witnessed the

induction of a Sulka man, Joe Quintaip, into the

Franciscan priesthood. This ceremony commenced

as a Catholic service presided over by the

Archbishop of Rabaul and ended with traditional

masked dances which included a hemlaut

performance. The masked performances were

doubly appropriate as Joe Quintaip’s ordination

was considered by his relatives as both an

initiation and a death—initiation into the

knowledge and rituals of the Church and a death

to normal village life.

The hemlaut for that occasion was designed to

illustrate a Christian story and since the ceremony

was conducted soon after Christmas, the theme

chosen was Bethlehem. Accordingly, beneath the

two metre diameter umbrella-like form

characteristic of the hemlaut mask, an elaborate

woven sculpture was created representing Mary,

Joseph and two angels kneeling around the baby

Jesus lying in a manger.

At Tagul, on this trip in 1993, I was most

fortunate to observe another complex hemlaut

mask, this time with a traditional Sulka story

represented beneath the ‘umbrella’ (Fig. 17). This

was the story of the two brothers named Noot, a

younger brother who lived in the hills and an older

brother who lived by the sea:

The younger brother was always coming to the older

brother’s place and seducing the women. This made

the older brother so angry that he captured a lot of

men of his own village and tied them by hands and

feet to poles and contemplated cooking and eating

them. But it didn’t look right so he released the men

and instructed them to bring pigs to take their place.

They had a big feast but this attracted the younger

FIGURE 16. Procession of Sulka susu masks, Tagul village, Wide Bay, East New Britain Province (Photo by B.

Craig, 7th January 1993). Two of these masks were purchased for the South Australian Museum (Accession

numbers A.74135, 74136) and are currently on display in the Pacific Gallery of the Museum.

FIGURE 17. Sulkanised Mengen hemlaut mask being danced at Tagul village, Wide Bay, East New Britain Province

(Photo by B. Craig, 7th January 1993). This mask has a woven sculpture beneath the ‘umbrella’, representing the

legend of two ancestral brothers named Noot.

brother again and this time he seduced the older

brother’s wife. The older brother began to plot his

younger brother’s death. He commenced the building

of a men’s house which required a very deep hole for

a carved and painted centrepost and lured his

younger brother into the hole, then dropped the post

into it, but the younger brother escaped by following

his dog, which had dug a tunnel out of there. Then

the older brother held a feast and invited the younger

brother, intending to kill him with a wooden club but

the younger brother sent an animated wooden

likeness of himself and again escaped death.

The two brothers then fought with slings and fire and

the younger brother took the older brother’s wife and

flew away, some say in an aeroplane to a great land

in the south, to become the ancestor of the white

people (B. Craig ms.).

The mask I saw danced at Tagul consisted of a

central figure representing the older brother Noot

holding up the ‘umbrella’, surrounded by a

representation of a man trussed up on a pole with

a trussed pig opposite, and an oval men’s house,

with a tall food storage bin in front, opposite the

younger brother’s dog.

The structure of the ‘Bethlehem’ and ‘Noot’

masks were the same: a central figure surrounded

by four significant elements of the story. The other

hemlaut masks danced at Tagul that day also

exhibited this four-part structure, though the

sculptural elements were simpler petal-like forms

or merely suggested by the design on the

underside of the ‘umbrella’®. But there is more to

these masking traditions than their forms: for

example, the songs that accompany their display

and performance. Rowena Hill, Conservator at the

PNG National Museum in 1982, recorded several

and had them translated (Hill 1982: 98-107). This

is a song to accompany a hemlaut mask:

A woman gave birth to a snake; her husband took

her to a magic reef where there lived a spirit called

kot. When the snake came out from her belly his

tongue was red and forked and [he] covered the

® I was struck by the similarity of these designs to those of mandalas

when, in September 1994, the South Australian Museum hosted a

group of Tibetan Buddhist monks who demonstrated the ‘sand-

painting’ of a Mandala for Peace. They set up their space

immediately in front of the old Sulka hemlaut mask in the Pacific

Gallery but missed the opportunity to allow onlookers to draw the

comparisons by covering up the case in which the Sulka mask was

displayed.

FOLLOWING THE TRACKS OF EDGAR WAITE 49

whole woman, wrapping himself around her like the

underneath of the ‘umbrella’ mask.

A song to accompany the dancing of the susu

masks:

A man was standing on top of a hill overlooking a

village called Indaru when he heard an echo,

bouncing from a cliff, of a woman singing. Out of the

waterfall came her two sons; they were as beautiful

as a pair of cockatoos.

They were sitting on top of the cliff, one on each side

of the waterfall. The man saw them both and as he

fixed his gaze they merged into the white flowing

waterfall and kept flowing and flowing.

The susu mask is dancing like a flower blowing in

the wind. Its skirt like leaves is swinging to and fro.

The mask is like the flower dancing on the stem.

One song which seems to be concerned with the

pan-human problem of communication between

men and women laments:

A woman was standing on top of a hill, and on her

face she wore a mask of [white] lime and plant juice.

When I tried to talk to her she refused to speak and

stood there facing the setting sun. When I

approached her she started to cry; I felt sorry for her

and started to cry in sympathy. Suddenly she

disappeared and I did not know what to do or say.

The rather magisterial and fearsome appearance

of the masks is belied by the sensitivity and

imagery of many of these songs.

After arranging to purchase two of the susu

masks for the South Australian Museum, and for

the ‘Noot’ mask to be put aside for consideration

by the PNG National Museum (the masks are

normally destroyed soon after performance), we

commenced our walk back to Guma. I fell into

step with the Provincial Minister for Education,

who is Sulka, and he assured me that the

Provincial Government is fully supportive of such

projects as the one we were engaged upon,

providing that there is full agreement among the

villagers and that the PNG National Museum is

supportive.

It rained heavily next morning and this

prompted the information that someone can be

commissioned to cause rain in order to spoil a

feast. Rain can be induced by placing stones,

carved as human heads, in a river and stopped by

taking them out. Another method is to saturate a

knotted rope with the juices of a particular plant

and then to spray the juices around by whirling

the rope vigorously; the spell is cancelled by

undoing the knots in the rope. The fear of rain

being brought on to spoil a feast is also common

among the New Irelanders.

That afternoon Chris and I went to the bush

shelter where the masks were being made for the

performance in Guma village. I continued

photographing the painting of the masks, and the

preparation of two masks called keipa. The keipa

mask is a simple cap of woven vine or rattan with

eyes painted on it and the usual body-covering of

palm leaves. They are sometimes referred to as

‘whipping masks’, as the wearers burst in while

the women are dancing during the first afternoon

of the festivities and lash those who are to receive

the food piled up in the middle of the village for

distribution. These masks are also used to ensure

a prompt supply of food to the young men who are

constructing and preparing the masks in the bush

shelter.

I was invited to offer myself as a recipient of the

attentions of the keipa masks that afternoon but

managed to convince my hosts that I needed to

make good use of my cameras during the event.

Sometimes the notion of ‘participant observation’

becomes distinctly uncomfortable!

After the whipping episode, the women returned

to their singing and in due course a man stood in

front of the heaps of tubers and pork and called

one-by-one the names of the recipients of the food.

Within a half hour the space had been cleared and

the food distributed to individual households.

At midnight I recorded an hour and a half of

singing and dancing. The men were clustered

around a painted sapling, something like the

European maypole, drumming and singing, with a

line of women circling them clockwise. Off to one

side a line of women and girls in traditional dress

sang separate songs. There were quite lengthy

pauses in the singing with some experimental

drumming and voice-testing as catalysts for what

appeared to be an impromptu selection of songs

from a widely known corpus. Chris told me that

some of the songs are in a very old language that

few people, if any, understand; that other songs

are metaphoric or contain oblique references to the

real meanings; and that others can be made up on

the spot—an opportunity for creativity. The

singing and dancing continued until dawn.

Next day the hemlaut masks were completed

and the palm-leaf body shrouds attached. The

shroud is in two parts—one attached to the lower

rim of the mask itself on a rigid frame and the

other on a flexible frame is worn by the dancer

suspended from shoulder straps. This enables the

mask to undergo amazing transformations in size.

When the masker crouches, the shroud diminishes,

concertina fashion, to about a metre in height and

the mask appears to be short and fat; when the

masker leaps into the air, the shroud extends fully

50 B. CRAIG

two metres in height and the mask appears

gigantic. West African maskers use the same

device to achieve identical effects.

The introductory women’s songs, to call the

masks, began about 3.30pm and, shortly after, the

first of the hemlaut masks was led into the village

by a crowd of singing men. When the mask

reached the men’s house at the other end of the

village where a young lad awaited circumcision,

the group of singing men stood back from the

hemlaut and allowed it to perform in a clear space

while they continued drumming and singing; then

it left—all over in about ten minutes. There was a

hiatus and eventually the women’s group

recommenced singing and drumming, calling the

second hemlaut. It was led in by the men and

when the space was cleared for its solo

performance, the mask was danced in sequences

where it was tilted 90° to the vertical to allow a

clear view of the design on the underside of the

‘umbrella’—a standard procedure during

performance. The crowd applauded the

performance. Then the two little girls who were

undergoing initiation were publicly presented,

followed by the circumcision of the lad in the

men’s house. The hemlaut continued dancing

around the men’s house and the men crowded

around whistling, yelling and singing, and

banging on the corrugated iron covering part of

the roof of the house so that the whole house

shook, to drown out the cries of fear and pain from

the boy. Men were drumming vigorously inside

the house as well. At about five o’clock in the

afternoon, dishes of cooked food were brought and

placed in palm leaves in front of the men’s house

and a squealing pig was secured to a pole and

carried off. The second hemlaut departed from the

village only to reappear later and attempt to

intimidate a crowd of children. Cooked pork was

distributed amongst the cooked tubers and then

women came with their dishes to collect their

portions.

That evening Chris relayed several criticisms of

the hemlaut performances. The first masker was

inexperienced and failed to ‘show the writing’, ie.

the underside of the ‘umbrella’, adequately. The

second masker was much better but made a

mistake by leaping up into the air during the

second song when he shouldn’t have. Apparently

everybody laughed at that; I had heard them

laughing but thought it was excitement.

In general, it was felt that the whole thing was

too rushed and should have taken place over four,

rather than two, days. The first day—when the

keipa masks whip the receivers of food—is called

elaton. The second stage, called mselor, should

take place three months later, to allow time to get

all the food ready; this requires the women to

perform their songs whilst the food is set out in

heaps for distribution. The third stage, elonpik,

should occur the next day; the women perform

again and the food is distributed to those invited

to the feast; the initiated men are invited to eat in

the bush shelter where the masks are being

prepared. On the final day, kamit, the masks are

danced as described above.

According to pre-colonial traditions, everybody

went through three stages of initiation. For males

this involved circumcision as boys, then the

wearing of the masks as youths, and finally teeth-

blackening (rarely performed these days). Girls

may have their ears and noses pierced at their first

initiation, but I was unable to discover what

actions, if any, were performed at the second stage

(which I assumed was first menses) and at the

third, which was when they were of marriagable

age’. The feast for circumcision and nose-piercing

is called pam-o-wlongtuk.

On the evening of the eighth day of my stay at

Guma, the boat arrived to take us and the masks

to Rabaul. When darkness fell, the hemlaut was

brought down to the beach and taken out to the

boat in the dinghy. The smaller masks had been

packed in cardboard boxes. The boat’s crew had

ensured there were no women or children on board

and we lowered the canvas awnings around the

deck to protect the hemlaut from sea spray and to

ensure the mask would not be seen by women and

children when we arrived in Rabaul in the

morning. Women and children are expected to

regard the masks as spirits, not as manufactured

items worn by men, which is why they are

normally destroyed after their performance.

We departed at 7.15 pm and the journey was

uneventful. We arrived at 5.15 am and had to wait

a couple of hours before the wharf was clear of

women and children and we could safely transfer

the hemlaut to a lock-up shed. I spent the rest of

the day and part of the next buying materials for a

large plywood crate, building the crate, securing

the masks inside and having it trucked out to

Airniugini’s cargo shed at the airport to await the

Dash-7 cargo plane scheduled for Tuesday. Chris

and I flew to Port Moresby on Sunday, passing

over an active volcano on the western side of New

Britain. The volcanoes around Rabaul, which were

to erupt in spectacular fashion during 1994, were

quiescent.

7. Such matters are not likely to be told to a male enquirer.

FOLLOWING THE TRACKS OF EDGAR WAITE 51

On Monday morning I received a call from

Rabaul saying the crate was too large for the

plane. Even though I had built it within the

dimensions provided by Airniugini staff in

Rabaul, they had forgotten the requirement for

walk space from the front of the plane to the rear.

I had to get back on a plane for Rabaul that

afternoon to take the crate apart and remove the

masks, which I then completely covered in yellow

plastic to disguise their nature. Next morning I

supervised the loading of the disassembled crate

and the masks, ensuring there was walkspace

through the cargo hold. On arrival in Moresby, we

unloaded the masks and crate into a hired truck

and transferred them to the PNG National

Museum where I arranged for fumigation and

space to reassemble the crate and repack the

masks.

DiIscussION

Airniugini sponsored the airfares of myself,

Chris Issac, the four New Guinea dancers and the

airfreight of the collection through to Sydney, thus

saving a considerable amount of expense. Even

so, the non-salary field expenses for the six-week

trip amounted to around $6 000 and freight from

Sydney to Adelaide, internal airfares and

accomodation for the four dancers, the purchase of

additional masks brought to the Symposium by

the New Ireland dancers, and the installation of

the material in the Pacific Gallery added about

another $5 000. Nevertheless, for the $11 000

spent, Symposium delegates were treated to two

magnificent demonstrations of masked

performances, the cultural significance of the

masks were explained by indigenous

spokespersons and a radical upgrade of two small

sections of the Pacific Gallery became possible.

The objects, photographs and information

obtained in New Ireland and New Britain

constitute a major additional resource linked to

existing collections of objects, photographs and

written material held by the Museum. The

possibility of collaborative exercises involving

Anthropology and Natural History researchers is

also indicated, especially in relation to the

ethnographic and natural history specimens

collected by Waite in New Ireland and Tabar

Islands.

One unforeseen outcome was the generation of

a debate about the motivations, ideology and

ethics of the whole exercise, including the Pacific

Arts Symposium itself (Chance & Zepplin 1993;

Zepplin 1993; Craig 1993b; Fergie 1994). For

example, Zepplin writes accusingly that:

Cultural objects and customs of the Pacific remain

appropriated within the intrepid, pith-helmeted

domains of anthropology, archaeology and natural

history museums (read: exotic, alien and boringly

impenetrable)—the authorised exposition of the

Other (1993: 15).

Although it is not apparent to the visitor in the

Pacific Gallery, the South Australian Museum is

committed to a program of consultation and

involvement of indigenous people with the cultural

material held in this institution. This has been the

major strategy in relation to Australian Aboriginal

material for at least a decade now. The exercise I

have described is one model for what might be

done in relation to the non-Aboriginal cultural

material. The stimulation of debate about the

success and propriety of the outcomes of the

opportunistic exercise I have described need not

be read as a threat, but rather as an indication that

we are moving in the direction of relevance to

contemporary concerns and have not run aground

in the shoals of nineteenth century attitudes and

values.

REFERENCES

CHANCE, I. & ZEPPLIN, P. 1993. Cannibal Cultures:

The (Un)making of the Modern Museum. Broadsheet

2 Bye ND, 11335

CORBIN, A. 1990. Salvage Art History among the Sulka

of Wide Bay, East New Britain, Papua New Guinea.

Pp.67—83 in: ‘Art and Identity in Oceania’ Eds. A.

Hanson & L. Hanson. Crawford House Press:

Bathurst, New South Wales.

CRAIG, B. 1993a. Sulka Danced Sculpture. Artlink 13

(2): 46-49.

CRAIG, B. 1993b. Letters: Cannibal Critics. Broadsheet

27) (ANID,

CRAIG, B. 1994. Masta bilong faiawud. Paradise 103:

29-31.

FERGIE, D. 1994. Letters: The cost of curiosity

cabinets. Broadsheet 23 (1): 8.

GASH, N. & WHITTAKER, J. 1975. ‘A Pictorial

History of New Guinea’. Jacaranda Press: Milton,

Queensland.

HALE, H. M. 1956. The First Hundred Years of the

South Australian Museum 1856-1956. Records of

the South Australian Museum, 12.

HELFRICH, K. 1973. ‘Malanggan.1: Bildwerke von

Neuirland’. Museum fur Volkerkunde: Berlin.

ay B. CRAIG

HILL, R. 1982. ‘Field Trip to the Sulka Area of Wide

Bay, East New Britain Province’. Conservation

Department, PNG National Museum & Art Gallery:

Waigani, Papua New Guinea.

JONES, P. G. 1992. The life of a ‘Museum Man’—

Edgar Waite’s diaries as an historical source. Records

of the South Australian Museum 26 (1): 73-15.

REGIUS, H. 1993. ‘Our Ethnographical Troops in the

Field’: Swedes and museum collecting in Melanesia

circa 1900. Paper delivered at 5th International Pacific

Arts Symposium, Adelaide, 12-17th April 1993.

THOMSETT, S. 1993. A History of the Pacific

Collections in the Australian Museum, Sydney.

Pacific Arts 7: 12-19.

WAITE, E. R. 1918. Anthropological Journey to the

Islands of New Guinea: The Personal Journal of Edgar

R. Waite, 1918. (Transcribed and assembled by Brian

R. Rutherford). Anthropology Archives, South

Australian Museum, Adelaide.

ZEPPLIN, P. 1993. Pacific Arts Symposium. Art and

Asia Pacific 1 (1): 15-18.

‘A SPECIAL INDUCEMENT”’: THE ESTABLISHMENT OF THE RECORDS OF THE

SOUTH AUSTRALIAN MUSEUM.

PHILIP JONES

JONES, P. G. 1995. ‘A special inducement’: the establishment of the Records of the South

Australian Museum. Records of the South Australian Museum 28(1): 53-59.

The first edition of the Records of the South Australian Museum was published on 24 May,

1918. While the journal has been published continuously for seventy-six years, the South

Australian Museum was nevertheless one of the last major Australian museums to launch its

own scientific journal. The delay was not due to any dilatoriness on the part of its staff in

producing scientific papers; it reflects instead the enduring success of the colony’s first scientific

journal, the Transactions of the Royal Society of South Australia. The crucial role of Edgar

Waite in the establishment of the Records is documented in this paper.

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

South Australia, 5000. Manuscript received 10 July, 1994.

Histories of colonial museums have generally

overlooked the role of scientific journals in

marking the transition to what Bassala has

described as ‘the final phase of a struggle to attain

an independent scientific tradition’ (quoted in

Moyal 1976: 4). These journals were crucial for

the professional legitimation of colonial science.

In Britain and Europe, scientific journals had

evolved from the reports produced of their

meetings by the academies, which had in turn

arisen from the Renaissance universities. In

colonial Australia, this long process was

compressed into little more than twenty or thirty

years. Adelaide’s Philosophical Society for

example, was formed in 1853, just seventeen years

after the colony’s foundation. It played a major

part in the foundation of the South Australian

Museum, and reconstituted as the Royal Society

of South Australia, produced the colony’s first

scientific journal, the Transactions of the Royal

Society, in 1878. This pattern was mirrored in the

other Australian colonies (Inkster & Todd 1986:

UD,

In their attempts to represent the gamut of

scientific endeavour, the journals of the scientific

academies and museums defied a trend towards

greater specialisation which had accompanied the

rapid increase in the volume of scientific journals

during the nineteenth century (Thornton & Tully

1971: 277-93). To a large degree, museum

journals have retained this eclectic character today.

Apart from Adelaide, the major Australian

museums of Sydney, Melbourne and Brisbane had

all founded their museum journals by the turn of

the century. The Records of the South Australian

Museum celebrated seventy-five years of

continuous publication in 1994, but while it is one

of the longest running museum journals, it was

founded considerably later than its sister

publications. The delay does not imply a relative

scientific immaturity or lack of independence in

Adelaide; cultural dependence upon Britain was

at least as high in other capitals. The historian of

science Jan Inkster has argued that:

Australian scientists remained peripheral to the

cultures of both the mother country and the colony.

They were spatially distant from Britain, but mentally

isolated from the provincial centres in which they

lived and worked (quoted in Sheets-Pyenson 1988:

14).

Adelaide was among the first Australian centres

to focus upon the issues of building indigenous

collections and regulating the unimpeded flow of

natural history and ethnographic specimens out of

the country (Kohlstedt 1984a; 1984b). Together

with the Canterbury Museum, the South

Australian Museum was also one of the first in

Australasia to establish an international reputation

for particular indigenous collections, notably

ethnology and palaeontology (Sheets-Pyenson

1988: 100).

The relatively late foundation of the South

Australian Museum’s Records can be attributed

to the very close relationship which its scientists,

and those of the University of Adelaide, had with

the Royal Society of South Australia. The

Society’s office holders were drawn principally

from the Museum and the University during the

decades of the 1880s, 1890s and 1900s, the years

in which an independent museum publication

54 P. G. JONES

might otherwise have arisen. The network of

Adelaide’s scientific and cultural institutions and

individuals was extremely tight, both physically

and socially. The proximity of the main

institutions and their libraries on North Terrace

fostered the growth of overlapping and

complementary scientific relationships. The South

Australian Museum’s role, particularly in the

fields of zoology and ethnology, was that of a

‘keystone’ institution, at least until the

establishment of the University’s Chair of Zoology

(1922), the Department of Entomology at the

Waite Institute (1927) and the Division of Animal

Nutrition, CSIRO (1929) (Edmonds 1986: 207).

Moreover, the siting of Adelaide’s Museum,

Library, Art Gallery, and University on the one

city block was unparalleled elsewhere in the

country. Inkster’s model of ‘mental isolation’ does

not seem suited to turn-of-the-century Adelaide.

The Museum’s Director from the 1880s to the

First World War was Professor Edward Stirling,

himself a founder of the University of Adelaide

Medical School and a Royal Society office holder.

His own prestigious award as a Fellow of the

Royal Society in London was made principally

because of his publications on Australian zoology

and palaeontology which appeared in the Society’s

Transactions. Stirling considered that the journal

was more than adequate to serve the needs of his

staff and the colony during this period. Stirling’s

University colleague and fellow Museum

Committee member, the geologist and zoologist

Professor Ralph Tate, used the Transactions as

his forum to remind readers of Adelaide’s capacity

as an independent centre of research. The British

Museum itself, he stated, ‘owes to us some of its

knowledge of the natural history of the colonies’

(Tate 1878).

Tate made these remarks in the 1878 inaugural

edition of the Transactions, for which he appears

to have been largely responsible. Tate played a

large role in reinvigorating natural science in

South Australia at this time:

From his arrival in 1876 Ralph Tate, professor of

natural science at the new university, the whole

status of the Adelaide [Philosophical] society was

changed, new rules were drawn up, and publication

was put on a sound and permanent basis by the

foundation of a regular journal. Scientific

contributions came in abundantly for the first time

and soon exchanges were arranged with societies

around the world (Inkster & Todd 1988: 114).

For the following forty years the Royal Society’s

journal remained the principal outlet for scientific

publications in South Australia. Through a period

in which the Museum was only beginning to

establish its collections and its scientific expertise,

the Royal Society, like others around the country,

provided a nucleus around which young scientific

communities could grow and mature. They had

attracted a broad-based audience for science and had

established comunication links within and beyond

Australia (Inkster & Todd 1988: 115).

In the meantime, the early years of the

Transactions were uncertain. The journal relied

upon the subscription of members with a

matching government grant and for several years

it was published irregularly. This fact prompted

the Museum’s Director of the early 1880s,

Wilhelm Haacke, to consider the publication of a

scientific journal under the auspices of the

Museum itself. The proposed journal would:

contain notices of new species of the Australian

fauna, anatomical monographs thereof, and notices

of new modes of mounting and preserving specimens

and apparatus thereof.’

Haacke, who had been appointed to his post

from the State Museum of Jena in Germany, went

so far as to propose a publisher—the firm of N.

Engelmann of Leipzig, Germany. The South

Australian Museum Committee instructed him to

make further enquiries as to the costs of printing

and publishing a journal, but by November 1884

Haacke had resigned and no further action was

taken. That year also saw the Museum’s official

amalgamation with the Public Library and the Art

Gallery under a new Act; this factor alone would

have lowered the chances of a new, independent

museum publication being developed.

The Transactions provided a convenient and

prestigious outlet for the publications of Museum

staff. Edward Stirling led the way as the

Museum’s Director, maintaining an active

publication record in the Transactions and

encouraging its use by other natural scientists and

ethnologists. During 1909 Stirling received a

request from the Perth Museum for a South

Australian Museum publication and he placed the

matter before his Committee for consideration.

Their conclusion was that any independent

publication by the Museum staff would ‘seriously

affect’ the Royal Society.2 This consideration

influenced any discussion concerning a scientific

journal for the following eight years.

The catalyst for a renewed debate was the

retirement of Edward Stirling from the directorate

1. Minutes of the Museum Committee, 9 January, 1883. GRG19/

364/1883, p.119, State Records Office, Adelaide.

2. Minutes of the Museum Committee, 8 September, 1909. GRG19/

364/1909, p.181, State Records Office, Adelaide.

THE ESTABLISHMENT OF THE RECORDS SS)

FIGURE 1. Edgar Ravenswood Waite, Museum Director

1914-1928, who was successful in convincing the

Museum Committee to recommend to the Board that the

Museum should issue its own scientific journal.

at the end of 1912 after thirty-one years of service,

and his replacement during April 1914 by Edgar

Waite (Fig. 1), previously Director of the

Canterbury Museum (Jones 1992).

Waite brought with him considerable experience

as an editor and contributor to scientific

publications. During his early career as Curator of

the Leeds Museum in England he had edited the

Museum’s journal The Naturalist. He had been an

active contributor to the Australian Museum’s

Records while Assistant Curator of Zoology, and

had gone on to found and edit the Records of the

Canterbury Museum in 1907. Waite was

conscious that the South Australian Museum

lagged behind other states in not publishing a

journal, but the Transactions continued to serve

his staff well. In fact, the average length of each

issue of the Transactions steadily increased during

the period after 1910: 257 pages in 1911, 273 in

1912, 496 in 1913, 528 in 1914 and 892 in 1915.

This in itself helped to focus the debate on the

need for an independent museum journal.

Waite first raised the issue of a scientific journal

for the consideration of the Museum Committee in

his Director’s Report of 1 November 1916. A Sub-

Committee was established to examine his

proposal and immediately encountered a familiar

obstacle. The main concern was that there had

been ‘a long standing idea that the Museum

Officers are compelled to supply original matter

for publication to the Royal Society of South

Australia only’. Unable to find any evidence of

any formal obligation of Museum staff to publish

in the Transactions, the Chairman of the Public

Library, Museum and Art Gallery Board asked the

Royal Society for clarification. The Society’s

Secretary, W. P. Gill, confirmed that no formal

obligation existed.*

This obstacle overcome, Waite was in a position

to advance the argument. He did so in a lengthy

formal proposal to the Board, dated 10 January,

ONG

The covert suggestion previously made and now

definitely formulated is that the S.A. Museum issue a

scientific publication to be devoted to the research

work of the staff, including records of the

expeditions, descriptions and illustration of objects in

the Museum, occasional notes and the work of

outside investigators dealing with Museum material.

The publication to be edited by the Museum Director,

to be issued at no stated intervals, but when material

to form a part is available, a suitable number of such

parts to form a volume.*

Waite stressed the benefits of an independent

scientific publication devoted to the Museum,

noting that an alternative suggestion had been

made for the launch of a publication representing

‘all the ramifications’ of the composite institution,

the Public Library, Museum and Art Gallery.® He

advanced five main arguments for an independent

publication. It would firstly relieve the pressure on

the Transactions which was forced to publish

unreasonably large volumes. It would enable the

Museum Library to benefit from receiving

publications in exchange from other scientific

institutions. These publications would be

‘precisely the kind of literature required for

Museum use’. Thirdly, in Waite’s view a scientific

3. J. R. G. Adams to W. J. Sowden, 6 December, 1916. Docket

no.103/1917/3737, State Records Office, Adelaide.

4. J. R.G. Adams to W. J. Sowden, 7 December, 1916; T. Gill to

General Secretary, Public Library, Museum and Art Gallery, 15

December, 1916. Docket no.103/1917/3748; 22524, State Records

Office, Adelaide.

5. ‘Proposals of the Museum Director’, 10 January, 1917', Docket

no.103/1917, State Records Office, Adelaide.

6. This suggestion appears to have been made by the Board’s

Chairman. Docket no.103/1917, State Records Office, Adelaide.

56 P. G. JONES

journal would raise the status of the Museum:

An individual may become quite famous locally but

unless he publishes the results of his labours he will

never be widely recognised and the same is true of an

Institution. The more widely known a Museum

becomes, the better the opportunities for exchange

and of attracting experts to examine and report upon

special collections or subjects.

Fourthly, Waite stressed the fact that without a

scientific journal, the South Australian Museum

was significantly out of step with other

Australasian museums; it was the only museum in

the region which was in a position to publish a

scientific journal, but did not. Brisbane, Sydney,

Melbourne and Perth Museums in Australia, and

Wellington and Canterbury in New Zealand, all

published their journals, as did the ‘principal

Museums in other British Colonies ... and most

Foreign Museums’. Waite’s final point was that

the journal would provide a ‘special inducement’

for Museum staff to publish their researches ‘in a

publication with which they were so closely

associated’. This, he considered, ‘would prove a

stimulus for diffident or indifferent workers’, but

added diplomatically ‘(I may here be thinking of

the future)’. Perhaps one of Waite’s most potent

arguments though, was stated almost as an

afterthought:

At present no new species can, for example, be

constitutionally published until December of any one

year, when the volume of the Royal Society of South

Australia appears; and recognising this, the Museum

Committee recently authorised the publication of new

records, out-side the pages of the Society.’

These arguments seemed convincing but Waite

was immediately countered by his authoritative

Curator of Insects, Arthur M. Lea. A prolific

author of scientific papers, Lea contrasted the

advantages of a Royal Society, as against a

Museum, publication. His conclusion was that the

latter still offered the best and most dependable

forum for Museum publications and that the Royal

Society’s library could not be bettered by the

Museum’s. Lea’s forthright letter to Waite ended:

The scientific workers in South Australia are

insufficient in numbers to justify two publications

dealing with practically the same subjects.’

The Museum Committee met on 31 January,

1917, the day following Lea’s letter, and decided

that:

7. ‘Proposals of the Museum Director’, 10 January, 1917, Docket

no.103/1917, State Records Office, Adelaide.

8. A.M. Lea to E. R. Waite, 30 January, 1917, Docket no.103/1917,

State Records Office, Adelaide.

consideration of the question of altering the existing

arrangements whereby scientific papers by Museum

Officials are published in the Transactions of the

Royal Society of South Australia be deferred until

after the European War.’

Waite was resigned to this setback and

continued with plans for the publication by the

Royal Society of the first illustrated catalogue of

South Australian fish species. It appears to have

been his specialised requirements for printing

illustrations to this catalogue which finally caused

the Museum Committee to reconsider Waite’s

suggestion for a museum journal. Events took a

sudden turn. Waite’s diary entry for Thursday, 18

September, 1917 recorded that:

the Committee will recommend that a publication be

issued by the Museum, my figure of £100 annually

being accepted as a basis.'°

Two days later the matter was dealt with by the

full Board of the Public Library, Museum and Art

Gallery. Waite recorded the result:

Board granted me the £25 requested for preparing

drawings of fishes and approved my suggestion to

issue a Museum “Bulletin”. I am to draw up

necessary details as size and general “get up”. My

present idea is for No.1 to contain my review of Aust.

Typhlopidae [Fig. 2], Notes on Fishes by McC

([McCulloch] and W. [Waite] and possibly a

catalogue of Australian lizards now being prepared

by Zietz.'!

The funds for the journal were to be drawn from

the Museum’s share of the Morgan Thomas

Bequest Account, which had been invested in

securities since its establishment in 1903. At this

early stage, the 100 pounds set aside for the

Records represented about one-tenth of the annual

income derived from the Museum’s share of the

bequest (Hale 1956: 75, 136).

It was considered necessary by the full Board of

Governors to mark their shift in publishing policy

in a letter to the Royal Society. The Board’s letter

contained a hint of apology,

gratefully acknowledging its indebtedness to the

society for making it possible for so many years

through the medium of its Journal to place before the

public the results of the scientific research of its

Museum officers. The benefit possibly may have been

of a mutual character, but the Board is sensible of

the fact that hitherto the expense of publishing at its

Own expense was not possible.

9. ‘Scientific Publications by Museum Officers’, 1 February, 1917,

Docket no.103/1917, State Records Office, Adelaide.

10. Waite, E. R. Diary no.64, 18 September, 1917. AA356,

Anthropology Archives, South Australian Museum, Adelaide.

11. Waite, E. R. Diary no.64, 21 September, 1917. AA356,

Anthropology Archives, South Australian Museum, Adelaide.

THE ESTABLISHMENT OF THE RECORDS

Phyllis Clarke, ged.

S AUSTRALIAN FISHULS.

FIGURE 2. A copy of Plate VII from McCulloch and Waite, Some New and Little-known

Australia, Records of the South Australian Museum, Volume 1(1).

Fishes from South

58 P. G. JONES

The Board still hopes, however, that it may continue

to offer for publication in the Transactions of the

Royal Society of S.A. papers that will be regarded as

acceptable. !

Such deference was irritating to Waite, who

was conscious of the Museum’s historic role in

providing the core of the Transactions’ scientific

publications. The careful reference to ‘mutual

benefit’ was insufficient for him, as revealed by

his private journal entry, following the Museum

Committee meeting which drafted the letter:

Howchin said that the Roy. Soc. would not in future

print Museum material & got a vote of thanks passed

to the Soc. for publishing the Museum papers in the

past, nothing said as to the value of the Museum

contribution to the Society!!'%

By early 1918 Waite had established the ‘size,

style, type, etc.’ for the new journal, and had been

able to assemble material for the inaugural issue.

He estimated its cost at ninety pounds. He also

prepared a statement for the Museum Committee,

of which Stirling was now the Chairman,

outlining the character of the new publication. It

was

to be devoted to research work of the Museum Staff,

including records of expeditions, descriptions and

illustrations of objects in the Museum or the property

of others, occasional notes, and the work of outside

investigators, dealing with Museum Material ... The

publication is to be edited by the Museum Director

and to be issued in parts at intervals when sufficient

matter is available: a suitable number of such parts to

form a volume. '4

An order was subsequently placed with the

Adelaide printers Hassell and Son, and by April

14th Waite had corrected the first proofs. Two

weeks later he received the news that the

opportunity for a collecting expedition to the

south-west Pacific had arisen. Progress on the

journal was now urgent, as Waite’s diary entry for

May 2nd reveals: ‘Now pushing on “Museum

Records” in case I have to go away’.'5

The ensuing six week period was a race against

time for Waite, with various administrative duties

to finalise, as well as final arrangements for the

journal. He pressured Hassells to publish as soon

as possible; finally they were able to produce the

12. J. R. G. Adams to the Hon. Secretary, Royal Society of South

Australia. 18 February, 1918, Docket no.103/1917, State Records

Office, Adelaide.

13. Waite, E. R. Diary no.64, 6 February, 1918. AA356, Anthropology

Archives, South Australian Museum, Adelaide.

14. E. R. Waite to Museum Committee, 31 January, 1918, Docket

no.103/1917, State Records Office, Adelaide.

15. Waite, E. R. Diary no.64, 2 May, 1918. AA356, Anthropology

Archives, South Australian Museum, Adelaide.

inaugural issue of the Records, together with

offprints, on the morning of June 24th, 1918 (the

official publication date was recorded as May

24th). At 4.30pm that June afternoon Waite left

by train to Melbourne, en route to his

disembarkation for the Pacific islands expedition.

The Museum’s Annual Report was written aboard

ship near Hinchinbrook Island, and posted from

Cairns.

The first issue of the Records contained

significant scientific contributions to the fields of

ichthyology and entomology. Contributions to

other branches of natural science and to

anthropology followed in the remaining three

issues of the first volume, published between 1918

and 1921. Waite was a prominent contributor to

this and subsequent volumes. His catalogue of

South Australian fishes, prepared with McCulloch

for the first issue, was subsequently reprinted in

1923 as one of the first Handbooks of the Flora

and Fauna of South Australia. Another prominent

contributor to the early issues of the Records was

Arthur Lea, whose strong advocacy of the

Transactions had presented a major obstacle to

Waite’s initiative. Lea contributed a major

summary of South Australian coleoptera to the

first four parts of the Records.

The Records of the South Australian Museum

enjoyed a decade of healthy activity under Waite’s

editorship until his sudden death in 1928. The

mantle then passed to Herbert Hale and eventually

to Norman Tindale, who took the journal through

to the 1960s. Waite’s model for the journal as a

forum for the publication of the ‘research work of

Museum staff’ has been largely adhered to

throughout its history, although the work of

‘outside investigators’ has become more

prominent in recent decades. At the close of the

first decade, three volumes of the Records had

been published, comprising sixty-seven papers, of

which forty-one had been prepared by members of

the Museum staff. The remaining papers, by

external researchers, dealt with material in the

Museum collections (Hale 1956: 136).

As a scientist who had published widely in both

hemispheres before his arrival in Adelaide, Waite

did not perceive or impose any restriction on the

journal’s geographical scope. In spite of this, the

journal naturally achieved a South Australian

focus. Waite’s eclecticism informed the journal’s

scientific trajectory through the 1920s and beyond.

Flute-player, theatre-goer, motor-cycle rider,

aquarium-keeper, Waite published in several

branches of natural history as well as in

anthropology. With Edward Stirling too ill to

THE ESTABLISHMENT OF THE RECORDS 59

complete his paper on the Aboriginal toas

acquired from the missionary J. G. Reuther, Waite

became the paper’s co-author and supervised its

1919 publication in the Records’ second Part.

Thus from the first volume, the journal represented

the main branches of scientific and cultural

investigation at the South Australian Museum.

Tindale’s era strengthened this cross-disciplinary

representation; he published strongly in

entomology, anthropology and, following his 1929

Devon Downs excavation, in archaeology.

The Transactions of the Royal Society did not

suffer the decline which might have been

anticipated. The revitalisation of South Australian

academic and research zoology which occurred

during the 1920s assured the Society of a

continuing flow of papers. Museum scientists

continued to publish in the Transactions as well,

a trend which has continued until the present.

After three-quarters of a century the five main

benefits which Waite predicted would flow from

the Museum’s publication of its own scientific

journal all appear to have been realised. The

pressure on the Transactions was relieved,

Museum staff were offered a ready inducement to

publish, the scientific status of the Museum was

enhanced and it was placed on a par with its sister

institutions. Most tangibly, the Museum’s library

benefited from a series of exchanges made

possible by the production of the Records.

Hundreds of periodicals have been received in

direct exchange for the twenty-seven volumes of

the Records published since 1918. These volumes

have contained more than 670 scientific papers,

shortly to be listed in a published cumulative index

assembled by Stan Edmonds (in prep.).

It is worth noting that the impetus for the

establishment of the Records came from the

vigorous efforts of Edgar Waite, newly arrived in

Adelaide. His impact on the publishing program

of the South Australian natural science community

can be likened to that of Ralph Tate a generation

earlier. Both were accomplished natural scientists

with eclectic interests, extensive publishing

experience and strong connections with national

and international scientific communities. Both

succeeded in reshaping the public face of scientific

research in South Australia.

REFERENCES

EDMONDS, S. (in prep.) Records of the South

Australian Museum Cumulative Index, Volumes

1(1)—27(2), 1918-1994.

HALE, H. M. 1956. ‘The First Hundred Years of the

South Australian Museum 1856-1956’. Adelaide,

South Australian Museum.

INKSTER, I. Scientific enterprise and the colonial

‘model’: Observations on the Australian experience in

historical context. Social Studies of Science 15: 677—

704.

INKSTER, I. & TODD, J. 1988. Support for the

scientific enterprise, 1850-1900. in R.W. Home (ed.)

“Australian Science in the Making’. Cambridge,

Cambridge University Press, pp.102-32.

JONES, P. G. 1992. The Life of a ‘Museum Man’—

Edgar Waite’s Diaries as an Historical Source.

Records of the South Australian Museum 26(1): 73-

IDs

KOHLSTEDT, S. G. 1984a. Historical records in

Australian Museums of Natural History. Australian

Historical Bibliography Bulletin 10: 61-82.

KOHLSTEDT, S. G. 1984b. Natural heritage: Securing

Australian materials in 19th Century Museums.

Museums Australia December, 1984: 15—22.

MOYAL, A. M. 1976. ‘Scientists in Nineteenth Century

Australia. A Documentary History’. Melbourne,

Cassell.

TATE, R. 1879. President’s Address. Transactions,

Proceedings and Report of the Royal Society of

South Australia xxxix—xli.

THORNTON JE a comULbiY, Roi J, 197k) Scientific

Books, Libraries and Collectors. A Study of

Bibliography and the Book Trade in Relation to

Science’. London, The Library Association.

REVISION OF THE HALIPLIDAE (COLEOPTERA) OF THE AUSTRALIAN REGION

AND THE MOLUCCAS.

BERNHARD J. VAN VONDEL

VONDEL, B. J. VAN, 1995. Revision of the Haliplidae (Coleoptera) of the Australian region

and the Moluccas). Records of the South Australian Museum. 28(1): 61-101.

The Haliplidae of the Australian region and the Moluccas are revised. Fifteen species are

recognized, of which Haliplus wattsi, Haliplus hydei, Haliplus storeyi and Haliplus timmsi are

described as new. A key to the species is provided and distribution maps are given. Types of

most species have been studied. Lectotypes have been designated for Haliplus signatipennis

Régimbart and Haliplus oberthuri Guignot. Haliplus nicholasi Watts is considered a junior

synonym of Haliplus ferruginipes Régimbart and Haliplus nigrolineatus Wehncke a junior

synonym of Haliplus testudo Clark.

B. J.van Vondel, Natuurmuseum Rotterdam. Correspondence: Roestuin 78, 3343 CV Hendrik-

Ido-Ambacht, The Netherlands. Manuscript received 1 November 1993.

Recently Watts (1988) revised the Australian

Haliplidae. He recognized eight species, of which

four were described as new. Because there are a

few more species outside Australia, but belonging

to the Australian faunal region, it seemed useful to

me to treat and illustrate these species together

with the Australian ones. Furthermore I would like

to propose some additions to Watts’ revision.

This revision treats the species present in

Australia, New Guinea, Seram and New

Caledonia. No Haliplidae are known from the

other islands in the region or from New Zealand.

METHODS

The methods used are those of Vondel (1991).

The terms used are explained in Figs 1 and 2.

MATERIAL

Material from the region, especially outside

Australia, is usually rare in the collections of the

institutions listed below. Expeditions by Michael

Balke and Lars Hendrich to West New Guinea,

very successful in collecting various water beetles,

did not produce Haliplidae. Manfred Jach could

collect only one Haliplus during his recent water

beetle expedition to the Central Moluccas

(Ambon, Seram).

Data on ecology are seldom present on labels.

According to Lawrence et al. (1987: 322)

Australian Haliplidae are algal-feeders and

bottom-dwellers and are usually found in lentic

fresh water, which in general also applies to most

non-Australian members of this family.

The material I had access to originates from the

following institutional and private collections:

ANIC — Australian National Insect Collection,

Canberra, Australia.

BMNH - Natural History Museum, London, UK.

BPBM - B. P. Bishop Museum, Honolulu,

Hawaii, USA.

CNCI -— Canadian National Collections,

Ottawa, Ontario, Canada.

CV — Collection B. J. van Vondel, Hendrik-

Ido-Ambacht, The Netherlands.

CW — Collection Dr C. H. S. Watts, Adelaide,

South Australia, Australia.

ISNB_ — Institut Royal des Sciences Naturelles

de Belgique, Brussels, Belgium.

MCZC — Museum of Comparative Zoology,

Cambridge, Massachusetts, USA.

MNHN -— Muséum National d’ Histoire Naturelle,

Paris, France.

MUNC — Memorial University of Newfoundland,

St. John’s, Canada.

MVMA-— Museum of Victoria, Abbotsford,

Victoria, Australia.

NHMV - Naturhistorisches Museum, Vienna,

Austria.

QMBA — Queensland Museum, South Brisbane,

Queensland, Australia.

— Queensland Department of Primary

Industries, Mareeba, Queensland,

Australia.

RMNH - Nationaal Natuurhistorisch Museum,

Leiden, Netherlands.

QPI

62 VAN VONDEL

FIGURE 1. Haliplus wattsi, dorsal view: an, antenna; as,

apical spur; bp, location of basal punctures, illustrated

for each species; cl, clypeus; el, elytron; fr, frons; 1.1, i.2,

1.3, etc, interval 1, 2, 3 etc; pr, pronotum; p.1, p.2, p.3,

etc., primary puncture-row 1, 2, 3 etc; st, setiferous

striole; su, suture; ve, vertex.

SAMA — South Australian Museum, Adelaide,

South Australia, Australia.

SMFD - Forschungsinstitut und Naturmuseum

Senckenberg, Frankfurt am Main,

Germany.

— University of Queensland, Brisbane,

Queensland, Australia.

UZMH -— Zoological Museum, Helsinki, Finland.

ZMUC — Zoological Museum, Copenhagen,

Denmark.

UQIC

SYSTEMATIC SECTION

The Haliplidae are represented in this region by

only one genus: Haliplus Latreille. The genus

Haliplus is divided into six subgenera, of which

FIGURE 2. Haliplus wattsi, ventral view: cp, metacoxal

lobe (or coxal plate); ep, elytral epipleuron; mp,

metasternal process; pe, proepisternum; pp, prosternal

process; ps, prosternum; s.5, S16 Se7euStemmite sO.)

(last sternite).

three occur in the Australian region. Guignot

(1935a) erected the subgenus Phalilus for H.

oberthuri. Guignot (1955) assigned H. bistriatus,

H. fuscatus and H. gibbus to the subgenus

Neohaliplus Netolitzky (1911).

The remaining species belong to the subgenus

Liaphlus Guignot (1928).

The species below assigned to the subgenera

Phalilus and Neohaliplus seem to form a group

having a combination of characters, which is so

far not found outside the Australian region. Most

characteristic of the species in this group is the

lack of genal lines behind the eyes while all other

world Haliplidae have one or two genal lines. The

type-species of Neohaliplus: Haliplus lineatocollis

(Marsham) has two genal lines. Further study,

based on a worldwide survey, is needed to

determine if the Australian species indeed belong

AUSTRALIAN HALIPLIDAE 63

to Neohaliplus, as they show strong differences

with the palaearctic species of Neohaliplus. The

species in the subgenus Liaphlus have two genal

lines. Representatives of Liaphlus in the presently

used context are spread worldwide, but according

to Beutel & Ruhnau (1990) there is, so far, no

evidence for the monophyly of this subgenus.

The Australian subgenera in their present

context can be distinguished by the following

characters:

1 — Pronotum with basal plicae (Fig. 3), right

paramere of male without apical digitus (Fig. 14)...

— Pronotum without basal plicae (Fig. 70), right

paramere of male with apical digitus (Fig. 78)........

Brora coo Acarteen enc rirondsbar er eaSdaco ere Rersparcs Liaphlus

2 — Last abdominal sternite with medial ridge (Fig.

NO EDD)) Renee key ements Sores Benen Oa” Phalilus

— Last abdominal sternite without medial ridge.........

coaseioniod Bossy aencde onan anocodacha Soom ane ee ta ay Neohaliplus

After the examination of about 390 specimens,

including types of all species except Haliplus

ferruginipes Régimbart and Haliplus bistriatus

Wehncke, I recognize the following species:

Haliplus oberthuri (Phalilus) Guignot, 1935b

Haliplus storeyi (Phalilus) sp. n.

Haliplus hydei (Neohaliplus) sp. n.

Haliplus fuscatus (Neohaliplus) Clark, 1862

Haliplus gibbus (Neohaliplus) Clark, 1862

Haliplus bistriatus (Neohaliplus) Wehncke, 1880

Haliplus australis (Liaphlus) Clark, 1862

Haliplus wattsi (Liaphlus) sp.n.

Haliplus testudo (Liaphlus) Clark, 1862

Haliplus nigrolineatus Wehncke, 1883. syn. n.

Haliplus signatipennis (Liaphlus) Régimbart,

1891

Haliplus ferruginipes (Liaphlus) Régimbart, 1891

Haliplus nicholasi Watts, 1988. syn. n.

Haliplus alastairi (Liaphlus) Watts, 1988

Haliplus timmsi (Liaphlus) sp. n.

Haliplus stepheni (Liaphlus) Watts, 1988

Haliplus sindus (Liaphlus) Watts, 1988

Key To Tue Species Or HALIPLIDAE OF THE

AUSTRALIAN REGION AND THE Mo.uccas

1 — Pronotal base usually with longitudinal plicae.

Between plicae strongly impressed (Fig. 3).

Prosternal process posteriorly strongly impressed

HWY TaaWTAINS: (CFE, 7) ascencnoscedocnecbane odoacuser dbasoonaonnnoteeds 2

— Pronotal base without longitudinal plicae (Fig. 70).

Prosternal process not strongly impressed in

Tail lea (igi 27/4) enema Su aanehs eNO a 8

2 — Last abdominal sternite with strong longitudinal

BIG SC KOLE KC | eereen etre ce tt nes sO tne ARLE oe ING 3

— Last abdominal sternite without ridge or keel....... 4

3 — Pronotal plicae absent or hardly recognizable (Fig.

15), base of pronotum strongly and coarsely

punctured. Last sternite with clearly visible ridge

in posterior half, which is hardly visible in lateral

WAIN (CENGS 222),, 213) saccooasadaconsaccesenodn Haliplus storeyi

— Pronotal plicae long and strongly bent (Fig. 3),

base of pronotum moderately punctured. Last

sternite with strong ridge, which is also clearly

visible in lateral view................ Haliplus oberthuri

4 — Pronotal plicae long, base of elytral puncture-row

5) SOD GROOVE sasconscocectnoasaeaqce Joounaecceoadcbannebecn 5

— Pronotal plicae short and straight, base of elytral

puncture-row 5 at most slightly grooved............... 6

5 — Pronotal basal plicae straight at base and

anteriorly hardly curved. Pronotum basally and

anteriorly at most vaguely darkened (Fig. 27).

Elytron with vague, hardly interrupted, dark lines

on primary puncture-rows. Metasternal process in

middle with dense hairy punctures (Fig. 31). Male:

protarsal claws clearly unequal in length (Fig. 30);

left paramere without recognizable solid digitus

and without hairs on apex (Fig. 34); penis narrow,

tip slightly widened, inner side gradually curved to

COPIGRI SENSO) Becera ce ewe te eet et ae rer Haliplus hydei

— Pronotal basal plicae curved and strongly bent

inwards before base. Pronotum distinctly darkened

along base and along anterior edge (Fig. 57).

Elytron with distinct interrupted dark lines on

primary puncture-rows. Metasternal process in

middle without dense hairy punctures (Fig. 61).

Male: protarsal claws equal in length (Fig. 60); left

paramere with solid digitus (sometimes wide and

then not always clearly recognizable) with hairs

on top (Fig. 64 or 67); penis wider in middle or

nearly parallel, inner side abruptly bent before top

(BUG, OS) OP OS) soscoascissocencaonsaoeos Haliplus bistriatus

6 — Females.................. Haliplus fuscatus or H. gibbus

SNA CS teste Saat wtvanace Sun re dl seen Trace ate Pree enh 7

7 — Penis very narrow (Fig. 45).........Haliplus fuscatus

— Penis broad (Fig. 55)...........:0:08 Haliplus gibbus

8 — Pronotum with longitudinal dark mark in middle.9

— Pronotum with dark mark at most anteriorly......10

9 — Secondary punctures on elytron weak and sparse.

Anterior elytral margin at most weakly serrate

(Fig. 97). Specimens from New Guinea..................

so pats hictmiek Mes cvestain cosy eel Haliplus signatipennis

— Secondary punctures on elytron very strong and

dense, almost as strong as primary punctures.

Anterior elytral margin clearly serrate (Fig. 106).

Specimens from Seram....... Haliplus signatipennis

10 — Uniformly yellow-brown. Pronotum with lateral

margins serrate (Fig. 147). Prosternal process

impressed in middle (Fig. 151).....Haliplus sindus

— Elytron usually with dark marks or stripes, but at

least with darkened punctures. Pronotum with

lateral margins not serrate. Prosternal process not

VTA! TN TRAN Les ccoscoccnccqvooscotecnesesocesoese50500008 11

64 VAN VONDEL

11 — Elytral puncture-row 5 basally ending in deep

transverse impression (Fig. 118)................00+ 12

— Elytral puncture-row 5 basally at most with few

punctures standing close together or more or less

COMPUTE Mt ee ees eee eee eee es, errr 14

12 — Elytron with dark band along base reaching to

puncture-row 5 (Fig. 136). Pronotum with or

without anterior dark mark (Fig. 136)..................

Soop tiene ane sees See ace tes Seeger eae Haliplus stepheni

— Elytral base at most with dark mark near

puncture-row 4. Pronotum without marks......... i)

13 — Metasternal process flat, at most with some

strong punctures on both sides close to each other

(Fig. 122). Elytron with secondary punctures near

base of primary row 5 strong, basal punctures of

primary row 6 strong. Elytron usually with well

defined maculation (Fig. 118)...Haliplus alastairi

— Metasternal process with clear pit on both sides

(Fig. 131). Elytron with secondary punctures near

base of primary row 5 weak. Basal punctures of

primary row 6 weak. Elytron usually with

interrupted dark lines, which are rarely connected

by vague marks (Fig. 127)........... Haliplus timmsi

14 — Elytron with extensive maculation, base broadly

darkened to puncture-row 5 (Fig. 112)....0......00.0.

Sean rs Seer ned UO th Haliplus ferruginipes

— Elytron without extensive maculation, base not

GiStincllyadanke Neda. == IS)

15 — Proepisternum strongly and densely, sometimes

even coarsely punctured. Prosternal process

usually slightly but gradually widening anteriorly,

not clearly narrowed before coxae (Fig. 83). Last

sternite with strong punctures on most of its

surface. Male: left paramere with small solid

GNBTING (BG, BD) orocoscrccnecvercnesere scorer Haliplus wattsi

— Proepisternum at most weakly punctured in

anterior part. Prosternal process slightly narrowed

before coxae, more or less sinuate, anteriorly

about as wide as posteriorly. Last sternite with

moderately strong to weak punctures on about

apiCallghial i¢eesee een aene eee (cen ee eee 16

16 — Elytron usually without dark stripes on primary

puncture-rows, only punctures darkened (Fig.

70). Row of secondary punctures along suture

very dense and in irregular row, elytral interval 2

with complete row of secondary punctures,

interval 4 with sparser puncture-row. Metasternal

process with lateral rows of strong punctures,

which usually lie in slight impression (Fig. 74).

Male: left paramere with small solid digitus (Fig.

(JO) RA Re SS 2 a, Sel Haliplus australis

— Elytron with continuous or hardly interrupted

dark stripes on primary puncture-rows (Fig. 88).

Row of secondary punctures along suture dense

and in regular row, interval 2 with about 6-12

punctures and interval 4+6 only anteriorly with a

few punctures. Metasternal process with clear

lateral impressions (Fig. 92). Male: left paramere

without solid digitus (Fig. 94).....Haliplus testudo

DESCRIPTION OF SPECIES

Haliplus oberthuri Guignot

(Figs 3-14)

Haliplus oberthuri Guignot, 1935a: 165.

Lectotype d (here designated) ‘N. Caled.;

TYPE; Det. Dr Guignot, Haliplus sg. Phalilus

oberthuri Guign., type; bistriatus Fauvel’

[Marais de l’anse Vata prés Nouméa]

(MNHN)[examined]

Haliplus bistriatus; sensu Fauvel 1883: 335, nec

Wehncke 1880. [Misidentification]

Haliplus bistriatus; Guignot 1935a: 36, 1935b:

164; Watts 1988: 25.

Haliplus oberthuri; Guignot 1955: 290.

Diagnosis

This species is easy to distinguish from other

species with pronotal plicae by the strong ridge on

last sternite.

Description

Length 3.0-3.3 mm, width 1.6-1.9 mm. Body

broad, parallel in middle (Fig. 3).

Head: Yellow-red, antenna yellow (Fig. 4),

palpi yellow, maxillary palpus with last segment

more than half length of penultimate segment.

Punctation anteriorly dense, sparser on vertex.

Distance between eyes 1.2x width of one eye.

Pronotum: Yellow-red, dark blotch along

anterior margin. Lateral border strongly convex,

finely margined, front corners strongly bent

downwards. Base wider than base of elytra, long

curved plicae at base reaching over half length of

pronotum, strongly impressed between plicae.

Punctation on disc dense, strong basally (Fig. 3).

Elytra: Yellow-red, dark lines on primary

puncture-rows 1+2 interrupted between dark

punctures in basal part, on rows 3-7 continous, on

rows 8+9 widely interrupted (Fig. 3). Primary

punctures except in basal part of row 1+2 very

dense and strong. About 45 punctures in row 1.

Basal 6-10 punctures of row 5 confluent, forming

clear plica. Secondary punctures very sparse and

usually very small. All punctures darkened except

in unstriped parts of outer rows. Suture apically

briefly margined. Laterally completely margined,

margin in middle hidden from above.

Underside: Body yellow-red; legs yellow-red,

darkened towards coxae; elytral epipleura yellow,

reaching sternite 5, strongly punctured in anterior

and posterior part. Prosternal process wide and

irregularly parallel-sided, strongly impressed in

AUSTRALIAN HALIPLIDAE

pe rieo oo

eee Te gcGtentel

Tay

o* oer ne SPEAR,

OCS ee TOOTS: Py MELD EA OS o's Y, ay

O09

eco oo?

Or mide p er

a © roca eneisin;o OR EROS e

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Sesion

FIGURES 3-14. Haliplus oberthuri, Paralectotype d: 3, dorsal view; 4, antenna; 5, punctures near elytral base and

suture; 6, tarsus of male fore-leg; 7, prosternal and metasternal process; 8, prosternal process in lateral view; 9, hind

tibia, 10, metacoxal lobes and sternites; 11, last sternite in lateral view; 12, left paramere; 13, penis; 14, right

paramere.

66 VAN VONDEL

apical half, strong punctured grooves on marginal

ridges, densely and strongly punctured on anterior

half, clearly margined on anterior edge (Fig. 7, 8).

Metasternal process with lateral ridges, formed by

row of strong punctures, strongly impressed

towards apical part, very sparsely and weakly

punctured (Fig. 7). Metacoxal lobes widely

rounded at apical part, finely margined on apical

comer, punctures fairly strong and dense, in

central part some coarse punctures (Fig.10).

Punctures on sternite 5+6 not forming clear row,

last sternite with strong ridge in middle (Fig.10,

11). Hind tibia without setiferous striole, longer

tibial spur clearly longer than first tarsal segment

(Fig. 9).

Male: Pro- and mesotarsomeres 1-3 widened,

tarsomere 1 more dilated ventrally, only

tarsomeres 1+2 with sucker hairs on ventral side.

Protarsal claws unequal in length (Fig. 6), meso-

tarsal claws slightly unequal in length. Penis and

parameres as in Figs 12-14.

Biology

The type material is found in a marsh (Fauvel

1883: 335).

Distribution (Fig. 153)

New Caledonia. Australia: Queensland. The

specimen I have seen labelled ‘New Zealand, N.

Cal.’ is obviously mislabelled.

Material examined: New Caledonia: | 6, lecto-

type; 1 ¢, N.elle Caledonie, Noumea, ex coll.

Gambey, R. Oberthur ded., paratype, Haliplus

(Phalilus) oberthuri Guignot (MNHN); 1 3, Coll.

E. Witte, Australien; 1 6, Coll. E. Witte,

bistriatus, N. Cal., N. Seeland [obviously

mislabelled](SMFD). Australia: 1 2, Yungaburra,

Q., Atherton Tab., Harvard Exp. Darlington,

Haliplus stepheni ms.nom, det. C. Watts

(MCZC); 1 3, Bne [Brisbane], H. fuscatus Clark,

Haliplus stepheni ms.nom, det. C. Watts 84 (CV);

1 d, unlabelled (QMBA).

Haliplus storeyi sp.n.

(Figs 15-26)

Type material: ¢ Holotype: Australia, N.T., 6

km E. Humpty Doo, 9.1i-4.iii.1987, R. I. Storey,

Haliplus bistriatus Wehncke, det. C. Watts 87,

T.12700 (QMBA); Paratypes: 5 2, same data as

holotype (3 in QPI, 2 in CV).

Diagnosis

This species differs from other species with

pronotal plicae by the ridge on last sternite and

from H. oberthuri by the faintly impressed

pronotal plicae and its smaller size.

Description

Length 2.4-2.5 mm, width 1.2-1.3 mm. Body

broad, parallel in middle (Fig. 15).

Head: Yellow-red, weakly punctured. Antenna

yellow, distal half of last segment darkened (Fig.

16), palpi yellow, maxillary palpus with last

segment about half length of penultimate segment.

Distance between eyes 1.5x width of one eye.

Pronotum: Yellow-red to yellow-brown. Lateral

borders strongly convex, finely margined, front

corners strongly bent downwards, hind corners

rectangular to slightly rounded. Long plicae at

base hardly visible or usually absent. Strongly

impressed along base. Moderately strongly, along

base strongly, densely and in hind corners coarsely

punctured (Fig. 15).

Elytra: Yellow-red to yellow-brown, vague

transverse maculation in middle and in apical part

(Fig. 15). Primary punctures strong and

moderately dense. About 29 punctures in row I.

Basal 6-7 punctures of row 5 confluent, forming

clear plica. Secondary punctures sparse and

usually very small along suture, almost absent on

intervals. All punctures darkened except in parts

of outer rows. Apical part of suture shortly

margined and with blunt dorsal tooth. Completely

margined, margin in middle invisible from above.

Underside: Body yellow-brown to brown, elytral

epipleura yellow-brown, reaching sternite 5, with

strong puncture-row in posterior part and two

strong rows in anterior part. Legs yellow-red to

yellow-brown, slightly darkened towards coxae.

Prosternal process wide and parallel-sided,

strongly impressed in apical half, strong punctured

grooves on marginal ridges, densely and strongly

punctured on anterior half, clearly margined on

anterior edge (Figs 19, 20). Metasternal process

with lateral ridges, formed by row of strong

punctures, only impressed anteriorly, very densely

punctured and hairy in middle (Fig. 19). Meta-

coxal plates not reaching sternite 5, widely

rounded at apical part, clearly margined along

posterior edge, punctures fairly strong. Fine

punctures on sternite 5+6 not forming clear row,

last sternite with keel in apical half, strongly

punctured (Figs 22, 23). Setiferous striole on

dorsal face of hind tibia on about half length,

longer tibial spur longer than first tarsal segment

(Fig. 21).

Male: Pro- and mesotarsomeres 1-3 widened,

tarsomere 1 more dilated ventrally, only

AUSTRALIAN HALIPLIDAE 67

FIGURES 15-26. Haliplus storeyi, Holotype d: 15, dorsal view; 16, antenna; 17, punctures near elytral base and

suture; 18, tarsus of male fore-leg; 19, prosternal and metasternal process; 20, prosternal process in lateral view; 21,

hind tibia; 22, last sternite; 23, last sternite in lateral view; 24, left paramere; 25, penis; 26, right paramere.

VAN VONDEL

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; 32, prosternal process in lateral view; 33,

27, dorsal view; 28, antenna; 29, punctures near elytral base and

suture; 30, tarsus of male fore-leg; 31, prosternal and metasternal process

hind tibia; 34, left paramere; 35, penis; 36, right paramere.

FIGURES 27-36. Haliplus hydei., Holotype 3;

AUSTRALIAN HALIPLIDAE 69

tarsomeres 1+2 with sucker hairs on ventral side.

Protarsal claws equal in length (Fig. 18). Penis

and parameres as in Figs 24-26.

Biology

Unknown

Distribution (Fig. 155) i

Only known from Humpty Doo, Northern

Territory, Australia.

Haliplus hydei sp.n.

(Fig. 27-36)

Type-material: Holotype ¢: Cardstone, Qld,

17-23.11.1966, K. Hyde, Haliplus bistriatus

Wehncke det. Watts, ANIC No. 111 (ANIC).

Paratypes: 10 d, 11 2, Same data as holotype (17

in ANIC, 4 in CV); 1 6, Cardstone, 32 km S. of

Ravenshoe, Q, 17.38S, 145.29E, 14.11.1968, K.

Hyde (ANIC); 3 6, 1 ¢, Australia, N. Qld,

Pinnarendi Stn, 60 km W. of Mt Garnet, 7.ii.1989,

D. Heiner (3 in QPI, 1 in CV).

Diagnosis

This species can be distinguished from H.

bistriatus, H. fuscatus and H. gibbus by the

pronotal plicae, being long and straight at base

and male with clearly unequal protarsal claws.

Description

Length 2.7-3.0 mm, width 1.4-1.6 mm. Body

oval, parallel to subparallel in middle (Fig. 27).

Head: Yellow-red to yellow-brown, weakly

punctured. Antenna yellow (Fig. 28). Palpi yellow

to yellow-brown. Distance between eyes about

1.2—1.6x width of one eye.

Pronotum: Yellow to yellow-brown, strongly

impressed base between long, curved to almost

straight plicae (Fig. 27). Lateral borders finely

margined, hind corners rectangular to slightly

rounded. Anterior half densely punctured; front

and hind corners strongly punctured; basal

depression weakly punctured.

Elytra: Yellow-brown to yellow-red. Vague

dark hardly interrupted stripes on primary

puncture-rows, slightly darkened along middle

part of suture. Primary punctures dense and

moderately strong, about 40 punctures in row 1.

Basal 6-8 punctures of row 5 in clear longitudinal

impression (Fig. 29). Puncture-row 7+8 not

reaching base, but united just behind base. All

punctures darkened except parts of row 8+9.

Sparse secondary puncture-row along suture. At

most some single secondary punctures on inter-

vals. Central base flat to weakly impressed. Com-

pletely margined, margin in middle invisible from

above.

Underside: Yellow to yellow-red, slightly darker

on prosternal and metasternal process. Legs

yellow to yellow-red, slightly darkened towards

coxae. Elytral epipleura yellow, reaching to

middle of sternite 5, with uncoloured punctures,

strong dense puncture-row on narrowed posterior

part. Prosternum anteriorly clearly margined.

Prosternal process nearly parallel, anteriorly wider

than posteriorly, in middle strongly impressed in

posterior 2/3, anterior edge strongly margined,

lateral ridges with groove formed by row of coarse

punctures, anterior part densely and coarsely

punctured, in posterior impression unpunctured

(Figs 31, 32). Metasternal process diverging

posteriorly, anteriorly impressed in middle, weakly

to moderately strongly punctured, in middle

usually slightly hairy, laterally with fine plicae

formed by punctures (Fig. 31). Metacoxal lobes

not reaching sternite 5, weakly and sparsely

punctured near suture, stronger and denser

punctured in lateral part. Sternite 4—6 with

posterior irregular puncture-row. Last sternite

especially on apex stronger punctured. Metatibia

without setiferous striole on dorsal face, but

posteriorly with kind of fine ridge on inner side,

longer tibial spur about as long as first tarsal

segment (Fig. 33).

Male: Pro- and mesotarsomeres 1—3 widened,

tarsomere 1 more dilated ventrally, only

tarsomeres 1+2 with sucker hairs on ventral side.

Protarsal claws clearly unequal in length (Fig. 30).

Penis and parameres as in Figs 34—36.

Biology

Unknown.

Distribution (Fig. 155).

Only known from North East Queensland.

Haliplus fuscatus Clark

(Figs 37-46)

Haliplus fuscatus Clark, 1862: 401. Holotype @,

no data [Clark gives Adelaide as locality]

(BMNH) [examined].

Haliplus fuscatus; Zimmermann 1920: 308; 1924:

141; Watts 1985: 27, 1988: 27; Lawrence et

al. 1987: 322.

Remarks

I failed to find reliable characters to distinguish

710 VAN VONDEL

between females of H. fuscatus and H. gibbus. As

the holotype of H. fuscatus is a 2, I am not sure

whether I have correctly associated it with males

which are treated here as H. fuscatus.

Both species have more or less the same

distribution. It is distinctly possible that the

holotype of A.fuscatus and the lectotype of

H.gibbus are conspecific, in which case at least

the ¢ specimens here treated as H. fuscatus

should belong to an undescribed species.

Diagnosis

This species is closely related to H. gibbus as

evidenced by general similarity, although the male

has a different penis, which is very narrow in H.

fuscatus.

Description

Length 2.4—3.2 mm, width 1.2-1.8 mm. Body

oval, but parallel to subparallel in middle (Fig.

3),

Head: Yellow-brown to brown-red, sparsely

punctured. Antenna yellow, last segment slightly

darker (Fig. 38). Palpi yellow to yellow-brown.

Distance between eyes about 1.3x width of one

eye.

Pronotum: Yellow-brown to brown-red, basally

strongly impressed and vaguely darkened between

short straight plicae, anteriorly with vague

transverse mark. Lateral borders finely margined,

hind corners rounded (Fig. 37). Strongly and

moderately densely punctured, in basal depression

only few sparse punctures.

Elytra: Yellow-brown to brown-red. Vague

markings on primary puncture-rows, along suture

and usually on intervals in posterior half and

along base to puncture-row 4 (Fig. 37). Primary

punctures dense and moderately strong, 25-35

punctures in row 1. Basal 4—5 punctures of row 5

not in clear longitudinal impression, but at most a

little confluent (Fig. 39). Puncture-row 7+8 not

teaching base, but united just behind base. All

punctures darkened except parts of row 8+9. Stria

(sometimes vague) along anterior 1/3 and

posterior 1/5 of suture. Fine secondary punctures

along suture and usually very fine punctures on

most of intervals. Surface with very fine, hard to

recognize micropunctuation in both sexes. Body

outline in posterior half a little bulbous and there

margin not visible from above. Central base

weakly impressed. Completely margined.

Underside: Brown-red, darker on prosternal and

metasternal process. Legs yellow-brown, slightly

darkened towards coxae. Elytral epipleura yellow,

reaching to middle of sternite 5, with uncoloured

punctures, only a few punctures on narrowed

posterior part, two puncture-rows on anterior part

weak and about equal in strength. Prosternum

anteriorly clearly margined. Prosternal process

parallel, in middle strongly impressed in posterior

3/4, curved anterior edge strongly margined,

laterally with narrow groove formed by row of

coarse punctures, weakly punctured (Fig. 41, 42).

Metasternal process diverging posteriorly, anteri-

orly impressed in middle, weakly punctured, late-

rally with fine plicae formed by punctures (Fig.

41). Metacoxal lobes not reaching sternite 5,

weakly and sparsely punctured near suture, a little

stronger punctured in lateral part. Sternite 4-6

posteriorly with very fine irregular puncture-rows.

Last sternite weakly punctured. Metatibia without

setiferous striole on dorsal face, longer tibial spur

about as long as first tarsal segment.

Male: Pro- and mesotarsomeres 1-3 widened,

tarsomere 1 more dilated ventrally, only

tarsomeres 1+2 with sucker hairs on ventral side.

Protarsal claws equal in length (Fig. 40). Metatar-

sal claws short and strongly bent. Penis and

parameres as in Figs 44-46.

Female: Metatarsal claws long and almost

straight.

Biology

This species occurs in swamps, temporary

ponds, rivers and lakes. A specimen was found in

a trout stomach. It is attracted to light.

Distribution (Fig. 157)

West Australia, South Australia, Victoria, New

South Wales ?(no males known), Queensland.

Material examined (identification based on ¢):

Australia: West Australia: 1 ¢, 1 2, Bickley Swp,

Rottnest Isl., x.1959, D.E. (CW, SAMA); 1 6, 1

?, Bulldozen, Rottnest Isl., W.A., x.1958, D.E.

(CW); 1 3, Rottnest Isl., Salmon Swp, x.1958,

D.E.; 1 2, Rottnest Isl., Barkers Swp, x.1959.,

D.E. (SAMA); 1 2, W.A., Rottnest Isl, x.1931,

Harvard Expedition Darlington (MCZC); South

Australia: holotype 2 (BMNH); Victoria: 1 3, 5

2, Vic. Eildon Weir, ix.1943, F. E. Wilson (5 in

MVMA, | in CV); 1 3d, Wyperfield Nat. Park,

Lowan Track, 35.35S, 142.05E, light trap,

16.xi.1973. S. Misko, Haliplus fuscatus Clark det.

T. A. Weir 1987 (ANIC); Queensland: 1 3, 4 @,

C. Qld, L. Galilee, 20.i.1983, Timms (SAMA).

Females of H. fuscatus or H. gibbus : Australia:

South Australia: 1 2, Parra Wirra, ix.1969, C.W.

(SAMA); Victoria: 1 2, Eilham, C. Oke; 1 2,

Birchip; 2 2, Carrum, 29.xii.1920, C. Oke; 1 2,

Coliban R., Trentham, F. E. Wilson; 1 9, E.

AUSTRALIAN HALIPLIDAE 71

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FIGURES 37-46. Haliplus fuscatus, 37, Holotype 2 ; 38-46, 3, Rottnest Island: 37, dorsal view; 38, antenna; 39,

punctures near elytral base and suture; 40, tarsus of male fore-leg; 41, prosternal and metasternal process; 42,

prosternal process in lateral view; 43, hind tibia; 44, left paramere; 45, penis; 46, right paramere.

72 VAN VONDEL

Moorabool R., 4 km W. Ballan, 10.vi.1976,

Neboiss; 1 2, Howitt coll.; 1 2, Clarkfield, 1.ix.1-

937, F. E. Wilson (MVMA); 1 2, Clarkfield,

4.xi.1941, A. D. Butcher, ex trout stomach (CV);

1 2, Dimboola, 36.27S, 142.02E, 22.x.1983, J. C:

Cardale, at light (ANIC); 1 @, Ringwood,

Haliplus gibbus Clark, Haliplus fuscatus det. Cc;

Watts 84; 1 2, Haliplus gibbus Clk, Howitt Coll.

(MVMA); New South Wales: 1 ?, Forest Reefs

(MCZC); 1 2, Bogan R., J. Armstrong (SAMA);

1 2, Willanora Bridge, 11 km N. of Mossgiel,

33.16S, 144.34E, dry swamp, 21.xii.1970, at light,

Britton, Misko & Pullen (ANIC); 1 2, Barrenbox

Swamp, interior N.S.W., 24.x.1979, Fields

(ZMUC); Federal state unknown: | ?, Sandham,

x.1935 (ANIC);

Haliplus gibbus Clark

(Figs 47-56)

Haliplus gibbus Clark, 1862: 400. Lectotype d

(designated by Watts, 1988), ‘S. Aust.,

Bakewell 59/24’ (BMNH)[examined].

Haliplus gibbus, Zimmermann 1920: 308; 1924:

142; Watts 1985: 27, 1988: 26; Lawrence et

al. 1987: 322.

Remarks

See remarks under H. fuscatus.

Diagnosis

This species closely resembles H. fuscatus,

from which the male can be distinguished by the

penis, which is broader in H. gibbus. I am not

able to distinguish the females.

Description

Length 2.4-3.2 mm, width 1.2-1.6 mm. Body

oval, but parallel to subparallel in middle (Fig.

47).

Head: Yellow to yellow-brown or yellow-red,

sparsely punctured. Antenna yellow, last segment

darker (Fig. 48). Palpi yellow to yellow-brown.

Distance between eyes about 1.3x width of one

eye.

Pronotum: Yellow to yellow-brown, strongly

impressed base slightly darkened between straight

plicae, which are 1/4 to 1/3 of length of pronotum,

anteriorly usually slightly darkened. Lateral fine

margins narrowed anteriorly, hind corners

rounded. Anterior half densely punctured, front

and hind corners coarsely punctured, in basal

depression row of strong punctures (Fig. 47).

Elytra: Yellow-brown to yellow-red. Dark

marks on parts of primary puncture-rows, along

suture and along base to puncture-row 5 and on

intervals. Primary punctures dense and moderately

strong, about 30 punctures in row |. Basal 34

punctures of row 5 in weak longitudinal

impression (Fig. 49). Puncture-row 7+8 not

reaching base, but united just behind base. All

punctures darkened except parts of row 8+9. Stra

along apical part of suture, sparse row of weak

secondary punctures along suture. No discernible

secondary punctures on intervals, except a few

large ones. Central base flat to weakly impressed.

Completely margined.

Underside: Yellow to yellow-red, slightly darker

on prosternal and metasternal process. Legs

yellow to yellow-red, slightly darkened towards

coxae. Elytral epipleura yellow, reaching to

middle of sternite 5, with uncoloured punctures.

Prosternum anteriorly clearly margined. Prosternal

process parallel, in middle strongly impressed in

posterior 2/3, anterior edge strongly margined,

laterally with groove formed by row of coarse

punctures, anterior part densely and coarsely

punctured, in posterior impression only weakly

punctured (Fig. 49). Metasternal process diverging

posteriorly, anteriorly impressed in middle,

moderately strongly punctured, laterally with

plicae formed by punctures (Figs 51, 52).

Metacoxal lobes not reaching sternite 5, weakly

and sparsely punctured near suture, stronger and

denser punctured in lateral part. Sternite 4-6 at

most with a few very fine punctures. Last sternite

with only a few very fine punctures, very short

fine keel on apical point. Metatibia without setife-

rous striole on dorsal face, longer tibial spur about

as long as first tarsal segment (Fig. 53).

Male: Pro- and mesotarsomeres 1-3 widened,

tarsomere 1 more dilated ventrally, only

tarsomeres 1+2 with sucker hairs on ventral side.

Protarsal claws equal in length (Fig. 50), metatar-

sal claws short and strongly curved. Penis and

parameres as in Figs 54-56.

Female: Metatarsal claws long and hardly

curved.

Biology

This species occurs in swamps, temporary

ponds and rivers and has been collected from

submerged vegetation at the margin of slow

moving streams (Watts, pers. comm.).

Distribution (Fig. 158)

Western Australia, South Australia, Victoria,

Tasmania, New South Wales ? (no males known

from this state).

AUSTRALIAN HALIPLIDAE

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FIGURES 47-56. Haliplus gibbus, Lectotype ¢: 47 dorsal view; 48, antenna; 49, punctures near elytral base and

suture; 50, tarsus of male fore-leg; 51, prosternal and metasternal process; 52, prosternal process in lateral view; 53,

hind tibia; 54, left paramere; 55, penis; 56, right paramere.

74 VAN VONDEL

Material examined (identification based on d):

Australia: West Australia: 1 6, Armadale,

vii.1960. D.E. (SAMA); 2 6, 1 @, Picton Junc-

tion, Swamp near Ferguson R., 30.xi.1965, Britton

& Uther Baker (2 in ANIC, 1 in CV); 1 36, 7 km

E. of Wuranga, xi.1981, K. & E. Carnaby (ANIC);

1 3, Bridgetown, 8.xi.1931, Australia Harvard

Expedition, Darlington (MCZC); South Australia:

lectotype 6 (BMNH); 2 d, 2 &, Williamstown,

ix.1961. C. Watts (SAMA); Victoria: 2 d, E.

Pomborneit, 24 km ESE Camperdown, temporary

pond, viii.1978-ii1.1979, P. S. Lake (ANIC); 1 o,

1 @,E. Vic., Yarra Riv., Warburton, F. E. Wil-

son; 2 6,1 2, Newhill Res., x.1945, F. E. Wilson

(MVMA); Tasmania: 1 3, 2 2, Launceston,

8.vi.1948 (ANIC); 1 3, Launceston (QMBA); 2

3, Launceston; 1 6, 3 2 (MVMA); 1 do, 1 &,

Launceston (ISNB); Federal state unknown: 3 2

syntypes (BMNH); 4 ¢ [no locality] (MVMA); 1

3, 1 2, Sandham, Goudie [?], x.1935 (MVMA,

CW),

Haliplus bistriatus Wehncke

(Figs 57-69)

Haliplus bistriatus Wehncke, 1880: 75. Type-

material: Not located. In the Wehncke collec-

tion in MNHN one @ is present originating

from Brisbane. In the description, however,

Adelaide is mentioned as type-locality.

Haliplus bistriatus sensu Fauvel 1883, nec

Wehncke 1880. [Misidentification].

Haliplus bistriatus; Fauvel 1883: 335;

Zimmermann 1920: 304; 1924: 71; Guignot

1935a: 36; 1935b: 164; Watts 1985: 27, 1988:

25; Lawrence et al. 1987: 322.

Diagnosis

This species can be distinguished from H.

fuscatus and H. gibbus by the long curved

pronotal plicae, from H. hydei by the basally

curved pronotal plicae and the metasternal process

not covered with dense hairy punctures and from

H. oberthuri and H. storeyi by the smooth last

sternite. Most males can be distinguished from

those of related species by the solid digitus on top

of the left paramere.

Description

Length 2.5-3.4 mm, width 1.2-1.6 mm. Body

oval, parallel to subparallel in middle (Fig. 57).

Head: Yellow to yellow-brown or yellow-red,

sparsely punctured. Antenna yellow, last segment

darker (Fig. 58). Palpi yellow to yellow-brown.

Distance between eyes about 1.2x width of one

eye.

Pronotum: Yellow to yellow-brown, strongly

impressed base narrowly darkened between long

curved plicae, anteriorly with dark central mark.

Lateral borders finely margined, hind corners

rounded. Anterior half densely punctured, front

and hind corners coarsely punctured, in basal

depression only a few sparse punctures (Fig. 57).

Elytra: Yellow-brown to yellow-red. Dark

interrupted stripes on primary puncture-rows,

darkened along suture and along base to puncture-

row 5. Primary punctures dense and moderately

strong, about 35 punctures in row 1. Basal 6-8

punctures of row 5 in clear longitudinal

impression. Puncture-row 7+8 not reaching base,

but united just behind base. All punctures

darkened except parts of row 8+9. Stria along

anterior 1/3 and posterior 1/5 of suture. No

discernible secondary punctures on intervals.

Central base flat to weakly impressed. Completely

margined.

Underside: Yellow to yellow-red, slightly darker

on prosternal and metasternal process. Legs

yellow to yellow-red, slightly darkened towards

coxae. Elytral epipleura yellow, reaching to

middle of sternite 5, with uncoloured punctures,

strong dense puncture-row on narrowed posterior

part. Prosternum anteriorly clearly margined.

Prosternal process parallel, in middle strongly

impressed in posterior 2/3, anterior edge strongly

margined, laterally with groove formed by row of

coarse punctures, anterior part densely and

coarsely punctured, in posterior impression only

weakly punctured (Figs 61, 62). Metasternal

process diverging posteriorly, anteriorly impressed

in middle, moderately strongly punctured, laterally

with fine plicae formed by punctures, longitudinal

and oblique backwards directed sutural lines

visible in middle (Fig. 61). Metacoxal lobes not

reaching sternite 5, weakly and sparsely punctured

near suture, stronger and denser punctured in

lateral part. Sternite 4-6 with posteriorly irregular

puncture-row. Last sternite especially on apex

stronger punctured. Metatibia without setiferous

striole on dorsal face, but posteriorly with kind of

fine ridge on inner side, longer tibial spur about as

long as first tarsal segment (Fig. 63).

Male: Pro- and mesotarsomeres 1-3 widened,

tarsomere | more dilated ventrally, only

tarsomeres 1+2 with sucker hairs on ventral side.

Protarsal claws equal in length (Fig. 60). Penis

and parameres as in Figs 64-69. Left paramere

with solid digitus (Fig. 64 or 67).

AUSTRALIAN HALIPLIDAE

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FIGURES 57-69. Haliplus bistriatus, 57-66, Brisbane; 67-69, Mt Mulligan: 57, dorsal view; 58, antenna; 59,

punctures near elytral base and suture; 60, tarsus of male fore-leg; 61, prosternal and metasternal process; 62,

prosternal process in lateral view; 63, hind tibia; 64, left paramere; 65, penis; 66, right paramere; 67, left paramere;

68, penis; 69, right paramere

76 VAN VONDEL

Biology

Specimens have been attracted to light in places

in open forest.

Distribution (Fig. 154).

Queensland and northern part of Western

Australia.

Material examined: Australia: Queensland: 1

3, Brisbane, Sharp, Dr Guignot visit 1939

(MNHN); 2 6, Caloundra, Qld, 24.iii.1963, C.

Watts (CW); 2 2, unlabelled (QMBA); 2 2, Bne

[Brisbane?], H. testudo Clark, Haliplus stepheni

ms.nom det C.Watts 84 (QMBA, CV); 1 6,1 &,

N. Qld, 21 E. Mareeba, 21.i.1991, at light, R. I.

Storey (QPI, CV); 1 2, Brisbane], Howitt Coll.;

1 6, unlabelled, Howitt Coll. (MVMA); 1 6,

NQ., Mt Spec., ii.1971, J. G. Brooks, Haliplus

stepheni ms.nom. det. C. Watts 84 (ANIC); 2 d,

5 9, nr Mt Mulligan, 31.i.1991, Larson &

Halfpap (MUNC); Western Australia: 1 °@,

CALM Site 13/4, 12 km S. of Kakumburu Mis-

sion, 14.25S, 126.38E, 7—11.vi.1988, T. A. Weir,

at light, in open forest (ANIC).

Haliplus australis Clark

(Figs 70-78)

Haliplus australis Clark, 1862: 400. Lectotype @

(designated by Watts, 1988), H. australis

Clark [yellow label], B.M., Type [white round

label with red margin], Lectotype [white round

label with blue margin], LECTOTYPE Female,

Haliplus australis Clark 1862, selected by C.

Watts, 1984. (BMNH)[examined].

Haliplus australis; Zimmermann 1920: 303;

ODA IA Wiatts aS oe 27 pelo oom)

Lawrence et al. 1987: 322.

Haliplus testudo; Watts 1985: 27, 1988: 23.

Remarks

Watts (1988) considered this species to be a

junior synonym of H. testudo. After examination

of type material of both names I regard them as

separate species.

Diagnosis

Males of H. australis have a solid digitus on

the apex of the left paramere, while H.testudo

lacks such a digitus. The penis of H. testudo is

more curved towards the tip. The elytra of H.

australis usually have no dark stripes on puncture-

rows, while H. testudo have well developed dark

stripes on elytral puncture-rows.

Description

Length 3.7-4.1 mm, width 2.1-2.3 mm. Body

wide oval, widest just in front of middle (Fig. 70).

Head: Yellow to yellow-brown, vertex slightly

darker, weakly and sparsely punctured, on vertex

a little stronger punctured. Antenna yellow (Fig.

71). Palpi yellow, last segment small, 1/3x length

of penultimate segment. Clypeus finely margined

anteriorly. Distance between eyes 1.3—1.6x width

of one eye. Eyes usually partly covered by

pronotum.

Pronotum: Yellow to yellow-brown, moderately

strongly and densely punctured along base and in

middle of anterior part, otherwise weaker and

sparser punctured, punctures in central part of

base and anterior part darkened. Punctures not

stronger than elytral punctures. Anterior margin a

little protruding between eyes, posterior margin

slightly impressed in middle. Lateral borders

straight, finely margined.

Elytra: Yellow without maculation, only suture

narrowly brown, punctures darkened but

darkening rarely forming continous lines. Primary

punctures strong and dense, about 48 punctures in

row 1. All punctures with clear hole in middle.

Puncture-row 1-7 not much differing in strength,

row 8 clearly weaker, row 9+10 very weak and

hardly darkened. Distance between row 10 and 9

and between row 9 and 8 clearly more than

between row 8 and 7. First three basal punctures

of row 5 confluent. Secondary punctures strong,

dense and partly doubled in sutural row on interval

1, dense in interval 2, 3, 5 and 7, less dense in

interval 4, in interval 6+8 only a few punctures in

basal and posterior part, in posterior part of

interval 9 continous dense row of weak punctures,

in row 10 a few weak punctures in central part.

Completely margined, anteriorly with about seven

very weak flat teeth, in apical part with about

twelve small sharp teeth.

Underside: Yellow to yellow-brown, elytral

epipleura yellow with uncoloured punctures. Legs

yellow to yellow-brown, slightly darkened towards

coxae. Prosternal process parrallel-sided, slightly

narrowed in central part, along both margins

impressed in posterior half, anterior edge clearly

margined, strongly punctured along margins and

with a few punctures in middle part (Figs 74, 75).

Prosternum strongly punctured and anteriorly

finely margined. Proepisternum at most weakly

punctured in anterior part. Metasternal process flat

in middle, strong punctures on both sides partially

confluent, more or less forming short impressions,

which do not reach anterior margin, otherwise

weakly punctured (Fig. 74). Metacoxal lobes fairly

AUSTRALIAN HALIPLIDAE

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FIGURES 70-78. Haliplus australis, 70-75, Lectotype ? ; 76-78, Paralectotype 6: 70, dorsal view; 71, antenna;

72, punctures near elytral base and suture; 73, hind tibia; 74, prosternal and metasternal process; 75, prosternal

process in lateral view; 76, left paramere; 77, penis; 78, right paramere.

78 VAN VONDEL

strongly punctured, in sutural area weaker punc-

tured. Outer protarsal claws of males seem to have

tooth on inner margin. Setiferous striole on dorsal

face of hind tibia about 1/8x length of tibia (hard

to see), it looks like there is on inner margin an-

other setiferous striole over apical half, longer tibi-

al spur about 2/3x length of first tarsal segment

(Fig. 73).

Male: Pro- and mesotarsomeres 1-3 widened

and with sucker hairs on ventral side. Penis and

parameres as in Figs 76-78. Left paramere with

small solid digitus (Fig. 76).

Biology

Specimens occur in rivers and ponds and have

been found in trout stomachs. Watts (pers. comm.)

found specimens in a well shaded woodland pond

that dries out completely in summer.

Distribution (Fig. 159)

Australia: South Australia,

Queensland.

Material examined: Australia: Federal state

unknown: Lectotype 2? (BMNH); 2 6, 3 &,

Howitt coll. (MVMA); Queensland: | 2, 1 6,

Glen Valley, ii.1951, VI-, F. E. Wilson (CW),

South Australia: 2 d syntypes, S. Australia, S.s.,

67-56, H. australis Clark, Australia (BMNH); 1

3,2 2, Chain of Ponds, 4.xii.1989, C.W. (CW);

1 3, Cheltenham, 20.iii.1925 (MVMA); Victoria:

1 ex., Flowerdale, 22.1.1968, R. E. Parrott (CNCD;

2 3, Jamieson, 20.iv.1943, F. E. Wilson; 1 @,

Yarra Riv., Melgrove, 4.1.1952, F. E. Wilson; 2

3,6 2, Glenmaggie, Weir, iv.1957, F. E. Wilson;

1 46,1 2, Lake Wendourec, ii.1945, F. E. Wilson;

2 6, Eildon Weir, ix.1943, F. E. Wilson; 1 6,

Melbourne, Howitt coll. (MVMA); 1 6,

Clarkfield, 31.x.1942, A. D. Butcher, Ex trout

stomach (CV); 1 6, E. Vic., Cann Riv., 28.1.196-

7, G. Monteith (UQIC).

Victoria,

Haliplus wattsi sp.n.

(Fig. 79-87)

Type-material: ¢ holotype, ‘Homehill, Qld,

7.4v.1963, CW.’ (CW). Paratypes: | ¢ and 1 2,

on same pin as holotype; 2 ¢, same data as holo-

type (CW, CV); | 6, Bandenberg, Queensland,

31.11.1963, CW, H. testudo det. C. Watts, 1991;

1 3, 25 km. N. Coen, Queensland, 29.i1x.1984, C.

Wrattis (Ow )s I oe, “Netra, IN, ©)...

Koombaloomba, 10.1.1962, E. B. Britton, B.M.

1962-153, At light (BMNH); 1 6, Nov. Holl.d.,

Rockkompt [?], Daniel (MNHN); | 2, unlabelled,

T12706; 1 6, Tambourine Mountain, Jan. 1898,

C.J. Wild, Haliplus alastairi ms.nom. det C.

Watts, T12707 (QMBA); 1 2°, S.E. Qld,

Atkinson’s Lagoon, 13 km N.W. Lowood,

10.ix.1978. J. King, Haliplus testudo Clark det.

C. Watts 1987; 1 6, 1 2, N. Qld, Mc Ivor River,

40 ml. N. of Cooktown, 7.v.1970, G. B. Monteith,

Haliplus testudo Clark, det. C. Watts 1987

(UQIC); 1 2, N. Qld, Iron Range, Cape York

Pen., 28.iv—-1.v.1968, G. B. Monteith, Haliplus

testudo Clark, det C. Watts 1987 (CV); 1 2, N.

Qld, Homestead, Silver Plains, Via Coen,

11.xii.1964, G. B. Monteith, Haliplus testudo

Clark Det. C. Watts 1987 (UQIC); 1 3d, 2 2,N.

W.A., Kununurra, 22.xii.1991-6.1.1992, R. I.

Storey; 1 6, 4 2, N. Qld, 10 km S. of Laura,

4 iii.1992, at light, J. Hasenpusch; 1 ¢, N. Qld,

Tolga, 27.i.1987, J.D. Brown, light trap, Haliplus

testudo Clark, det. C. Watts 1987; 1 d, Qld, 21

km S. Mareeba, 22.1.1991, R. I. Storey; 1 d, N.

Qld, Cow Bay, N. of Daintree, 25.i-7.11.1984, LC.

Cunningham, H.testudo C. Watts det. 1987; 1 6,

N. Qld, 7 km NE of Tolga, Feb. 1988, Storey &

De Faveri, light trap, Haliplus testudo Clark det.

C. Watts 1988; 1 3d, N. Qld, 7 km NE of Tolga,

MAR. 1987, Storey & De Faveri, light trap,

H.testudo Clark det C. Watts 1987; 2 3, 2 &,

N.T., 6 km E. Humpty Doo, 9.ii-4.1i1.1987, R. I.

Storey (QPI); 1 6, Papua, Loloki, c. 10 m. N. of

Pt. Moresby, 19.iii.1965, Stn No. 205, M. E.

Bacchus, B.M. 1965-120 (BMNH); | 2,

Cardstone, Qld, 17—23.11.1966, K. Hyde, Haliplus

testudo Clk det. C. Watts; 1 2, S. of Charleville,

Q., 9.v.1973, M.S. Upton, Haliplus testudo Clk

det. C. Watts; 1 ¢, Katharine N.T., at light,

9.i1.1968, J. A. L. Watson, Haliplus testudo Clk

det. C. Watts; 1 ¢, Coastal Plain Rsch. Station,

C.S.1L.R.O. nr Darwin, N.T., at light, 1x.1966, E.

C. B. Langfield (ANIC); 1 ¢, Cahills Crossing,

N.T., East Alligator River, 12.26S, 132.58E,

2.v.1973, E.G. Matthews, at light, Haliplus

?australis Clark, det T. A. Weir, 1980, Haliplus

testudo Clk det. C. Watts; 1 d, N.T., 6 km E.

Humpty Doo, 9.ii—4.iii.1987, R. I. Storey; 1 2, N.

Qld, 7 km NE of Tolga, Feb. 1988, Storey & De

Faveri, light trap, Haliplus testudo Clark det. C.

Watts 1988; 1 3, N. Qld, 10 km S. of Laura,

4.iii.1992, at light, !. Hasenpusch; 1 d, N. W.A.,

Kununurra, 22.xii.1991-6.1.1992, R. I. Storey

(CV); 1 2, Queensland, 160 km S. of Cooktown,

500 m, 28.i.1964, J. Sedlacek (BPBM); 1 3,1 &,

Qld, Port Douglas, 8.i.1991, D. Larson; 2 3, Qld,

5 km S. of Mareeba, 15.i.1991, Larson (MUNC).

Diagnosis

This species can easily be distinguished from

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IGURES 79-87. Haliplus wattsi, Holotype ¢: 79, dorsal view; 80, antenna; 81, punctures near elytral base and

86, penis; 87, right paramere.

suture; 82, hind tibia; 8

80 VAN VONDEL

H. australis and H. testudo by the proepisternum

being strongly and densely or even coarsely

punctured. It can also be distinguished from H.

testudo by the absence of dark stripes on the basal

area of the elytra, the prosternal process usually

being wider anteriorly than posteriorly, the

uncoloured pronotal anterior punctures, the weakly

serrate or nearly smooth apical elytral margin and

in the male the penis not being gradually curved

to the top and the left paramere having a solid,

sometimes very small, digitus.

Description

Length 3.4-3.6, width 1.9-2.1 mm. Body oval,

widest behind shoulders, strongly tapering behind

middle (Fig. 79).

Head: Yellow-red, darkened near antennae,

moderately strongly punctured. Antenna yellow to

yellow-red, first two segments brown (Fig. 80).

Palpi yellow-brown, last segment twice as long as

penultimate segment. Distance between eyes 1.3—

1.5x width of one eye. Eyes partly covered by

pronotum.

Pronotum: Yellow to yellow-red, moderately

densely and strongly punctured, base with a few

stronger punctures opposite elytral puncture-row

5. Lateral border straight, finely margined, margin

not reaching front corner. Base a little impressed

opposite elytral puncture-row 3—S.

Elytra: Yellow to yellow-red, suture and

puncture-rows darkened in posterior 2/3,

sometimes connected by vague marks. Primary

puncture-rows dense and moderately strong, 35—

40 punctures in row 1, anterior part of first 2 rows

weaker, basal 2 or 3 punctures of row 5 usually

wide and confluent. Row 9 very dense and

impressed in middle. Secondary punctures along

suture relatively strong and as dense as primary

punctures, sparser but still strong on intervals,

except on hardly punctured or unpunctured

interval 6+8. All punctures darkened, except some

in lateral rows, and with hole in middle. Complete

margin not always visible from above, anteriorly

serrate, posteriorly weakly serrate or sinuate.

Underside: Yellow-red to yellow-brown, some

darkening near prosternal and metasternal process.

Elytral epipleura yellow, almost reaching last

sternite, with strong weakly darkened punctures.

Legs yellow-red, femur and coxa brown to dark-

brown. Prosternum margined anteriorly, strongly

and densely punctured. Proepisternum strongly

and densely or even coarsely punctured. Prosternal

process slightly to clearly gradually diverging

anteriorly, laterally grooved in especially posterior

half, anterior edge clearly margined, moderately

strongly punctured (Figs 83, 84). Metasternal pro-

cess flat to slightly elevated in middle, weakly

punctured, laterally with a few confluent punctu-

res (Fig. 83). Metacoxal lobes not reaching

sternite 5, moderately strongly, near suture weaker

punctured. Sternites strongly punctured, last

sternite with very short fine ridge on apical point.

Setiferous striole on dorsal face of hind tibia about

1/6x length of tibia, longer tibial spur about 3/4x

length of first tarsal segment (Fig. 82), claws at

most 1/2x length of last tarsal segment.

Male: Pro- and mesotarsomeres 1-3 slightly

widened and with sucker hairs on ventral side.

Penis and parameres as in Figs 85-87. Left

paramere with small solid digitus (Fig. 85). Penis

dorsally dilated (Fig. 86).

Biology

Specimens have been found in rivers and are

attracted to light.

Distribution (Fig. 161).

Australia: Queensland, Northern Territory,

northern part of Western Australia. Papua New

Guinea.

Haliplus testudo Clark

(Figs 88—96)

Haliplus testudo Clark, 1862: 400. Lectotype ?

(designated by Watts, 1988), ‘lectotype [white

round label with blue margin], type [white

round label with red margin], 67-56, H.

testudo Clark, Australia, Lectotype female,

Haliplus testudo Clark, 1862, selected by C.

Watts 1984’. [2 2 2 on card, right one marked

as type]. (BMNH)[examined].

Haliplus australis sensu Watts 1985, nec Clark

1862. [Misidentification]

Haliplus nigrolineatus Wehncke 1883: 145.

syn.n.

Haliplus testudo; Zimmermann 1920: 316, 1924:

141; Watts 1985: 27, 1988: 23; Lawrence et

al. 1987: 322.

Haliplus australis; Watts 1985: 27, 1988: 23.

Remarks

Watts (1988) regarded this species as a senior

synonym of H. australis. I consider the two

species to be distinct, because of morphological

differences in penis and left paramere (see also

remarks under H. australis).

In the course of my study of the Neotropical

AUSTRALIAN HALIPLIDAE

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tures near elytral base and suture; 91, hind tibia; 92, prosternal and metasternal process; 93, prosternal process in

FIGURES 88-96. Haliplus testudo, 88-93, Lectotype 2 ; 94-96, Sydney: 88, dorsal view; 89, antenna; 90, punc-

lateral view; 94, left paramere; 95, penis; 96, right paramere.

82 VAN VONDEL

Haliplidae I examined a syntype of Haliplus

nigrolineatus Wehncke from Uruguay and

concluded that it is conspecific with H. testudo.

This is obviously a case of mislabelling (further

treatment in the revision of the Neotropical

Haliplidae)

Diagnosis

This species can be distinguished from H.

australis by the presence of dark stripes on the

basal area of the elytra, the prosternal process

usually not being wider anteriorly than posteriorly,

the coloured pronotal anterior punctures, the

clearly serrate apical elytral margin and in the

male the penis being gradually curved to the top

and the left paramere not having a solid digitus. It

can distinguished from H. wattsi by the

proepisternum being hardly punctured.

Description

Length 3.5-4.0 mm, width 1.9-2.2 mm. Body

oval, widest just in front of middle (Fig. 88).

Head: Yellow to yellow-brown, weakly

punctured, on vertex stronger punctured, clypeus

not margined anteriorly. Antenna yellow to

yellow-brown (Fig. 89). Palpi yellow to brown,

last segment 1/3x length of penultimate segment.

Distance between eyes 1.5—1.7x width of one eye.

Eyes partly covered by pronotum

Pronotum: Yellow-brown, strongly punctured in

central anterior area, along base and near lateral

margins; transverse area in middle almost

unpunctured, most punctures darkened. Lateral

borders slightly convex, finely margined. Slight or

no impression in front of base.

Elytra: Yellow-brown, black lines on primary

puncture-rows continous or sometimes weakly

interrupted. Suture darkened, most of base

narrowly darkened to puncture-row 5 or 6. Primary

punctures moderately strong and dense, about 35

punctures in row 1. Interval 9+10 not much wider

than interval 8. Puncture-row 9+10 with separated

black punctures except in apical part where short

lines may be present. First three or four basal

punctures of row 5 confluent. Secondary punctures

moderately strong, sometimes as strong as primary

ones, sutural row continuous with about 40-45

punctures, 7-10 punctures in interval 2, about 16

in interval 3, interval 4 with only about 4

punctures near base; interval 5,7+9 with sparse

row, interval 6,8+10 at most with some punctures

in basal area. Completely margined. Shoulders

weakly serrate, apical part with about 14 short

teeth.

Under side: Yellow-red to yellow-brown, legs

yellow-brown to brown, darkened near coxae.

Elytral epipleura yellow, reaching to sternite 7.

Prosternum margined anteriorly, sparsely

punctured. Prosternal process parallel, hardly

narrowed near coxae, on both sides with strong

groove over almost its total length formed by

strong punctures, anterior margin with clear ridge,

elsewhere weakly and very sparsely punctured

(Figs 92, 93). Metasternal process flat, on both

sides an almost continous groove formed by strong

punctures, elsewhere weakly and sparsely

punctured (Fig. 92). Metacoxal lobes strongly and

not very densely punctured, near suture weaker

punctured. Setiferous striole on dorsal face of hind

tibia weak, about 1/5x length of tibia, longer tibial

spur 1/2—2/3x length of first tarsal segment (Fig.

91).

Male: Pro- and mesotarsomeres 1—3 widened

and with sucker hairs on ventral side. Penis and

parameres as in Figs 94-96. Left paramere

without solid digitus (Fig. 94).

Biology

This species is found in rivers and in lentic fresh

water and is attracted to light.

Distribution (Fig. 160)

Australia: South Australia, Victoria, New South

Wales, Queensland, Northern Territory. The

specimen I have seen from Sumatra is obviously

mislabelled.

Material examined: South Australia: | d, 1 9

(ANIC); 2 6, 5.ii.1908, C. French (MVMA);

Victoria: 3 ex., Melbourne (ANIC); 1 &, Vic.,

Yarra Riv., Mellgrove, 4.i.1952, F. E. Wilson; 12

2, Vic., Jamieson, 20.iv.1943, F. E. Wilson; | 3,

1 , Victoria, Melbourne; 1 2, Victoria, Howitt

Coll. (MVMA); 5 6, 2 2, Vic., Moorobool R.,

iv.1932 & iv.1951, F. E. Wilson (6 in MVMA, 1

in CV); New South Wales: 2 6, Surr. Sydney,

N.S.W., Nikitin 1958 (ISNB); 1 2, N.S.W.,

Hornsby, 22.vii.1931, Harvard Exp. Darlington

(MCZC); Queensland: Lectotype 2; paralectotype

(on same card as lectotype) (BMNH); 1 °,

Caloundra, 21.iii.1963, C. Watts (CW); 2 ex.

Nov. Holl.d, Cape York (MNHN); 1 2°,

Queensland, Biggenden, 22.1.1975. H. & A.

Howden (CNCI); 2 2, Queensland (MVMA); 1

2, N. Qld, 11 km WSW of Petford, 3/4.iv.1988,

R. I. Storey, at light; (QP); 2 6, N. Qld, 7 km NE

of Tolga, ii.1988, R. I. Storey & De Faveri, light

trap (QPI, CV); 1 2, Queensland, Bundaberg,

31.11.1963, C.Watts; 1 ¢, Queensland, Gin Gin,

2.iv.1963, C.Watts (CW); 1 2, Q., Brisbane, N.

Pine R., 6.iii.1932, Harvard Exp. Darlington

(MCZC); 1 6, Ashgrove, 2.v.1931, H. Hacker; 1

3,1 @,N. Pine R., 10.iv.1933, H. Hacker; 2 6,

AUSTRALIAN HALIPLIDAE 83

Brisbane, x.1892, C. Wild (QMBA); 1 6, N. Qld,

Split Rock, 14 km S. of Laura, 23/26.vi.1975, C.

Monteith (CV); 1 ¢, Brisbane (QMBA); | 2, Q.,

Highvale, 8.1x.1965, B. Cantrell (UQIC); 1 6,

Qld, Brisbane, 2.viii.1964, B. Cantrell (UQIC); 3

ex., Catherine Cr. nr Collins Weir, 20.11.1990; 22

ex., nr Mt Mulligan, 31.1.1991, Larson & Halfpap

(MUNC); Northern Territory: 1 ¢, Goose Lagoon,

16.10S, 136.1E, 11 km SW by S of Borroloola,

17.1v.1976, at light, J. E. Feehan (ANIC); 1 °,

N.T., Horn Islet, Pellew Group, 25/31.i.1968, B.

Cantrell (UQIC); Federal state unknown: | ex.

(ANIC); 1 ex. without data; 1 ex. Nov. Holl. ex

Museo Thorey (MNHN); 1 6, 1 2, Australia,

coll. Wager (ISNB); 7 ¢, 5 2, Howitt coll.; 1 ¢,

1 2, unlabelled; 1 ¢, C. G. Oke (MVMA); 1 9,

unlabelled (QMBA). — Indonesia: 1 2, Fort de

Kock, Sumatra, 920 m., 1925, E. Jacobsen,

Haliplus pulchellus det. A. Zimmermann

{obviously mislabelled](ISNB). — Uruguay: | 6,

Montevideo, syntype Haliplus nigrolineatus

Wehncke [obviously mislabelled]|(MNHN).

Haliplus signatipennis Régimbart

(Figs 97-111)

Haliplus signatipennis Régimbart, 1891: 979.

Lectotype 2 (here designated), ‘?; N.Guinea

mer., Rigo, luglio 1889, L.Loria; signatipennis

Rég.; Museo Civ. Genova; Museum Paris, coll.

Maurice Régimbart 1908’ (MNHN)

{examined].

Haliplus signatipennis; Régimbart 1899: 187;

Zimmermann 1920: 316; 1924: 141; Watts

1988: 23.

Diagnosis

This species can be distinguished from other

species in the region by the longitudinal dark

mark on the pronotum.

Remarks

The specimen collected on Seram (Figs 106—

111) is differing from the typical form in the

secondary puncture-rows being almost as strong

and dense as primary puncture-rows. As this only

known specimen from Seram is a female it is not

clear if it represents a separate species.

Description

Length 3.4-3.7 mm, width 1.9-2.1 mm. Body

oval, parallel behind sligthly protruding shoulders

(Fig. 97).

Head: Red-brown to brown, darkened near

antennae. Weakly punctured. Antenna yellow-

brown to yellow towards end, first five segments

not longer than wide (Fig. 98). Palpi yellow-

brown, last segment very short, about 1/4x

penultimate segment. Distance between eyes 1.1—

1.2x width of one eye.

Pronotum: Yellow-brown, large dark

longitudinal mark in middle. Impressed basally,

opposite elytral puncture-row 5—7 with strong,

posteriorly well bordered impression. Moderately

strongly and densely punctured, near hind corners

a few widened punctures. Laterally not margined.

Elytra: Yellow-brown, very extensive dark

maculation along base, suture and on intervals.

Completely margined, no clear serration. Interval

2 slightly impressed. Primary puncture-rows

moderately strong, about 32 punctures in row 1,

row 4-7 stronger than adjacent rows. Secondary

punctures along suture dense and moderately

strong (Fig. 99), secondary puncture-rows on all

intervals, on interval 9 behind central dark mark

very dense row of secondary punctures. All

punctures darkened and with hole in middle.

Underside: Red-brown to dark brown. Elytral

epipleura yellow-brown, reaching to sternite 6.

Legs yellow-brown (tarsus) to dark brown

(femur). Prosternal process sinuate in anterior part

and close to apex, anterior edge margined,

laterally with clean grooves, in anterior third part

moderately strongly punctured, in posterior 2/3

weakly punctured (Figs 101, 102). Metasternal

process flat with lateral impressions, weakly

punctured (Fig. 101). Metacoxal lobes not

reaching sternite 5, moderately strongly, near

suture weaker punctured. Sternite 5+6 with dense

irregular puncture-rows, last sternite moderately

strongly punctured, apical point with short ridge.

Setiferous striole on dorsal face of hind tibia

about 1/5Sx length of tibia, longer tibial spur about

4/Sx length of first tarsal segment (Fig. 100).

Male: Pro- and mesotarsomeres 1—3 widened

and with sucker hairs on ventral side. Penis and

parameres as in Figs 103-105.

Biology

Specimens are collected in a stock pond, in a

Sago swamp and in bomb craters.

Distribution (Fig. 162)

Papua New Guinea. Indonesia: Seram.

Material examined: Papua New Guinea:

Lectotype 2 (MNHN); 3 d, 1 2, Madang, 5 km

N. Alexishafen, sago swamp, 9.iii.1991; 1 ¢,

Madang, 2 km W. Alexishafen, bomb craters,

l.iv.1991; 2 6, Ramu Villy, Brahman Mission,

84 VAN VONDEL

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te ey. <<

—

FIGURES 97-105. Haliplus signatipennis, 97-102, Lectotype 2; 103-105, ¢ from Ramu Vlly: 97, dorsal view;

98, antenna; 99, punctures near elytral base and suture; 100, hind tibia; 101, prosternal and metasternal process;

102, prosternal process in lateral view; 103, left paramere; 104, penis; 105, right paramere.

AUSTRALIAN HALIPLIDAE 85

a eo

POEs tae, Fi Viton Men deat Dae Q

FIGURES 106-111. Haliplus signatipennis, 2 from Seram: 106, dorsal view; 107, antenna; 108, punctures near

elytral base and suture; 109, hind tibia; 110, prosternal and metasternal process; 111, prosternal process in lateral

view.

si@el< frome, U7/sby OO, larson i 6, i ©,

Madang, Brahman Mission, 21 & 27.vi.1991, D.

& M. Larson (MUNC). Indonesia: 1 2, Seram nr

Wahai, leg. Jach 1989 (NHMV).

Haliplus ferruginipes Régimbart

(Figs 112-117)

Haliplus ferruginipes Régimbart, 1891: 979. The

type material (Papua New Guinea, Rigo) could

not be found in Genoa (MCSN), where it

should be, nor in Paris (MNHN).

Haliplus nicholasi Watts, 1988: 23. Holotype @,

‘Townsville, Qld Feb. 1972, T. Ingeldew, T-—

10793’ (MVMA) [examined] syn. n.

Haliplus ferruginipes; Zimmermann 1920: 305;

1924: 142; Watts 1988: 23.

Remarks

A specimen from Merauke, New Guinea is

considered to belong to this species as it matches

the description of H. ferruginipes, although its

length is 3.0 mm opposed to 3 4/5 mm as

mentioned in the description. This specimen also

resembles very much the types of H. nicholasi.

Although both species are only known from

females making checking of male aedaegi

86 VAN VONDEL

FIGURES 112-117. Haliplus ferruginipes, Holotype 2 of Haliplus nicholasi: 112, dorsal view; 113, antenna; 114,

punctures near elytral base and suture; 115, hind tibia; 116, prosternal and metasternal process; 117, prosternal

process in lateral view.

impossible, I consider H. ferruginipes and H.

nicholasi conspecific.

Diagnosis

This species can be distinguished from related

species by the widely darkened elytral base in

combination with puncture-row 5 not impressed

basally.

Description

Length 3.3-3.6 mm, width 1.8-1.9 mm. Body

oval, widest in middle (Fig. 112).

Head: Yellow-brown to yellow-red, anteriorly

near antennae darkened, moderately strongly

punctured. Antenna (Fig. 113) and palpi yellow-

red to yellow-brown. Distance between eyes about

1.5x width of one eye.

Pronotum: Yellow-red, strongly punctured, on

disc slightly weaker and sparser punctured, base

slightly impressed. Lateral borders margined,

margins narrowed anteriorly and hardly reaching

front corner. Hind corners a little rounded.

Elytra: Yellow-red to yellow-brown, extensive

dark confluent maculation along base to puncture-

row 6, along suture to at least puncture-row | and

on intervals. Primary puncture-rows moderately

strong and dense, about 30 punctures in row it.

basal punctures of row 5 not in an impression.

AUSTRALIAN HALIPLIDAE 87

Puncture-row 7+8 strong and a little impressed in

middle, united to one row long before reaching

base. Secondary punctures along suture nearly as

strong as primary row | (Fig. 114), sparse but

strong secondary punctures on interval 2, 3, 5,

7+9; interval 4, 6+8 unpunctured. All punctures

darkened and with hole in middle. Completely

margined, weakly and sparsely. serrate in anterior

part and weakly sinuate in apical part.

Underside: Yellow-red to yellow-brown,

slightly darkened near prosternal en metasternal

process, tarsi and tibia yellow-brown, femora

brown. Elytral epipleura yellow with strong unco-

loured punctures, reaching to sternite 6. Prosternal

process sinuate in middle and just before apex,

anteriorly finely margined, lateral puncture-rows

in slight impression, in anterior half strongly

punctured, in posterior half in middle sparsely

punctured (Figs 116, 117). Prosternum anteriorly

weakly margined. Metasternal process flat,

moderately strongly but sparsely punctured, a few

lateral punctures usually in slight impression (Fig.

116). Metacoxal lobes not reaching sternite 5,

strongly punctured, near suture weakly punctured.

Sternite 5+6 with strong and dense transverse

puncture-row. Sternite 7 sparsely punctured,

bulbous in lateral view, short clear ridge on apical

point. Setiferous striole on dorsal face of hind tibia

about 1/5x length of tibia, longer apical spur about

3/4x length of first tarsal segment (Fig. 115).

Male: unknown

Distribution (Fig. 156)

Australia: Northern Territory, northern part of

Queensland. Papua New Guinea. Indonesia: West

New Guinea.

Material examined: West New Guinea: 1 @, S.

Neth. New Guinea, Merauke, sea level, 1.iv.1955,

L. D. Brongersma (RMNH). Australia: 1 ?aholo-

type of Haliplus nicholasi Watts (MVMA); 1

paratype 2 [not ¢ as label suggests], Homehill,

Qld, 7.iv.1963, C.W., Nicholasi C. Watts 1984; 2

paratypes 2, Cairns, Qld, 16.iv. 1963, C.W.,

nicholasi C. Watts 1984 (SAMA); 1 2, N.T., 6

km E. of Humpty Doo, 9.ii-4.iii.1987, R. I. Storey,

Haliplus nicholasi ms.nom Det. C. Watts 1987

(QPI).

Haliplus alastairi Watts

(Figs 118-126)

Haliplus alastairi Watts, 1988: 24. Holotype 6,

12°36’S 132°52’E Magela Creek, N.T. 1 km

NNW of Mudginbarry HS. 25.v.1973,

Matthews & Upton (ANIC).

Remarks

Part of the material Watts (1988) considered to

belong to this species, belongs to a new species,

Haliplus timmsi, described in this revision.

Diagnosis

This species can be distinguished from the

related H. timmsi by the flat metasternal process

of H. alastairi opposed to the metasternal process

pitted on both sides of H. timmsi.

Description

Length 3.0-3.6 mm, width 1.7—2.0 mm. Body

oval, widest behind shoulders, clearly tapering in

apical 1/3 part (Fig. 118).

Head: Yellow-brown, clypeus and frons weakly

punctured, posterior part of vertex moderately

strongly punctured. Antenna yellow to yellow-

brown, fourth segment shorter than adjacent

segments (Fig. 119). Palpi yellow, last segment

about 1/2x length of penultimate segment.

Distance between eyes 1.4—1.7x width of one eye.

Pronotum: Yellow-brown. Moderately strongly

punctured, basal punctures opposite elytral

puncture-row 4+5 slightly stronger and lying in

transverse impression. Base hardly impressed.

Lateral borders weakly margined, margin

narrowed anteriorly and not reaching front corner.

Anterior edge slightly protruding in middle.

Elytra: Yellow-brown with distinct dark

markings. Darkened along most of suture and on

parts of intervals, darkening on parts of puncture-

rows. Primary puncture-rows moderately strong,

first three rows weaker and denser than others,

about 42 punctures in row 1. Row 5 abruptly bent

outwards and ending in an inwards directed strong

transverse impression, row 6 basally strong.

Sutural row of secondary punctures about as

strong as primary row 1 (Fig. 120). Secondary

punctures near basal impression of primary row 5

strong. Sparse row of secondary punctures in

interval 2, 3, 5+7. Interval 4, 6+8 with only a few

punctures in anterior part. Completely margined,

shoulders and apical margin serrate.

Underside: Yellow-brown to dark brown.

Elytral epipleura yellow to yellow-brown, reaching

to sternite 5. Legs yellow-brown to dark brown.

Prosternum anteriorly margined, strongly, but not

densely punctured. Prosternal process flat, nearly

parallel, slightly narrowed just before apex,

slightly diverging anteriorly, anterior edge

margined, posteriorly weakly punctured, anteriorly

stronger punctured (Figs 122, 123). Metasternal

process flat with on each side some strong

punctures, elsewhere weakly punctured (Fig. 122).

VAN VONDEL

"fees eeee 60 2 @

FIGURES 118-126. Haliplus alastairi, Holotype d: 118, dorsal view; 119, antenna; 120, punctures near elytral

base and suture; 121, hind tibia; 122, prosternal and metasternal process; 123, prosternal process in lateral view;

124, left paramere; 125, penis; 126, right paramere.

AUSTRALIAN HALIPLIDAE 89

Metacoxal lobes not reaching sternite 5, modera-

tely strongly to weakly punctured towards suture.

Sternite 5+6 moderately, but especially laterally

densely punctured. Sternite 7 fairly weakly

punctured. Setiferous striole on dorsal face of hind

tibia on posterior 2/3, longer tibial spur half as

long as first tarsal segment (Fig. 121).

Male: Pro- and mesotarsomeres 1-3 widened

and with sucker hairs on ventral side. Penis and

parameres as in Figs 124-126.

Biology

Specimens were attracted to UV light placed in

open forest.

Distribution (Fig. 163)

Australia: Northern part of Western Australia,

Northern Territory, Queensland.

Material examined: Western Australia: 1 2, N.

W.A., Kununurra, 22.xii.1991-6.i1.1992, R. I.

Storey (CV); 1 6, CALM site 13/4, 12 km S. of

Kalumburu Mission, 14.25S, 126.38E, 7-

11.v1.1988, T. A. Weir, open forest (ANIC);

Northern Territory: holotype d; 1 2 paratype

Katherine, 9.11.1968, at light, J. A. L. Watson

(ANIC); 2 2, N.T., Grotty Pond, Newry Stn,

8.11.1986, M. Tyler, M. Davies & G. Watson

(SAMA); 1 3, Arnhem Land, Maningrida, 5 m.,

16.11.1961, J. L. Gressitt, light trap (BPBM); 2 6,

1 2,N.T., Horn Islet, Pellew Group, 25/31.1.1968,

B. Cantrell (UQIC, CV); 1 2, N.T., 6 km E. of

Humpty Doo, 9.ii-4.iii.1987, R. I. Storey (QPI);

Queensland: 2 ¢ [on label indicated as ? 2],

Cairns, B. Allen (SAMA); 1 6, Q., Cairns,.

Darlington (MCZC); 1 2, N. Qld, Iron range,

Cape York Pen., 28.iv—4.v.1968, G. Monteith

(UQIE); 1 ex., Caims, €. J. W., PARATYPE

Haliplus alastairi Watts 1984, T.11164 (QMBA);

1 3,N. Qld, Weipa, 15/16.iii.1989, G. Dickinson,

at UV light (CV); 2 2, N. Qld, 10 km S. of Laura,

4 111.1982, at light, J. Hasenpusch (QPI); 1 d, 1 2

, Cardstone, 4-16.i1.1968, K. Hyde, paratype; 1 °,

Cook Town, N.Q., i.1971, G.B., paratype; 1 @,

King River, 14.30S, 143.20E, 22.vi.1968, F.

Parker, paratype (ANIC); 1 6, Qld, nr Mt

Mulligan, 31.1.1991, Larson & Halfpap (MUNC);

Federal state unknown: 1 d, Australia or Tasma-

nia, Ploson [?], Mackay (MNHN).

Haliplus timmsi sp.n.

(Figs 127-135)

Type material: Holotype ¢, Lake Buchanan,

Qld, 21.30S/45.50E, B. Timms, 25.1x.1953,

PARATYPE Haliplus alastairi C. Watts 1984

(SAMA). Paratypes: 4 2, N. T., Grotty Pond,

Newry Stn, 8.11.1986, M. Tyler, M. Davies & G.

Watson, Haliplus alastairi sp.nov. det. C. Watts

*86 (SAMA); 1 d, 1 2, N. Qld, Pinnarendi Stn

60 km W. of Mt Gamet, 7.1i.1989, D. Heiner

(QPI, CV); 1 db, 2 2, N. Qld, 10 km S. of Laura,

4.111.1992, at light, J. Hasenpusch (2 in QPI, 1 in

CV); 3 3,9 2,N. Qld 7km NE of Tolga, ii.1988,

Storey & De Faveri, light trap (12 in QPI, 1 in

CV); 1 2, N. Qld, 7 km NE of Tolga, iii.1987,

Storey & De Faveri, light trap (QPI); 1 6,1 2, 11

km WSW of Petford, 3/4.iv.1988, R. I.Storey, at

light (QPI, CV); 1 d, Katharine, N.T., at light,

911.1968, J. A. L. Watson, Haliplus sp.nov. det.

T. A. Weir, paratype Haliplus alastairi C. Watts

1984 (ANIC); 1 d, 2 2, Edge Hill, N.Q., ii.1954,

G.B. (2 in ANIC, 1 in CV).

Remarks

Some paratypes of Haliplus alastairi Watts

belong to this new species.

Diagnosis

This species can be distinguished from the

related H. alastairi by the metasternal process

being impressed on both sides.

Description

Length 3.0-3.6 mm, width 1.7—2.0 mm. Body

oval, widest behind shoulders, clearly tapering in

apical 1/3 part (Fig. 127).

Head: Yellow-brown, clypeus and frons weakly

punctured, posterior part of vertex moderately

strongly punctured. Antenna yellow to yellow-

brown, fourth segment shorter than adjacent

segments (Fig. 128). Palpi yellow, last segment

about 1/2x length of penultimate segment.

Distance between eyes 1.5—2.0x width of one eye.

Pronotum: Yellow-brown. Moderately strongly

punctured, basal punctures opposite elytral

puncture-row 4+5 slightly stronger, base with

clear impression. Lateral borders weakly

margined, margin narrowed anteriorly and not

reaching front corner. Anterior edge slightly

protruding in middle.

Elytra: Yellow-brown. Darkened along most of

suture, darkening on parts of puncture-rows, or

rarely with marks connecting puncture-rows.

Primary puncture-rows moderately strong, first

three rows weaker and denser than others, about

42 punctures in row 1. Row 5 abruptly bent

outwards and ending in an inwards directed strong

transverse impression. Row 6 basally weak.

Sutural row of secondary punctures weak. Sparse

row of secondary punctures in interval 2, 3, 5+7.

VAN VONDEL

cgi ren

: °

SRO RH n ono pio intens: a

: Ge RIGS trreroleic: oo gata © 92.

° C

FIGURES 127-135. Haliplus timmsi, Holotype d: 127, dorsal view; 128, antenna; 129, punctures near elytral base

and suture; 130, hind tibia; 131, prosternal and metasternal process; 132, prosternal process in lateral view; 133, left

paramere; 134, penis; 135, right paramere.

AUSTRALIAN HALIPLIDAE il

Interval 4, 6+8 with only a few punctures in ante-

rior part. Secondary punctures near basal impres-

sion of row 5 weak. Completely margined, shoul-

ders and apical margin serrate.

Underside: Yellow-brown to dark brown.

Elytral epipleura yellow to yellow-brown, reaching

to sternite 5. Legs yellow-brown to dark brown.

Prosternum anteriorly margined, strongly, but not

densely punctured. Prosternal process flat, nearly

parallel, slightly narrowed just before apex,

slightly diverging anteriorly, anterior edge

margined, posteriorly weakly punctured, anteriorly

stronger punctured (Figs 131, 132). Metasternal

process flat with on each side small very deep

impression, weakly punctured (Fig. 131).

Metacoxal lobes not reaching sternite 5, modera-

tely strongly to weakly punctured towards suture.

Sternite 5+6 moderately, but especially laterally

densely punctured. Sternite 7 fairly weakly

punctured. Setiferous striole on dorsal face of hind

tibia on posterior 2/3, longer tibial spur almost as

long as first tarsal segment (Fig. 130).

Male: Pro- and mesotarsomeres 1-3 widened

and with sucker hairs on ventral side. Penis and

parameres as in Figs 133-135.

Biology

Specimens were attracted to light.

Distribution (Fig. 164)

Australia: Queensland, Northern Territory.

Haliplus stepheni Watts

(Figs 136-146)

Haliplus stepheni Watts, 1988: 25. Holotype @,

Australia, N.T., Humpty Doo, 6 km E., 9.ii—

4.111.1987, R. I. Storey (SAMA)[examined].

Diagnosis

This species can be distinguished from related

species by the combination of a darkened elytral

base and elytral puncture-row 5 with a strong

transverse impression on base.

Description

Length 2.8-3.0 mm, width 1.5-1.6 mm. Body

long oval, widest in middle (Fig. 136).

Head: Yellow, weakly punctured. Antenna (Fig.

137) and palpi yellow. Last segment of maxillair

palpus about half length of penultimate segment

(Fig. 139). Last segment of labial palpus about 2/

3x length of penultimate segment (Fig. 140).

Distance between eyes |.5—1.6x width of one eye.

Pronotum: Yellow to yellow-red, usually dark

blotch on anterior central part. Moderately

strongly punctured, basally opposite elytral

puncture-row 4 transverse impression, surrounded

by strong punctures, lateral borders slightly

concave, margined except near front corner,

margin stronger posteriorly.

Elytra: Yellow to yellow-red, distinct dark

maculation consisting of: black suture reaching

secondary row 1 or in apical part reaching to

primary row 1, black band along base to puncture-

row 5, marks confluent with suture on disc and in

posterior part, large marks in anterior, in central

and in posterior part. Primary punctures strong

and with clear central hole, about 28 punctures in

row 1, basal punctures of row 5 in strong impres-

sion. Punctures in row 1+2 less strong than in row

3-6 or 7. Secondary punctures usually strong,

generally restricted to odd intervals. All punctures

darkened. Lateral sides margined, slightly

constricted in posterior part, serrate in anterior part

(about 7 teeth) and in posterior part (about 7

teeth).

Underside: Yellow to yellow-brown, legs

yellow to brown towards coxae. Elytral epipleura

yellow, reaching sternite 6, in anterior part with

strong darkened punctures. Prosternum margined

anteriorly, strongly punctured. Prosternal process

about parallel-sided, sligthly narrowed near coxae,

grooved along both sides, strongly and densely

punctured, clearly margined anteriorly (Fig. 142,

143). Metasternal process flat or slightly elevated

in middle, grooved along both sides, weakly

punctured (Fig. 142). Metacoxal lobes moderately

strongly, near suture weaker punctured, not

reaching posterior margin of sternite 4. Sternite

5+6 posteriorly with complete puncture-row,

sternite 7 with a few punctures in apical part.

Setiferous striole on dorsal face of hind tibia about

1/3x tibia-length, longer tibial spur 2/3x length of

first tarsal segment (Fig. 141).

Male: Pro- and mesotarsomeres 1-3 widened

and with sucker hairs on ventral side. Penis and

parameres as in Figs 144-146.

Biology

Specimens were collected in a creek and were

attracted to light.

Distribution (Fig. 162)

Australia: Western Australia,

Territory, Queensland.

Material examined: Western Australia: 1 6, N.

W.A., Kununurra, 22.xii.1991-6.1.1992, R. I.

Northern

92 VAN VONDEL

GLI 136-146. Haliplus stepheni, 136-143, Holotype 2; 144-146, Paratype 3: 136, dorsal view; 137, anten-

na; 138, punctures near elytral base and suture; 139, maxillair palpus; 140, labial palpus; 141, hind tibia; 142,

prosternal and metasternal process; 143, prosternal process in lateral view; 144, left paramere; 145, penis; 146, right

paramere.

AUSTRALIAN HALIPLIDAE 935

Storey (CV); Northern Territory: Holotype @?

(SAMA); 8 paratypes with same data as holotype

(1 do in CW; 4 3, 3 & in QPI); Queensland: 1 2,

N. Qld, 11 km WSW of Petford, 3/4.iv.1988, R. L.

Storey, at light (QPI); 2 ex. Gunshot Ck, 13 km

WNW of Heathlands, 11.43S, 142.26E,

18.11.1992, at light, D. C. F. Rentz (ANIC, CV);

1 2, Qld nr Mt Mulligan, 31.1.1991, Larson &

Halfpap (MUNC).

Haliplus sindus Watts

(Figs 147-152)

Haliplus sindus Watts, 1988: 22. Holotype 2, Qld

Bentinck Is. ‘Ninyilki’ 6 June 1963. P. Aitken,

N. B. Tindale. (SAMA)[examined].

Diagnosis

This species is easy to distinguish from others

in the region by its small size and the serrate

lateral margins of pronotum.

Description

Length 1.7—2.2 mm, width 1.1-1.3 mm. Body

oval, strongly tapering posteriorly, widest in front

of middle (Fig.147).

Head: Yellow to yellow-red, weakly and

sparsely punctured. Antenna (Fig. 148) and palpi

yellow. Last segment of maxillair palpi 1/3x

length of penultimate segment. Last segment of

labial palpi almost as long as penultimate

segment. Distance between eyes 1.4—1.5x width of

one eye.

Pronotum: Yellow to yellow-red, weakly

punctured, in anterior central part more strongly

and densely punctured, base impressed. Lateral

borders slightly convex, anteriorly bent inwards,

lateral margins clearly serrate, not reaching

anterior corner (Fig. 147).

LOS a2) Oa ay ae

148

(9)

b ©

b 0

b) ©

19)

fh °

151

FIGURES 147-152. Haliplus sindus, Holotype 2: 147, dorsal view; 148, antenna; 149, punctures near elytral base

and suture; 150, hind tibia; 151, prosternal and metasternal process; 152, prosternal process in lateral view.

94 VAN VONDEL

FIGURE 153. Distribution of Haliplus oberthuri (dots) and Haliplus sindus (triangle, black: examined; open:

unexamined paratype).

FIGURE 154. Distribution of Haliplus bistriatus.

AUSTRALIAN HALIPLIDAE

co waa

= co 2S

(eae °

FIGURE 156. Distribution of Haliplus ferruginipes. (Dot: examined, circle: not examined type.)

96 VAN VONDEL

FIGURE 158. Distribution of Haliplus gibbus (dots: males; circles: females of H. gibbus or H. fuscatus).

AUSTRALIAN HALIPLIDAE

FIGURE 159. Distribution of Haliplus australis.

FIGURE 160. Distribution of Haliplus testudo (dots: locality known; circle: specific locality in S.A. unknown).

OF,

98 VAN VONDEL

FIGURE 162. Distribution of Haliplus signatipennis (dots) and Haliplus stepheni (triangles).

AUSTRALIAN HALIPLIDAE

FIGURE 163. Distribution of Haliplus alastairi.

FIGURE 164. Distribution of Haliplus timmsi.

100 VAN VONDEL

Elytra: Yellow to yellow-red. Primary punctures

weak and sparse, row 1+2 not as strong as row 3-

7, base of row 5 ending in transverse strong

impression, about 25 punctures in row lI.

Secondary punctures scattered and almost

restricted to odd intervals, sutural row almost as

strong, but not as dense as primary row | (Fig.

149). Punctures not darkened. Interval between

puncture-row 8 and 9 slightly elevated in anterior

part. Lateral sides margined and clearly serrate

near shoulders and near apex.

Underside: Yellow to yellow-red, elytral

epipleura yellow to yellow-red, legs yellow-red to

yellow-brown, slightly darkened near coxae.

Elytral epipleura reaching end of sternite 5. Pros-

ternum completely margined anteriorly, not or

hardly punctured. Prosternal process parallel-

sided, slightly wider posteriorly, anterior edge

margined, slightly impressed in posterior half,

weakly punctured, stronger punctures in

impression (Figs 151, 152). Metasternal process

hardly wider than prosternal process, clearly

impressed anteriorly, moderately strongly

punctured (Fig. 151). Metacoxal lobes reaching to

end of sternite 4, sparsely punctured, punctures

weaker towards suture. Sternite 5+6 with

complete puncture-rows, sternite 7 completely, but

sparsely punctured. Setiferous striole on dorsal

face of hind tibia short, with about 3 punctures,

both tibial spurs about 2/3x length of first tarsal

segment (Fig. 150).

Male: unknown

Distribution (Fig. 153)

Only known from type localities in Queensland:

Bentinck Island (holotype) and Homehill

(unexamined paratype).

Material examined: Only holotype °.

ACKNOWLEDGMENTS

I am very grateful to Dr C. H. S. Watts (Adelaide,

SAMA) for his very valuable help and to Dr D. J. Larson

(St. John’s, MUNC) and Dr O. Bistrom (Helsinki,

UZMB) for critical comments.

I wish to express my sincere thanks to the following

persons for placing material or information at my

disposal: Dr L. Baert (Brussels, ISNB), Mr M. J. D.

Brendell (London, BMNH), Dr S. P. Cover (Cambridge,

MCZC), Dr K. Desender (Brussels, ISNB), Dr M.

Hansen (Copenhagen, ZMUC), Dr S. J. Hine (London,

BMNH), Dr M. Holmen (Copenhagen, ZMUC), Dr M.

Jach (Vienna, NHMV), Dr J. Krikken (Leiden, RMNH),

Dr G. B. Monteith (South Brisbane, QMBA); Mrs H.

Perrin (Paris, MNHN), Dr R. Poggi (Genoa, MCSN); Dr

G. A. Samuelson (Honolulu, BPBM), Ms M. Schneider

(Brisbane, UQIC); Dr A. Smetana (Ottawa, CNCI), Mr

R. I. Storey (Mareeba, QPI); Dr R. zur Strassen

(Frankfurt am Main, SMFD), Mr K. Walker (Abbots-

ford, MVMA); Mr T. A. Weir (Canberra, ANIC).

The Uyttenboogaart-Eliasen Foundation is

acknowledged for financial support.

REFERENCES

BEUTEL, R. G., & S. RUHNAU, 1990. Phylogenetic

analysis of the genera of Haliplidae (Coleoptera)

based on characters of adults. Aquatic Insects 12: 1—

CLARK, H., 1862. Catalogue of the Dytiscidae and

Gyrinidae of Australasia with descriptions of new

species. Journal of Entomology 1: 399-421.

FAUVEL, A., 1883. Les Coléoptéres de la Nouvelle-

Calédonie et dépendances avec descriptions, notes et

synonymies nouvelles, Suite. Revue d’Entomologie 2:

335-360.

GUIGNOT, F., 1928. Notes sur les Haliplus des groupe

fulvus F. Annales de la Société Entomologique de

France 97: 133-151.

GUIGNOT, F., 1935a. Douziéme note sur les

Hydrocanthares. Bulletin de la Société entomolo-

gique de France 40: 36-40.

GUIGNOT, F., 1935b. Treiziéme note sur les

Hydrocanthares (Col. ). Bulletin de la Société

entomologique de France 40: 164-169.

GUIGNOT, F., 1955. Sur la systématique des Haliplus

(Col. Haliplidae). Mémoires de la Société Royale

d’Entomologie de Belgique 27: 289-296.

LAWRENCE, J. F., T. A. WEIR & J. E. PYKE, 1987.

Haliplidae. Pp. 321-322 in ‘Zoological catalogue of

Australia Vol. 4. Coleoptera: Archostemata,

Myxophaga and Adephaga’. Ed. D.W. Walton.

Australian Government Publishing Service: Canberra.

NETOLITZKY, F., 1911. Die Parameren und das

System der Adephaga (Col. Caraboidea). Deutsche

Entomologische Zeitschrift: 271-283.

REGIMBART, M., 1890-1891. Viaggio di Lamberto

Loria nella Papuasia Orientale IV. Haliplidae, Dytis-

cidae et Gyrinidae. Annali del Museo Civico di Storia

Naturale di Genova Série 2a, Vol. 10: 978-997.

REGIMBART, M., 1899. Révision des Dytiscidae de la

région Indo-Sino-Malaise. Annales de la Société

Entomologique de France 68: 186-367.

VONDEL, B. J. van, 1991. Revision of the palaearctic

species of Haliplus subgenus Liaphlus Guignot

(Coleoptera: Haliplidae). Tijdschrift voor

Entomologie 134: 75-144, Figs 1-312.

AUSTRALIAN HALIPLIDAE 101

WATTS, C. H. S., 1985. A faunal assessment of _WEHNCKE, E., 1883. Neue Halipliden. Deutsche

Australian Hydradephaga. Proceedings of the Aca- Entomologische Zeitschrift 27: 145-146.

demy of Natural Sciences of Philadelphia 137: 22- ZIMMERMANN, A., 1920. Dytiscidae, Haliplidae,

28. Hygrobiidae, Amphizoidae. Coleopterorum Cata-

WATTS, C. H. S., 1988. Revision of Australian logus 71: 1-325.

Haliplidae (Coleoptera). Records of the South Austra~ 7 )4MMERMANN. A.. 1924. Die Halipliden der Welt.

lian Museum 22(1): 21-28. Entomologische Blatter (fiir Biologie und Systematik

WEHNCKE, E., 1880. Neue Haliplus. Stettiner der Kafer) 20(1): 1-16; (2): 65-80; (3): 129-144;

Entomologische Zeitung 75: 72-75. (4): 193-213.

MOLLUSC TYPE SPECIMENS IN THE SOUTH AUSTRALIAN MUSEUM. 6. ADDITIONS

AND CORRECTIONS TO PART 1 (CEPHALOPODA AND SCAPHOPODA) AND PART 3

(POLYPLACOPHORA).

K. L. GOWLETT-HOLMES & W. ZEIDLER

GOWLETT-HOLMES, K. L. & ZEIDLER, W. 1995. Mollusc type specimens in the South

Australian Museum. 6. Additions and corrections to Part 1 (Cephalopoda and Scaphopoda) and

Part 3 (Polyplacophora). Records of the South Australian Museum 28(1): 103-111.

This paper lists the recent additions made to the South Australian Museum’s type collections

of Chitons (Zeidler and Gowlett, 1986) and Cephalopoda (Zeidler and Macphail, 1978). Since

publication of these type lists type material of twelve taxa of Polyplacophora and nine taxa of

Cephalopoda have been added to the Museum’s collections. Further additions to type material

of species listed previously, changes in type status and nomenclature are also noted, reflecting

the importance of the mollusc collection in the South Australian Museum.

K. L. Gowlett-Holmes and W. Zeidler, South Australian Museum, North Terrace, Adelaide,

South Australia 5000. Manuscript received 18 October 1994.

There have been several important additions to

the collections since the first publication in the

series (Zeidler & Macphail, 1978) cataloguing the

mollusc type specimens in the South Australian

Museum (SAM). One of the most significant

additions since then has been some historical

cephalopod types described by S. Stillman Berry

in 1921 and 1932, which until recently had been

thought lost. These were found amongst Berry’s

vast collection, most of which was bequeathed to

the Santa Barbara Museum of Natural History

upon his death on 9 April 1984 (Hochberg, 1985),

except for the cephalopod collection, which was

left to the Smithsonian. Dr Clyde Roper

subsequently curated Berry’s cephalopod material

and returned the specimens to the South

Australian Museum in 1986. Two of the types had

been out on loan for 65 years!

Other additions have resulted from subsequent

work on the collections, donations from other

workers and nomenclatural changes, reflecting the

importance of the mollusc collection in the South

Australian Museum. The types of 9 taxa of

cephalopods and 12 taxa of Polyplacophora have

been added to the SAM collections. There have

been changes to the status or additions to the types

of 7 taxa of Polyplacophora listed previously

(Zeidler & Gowlett, 1986). There have been no

additions to the gastropod families catalogued to

date; Conidae (Zeidler, 1985), Marginellidae

(Hewish & Gowlett-Holmes, 1991), Cypraeidae,

Triviidae and Ovulidae (Gowlett-Holmes &

Zeidler, 1993). In the following list species are

arranged alphabetically in families under the

original name at the time of description. In the

Polyplacophora, specimens are dry unless

otherwise indicated, and are listed as ‘entire’ when

the articulated valves and girdle are present or as

‘entire with animal’ when the animal is also

present.

Other abbreviations used in the text are as

follows; AM = Australian Museum, Sydney.

BMNH = The Natural History Museum, London

(formerly British Museum, Natural History).

LACM = Los Angeles County Museum of Natural

History, California. MNHN = Museum National

dHistoire Naturelle, Paris. NMNZ = Te Papa

Tongarewa Museum of New Zealand, Wellington

(formerly National Museum, New Zealand). NUV

= Museum of Victoria, Melbourne (formerly

National Museum of Victoria). QM = Queensland

Museum, Brisbane. RSM = Royal Scottish

Museum, Edinburgh. SBMNH = Santa Barbara

Museum of Natural History, California. SEM =

Scanning Electron Micrograph. USNM = National

Museum of Natural History, Smithsonian

Institution, Washington, DC. WAM = Western

Australian Museum, Perth.

Class POLYPLACOPHORA

Family ABYSSOCHITONIDAE

Genus Ferreiraella Sirenko, 1988

Ferreiraella caribbensis Sirenko, 1988

Zool. Zh. 67(12): 1778-81, figs 1, 2.

104

Paratype: D18762, one specimen with animal in

spirit, with anterior and two median valves

disarticulated, dredged in 6740-6780 m, Cayman

Trench, Caribbean Sea [19°39'N, 76°37'W], RV

‘Ac. Kurchatov’, Stn 1267, collected by L.

Moskalev, 24-5 March, 1973.

Note: Donated to SAM collections by B. I.

Sirenko, St Petersburg, Russia. The holotype and

additional paratypes are in the Academy of

Sciences, St Petersburg, Russia.

Family ACANTHOCHITONIDAE

Genus Acanthochites Risso, 1826

Acanthochites kimberi Torr, 1912

Trans. R. Soc. S. Aust. 36: 167, pl. 6, figs 5a-f.

= Acanthochitona kimberi (Torr, 1912).

Syntype: D17586, entire specimen with animal,

from Aldinga, near Adelaide, South Australia,

collected by W. J. Kimber, date of collection

unknown.

Note: An additional syntype to the series listed by

Zeidler and Gowlett (1986) found recently in the

collections. Specimen with small label “TYPE 59’,

typical of other Torr type specimens. Present status

according to Kaas and Van Belle (1980).

Genus Acanthochiton Gray, 1821 em.

Iredale, 1915

Acanthochiton mayi Ashby, 1922

Trans. R. Soc. S. Aust. 46: 12, pl. 3, figs 1a—b.

= genus uncertain

Syntypes: D12799, five median valves, in 100 fm

(183 m), 7 miles east of Cape Pillar, and from

Schouten Island, Tasmania, collected by W. L.

May, 18 Dec. 1907 (Cape Pillar). D17942, one

median valve, in 60 fm (110 m), off Port Arthur,

Tasmania, collected by W. L. May, date of

collection unknown.

Note: May Collection No. 260A. Type status was

not indicated on the original labels, but there is no

doubt that these are the specimens that Ashby

(1922) examined, and may include the valve he

illustrated. Ashby states that one syntype was

lodged with the Tasmanian Museum, but Turner

and Dartnall (1971) do not list this specimen, and

we have been unable tolocate it there. Kaas and

Van Belle (1980) place this species in the genus

Notoplax, but it does not fit into this genus as

defined by Gowlett-Holmes (1991), and its current

generic status is uncertain.

K. L. GOWLETT-HOLMES & W. ZEIDLER

Genus Acanthochitona Gray, 1821

Acanthochitona saundersi Gowlett-Holmes &

Zeidler, 1987

Trans. R. Soc. S. Aust. 111(2): 111, figs 1, 2.

Holotype: D16699, entire specimen with animal,

from on edge of granite slope, under sand, in 8 m,

in cove off northwest point of East Franklin

Island, Nuyts Archipelago, South Australia,

collected by K. L. Gowlett, 20 July, 1983.

Paratypes: D16698, one specimen in spirit,

disarticulated, valves and SEM stub of radula in

dry collection, from on granite ledge, under sand,

in 6 m, on inside of reef off southwest side of East

Franklin Island, Nuyts Archipelago, South

Australia, collected by K. L. Gowlett, 18 July,

1983. D17441, two entire specimens with

animals, from on smooth rock under sand, in 12

m, on Far West Bottom, Tiparra Reef, Spencer

Gulf, South Australia, collected by K. L. Gowlett,

13 May, 1982. D17475, one entire specimen with

animal in spirit, from on granite fragment in sand

pocket on reef, in 7 m, off Point Gilbert, Port

Moorowie, Waterloo Bay, Yorke Peninsula, South

Australia, collected by N. J. C. Holmes, 29 March,

1986.

Genus Notoplax Adams, 1861

Notoplax lancemilnei Gowlett-Holmes, 1988

Trans. R. Soc. S. Aust. 112(4): 169-71, figs 1-3.

Paratype: D18436, entire specimen with animal in

spirit, trawled by F. R. V. ‘Kapala’ in 439 m, off

Wollongong, New South Wales [34°21-19'S,

151°23-25'E], collected by K. Graham, 13

December, 1978.

Note: Holotype in AM (C151130).

Family CHITONIDAE

Genus Chiton Linnaeus, 1758

Chiton (Rhyssoplax) kimberi Ashby, 1928

Trans. R. Soc. S. Aust. 52: 170, pl. 12, figs 10-12.

= Rhyssoplax kimberi (Ashby, 1928)

Holotype: D12392, eight disarticulated valves,

fragments of girdle, plus radula mounted on slide,

from Capricorn Group, Queensland, collected by

W. J. Kimber, date of collection unknown.

Note: This species was listed by Zeidler and

Gowlett (1986). The three median valves listed

there as presumed lost have now been located.

MOLLUSC TYPES 6: ADDITIONS AND CORRECTIONS

The two lots have been combined and registered

under the one number, as the pieces all belong to

the one specimen, the unique holotype.

This species was listed by Kaas and Van Belle

(1980) as Chiton (Rhyssoplax) kimberi, but

Rhyssoplax is now considered a valid genus

(Bullock, 1988).

Chiton tulipa alfredensis Ashby, 1928

Proc. Malac. Soc. 18(2): 87, pl. 8, figs 19-21.

= Rhyssoplax tulipa (Ashby, 1928)

Holotype: D10983, two slides, each with a piece

of what appears to be a single radula, from Port

Alfred, South Africa, collected by W. H. Turton,

date of collection unknown.

Note: This material most likely belongs to the

holotype listed by Zeidler and Gowlett (1986).

This species was listed by Kaas and Van Belle

(1980) as Chiton (Rhyssoplax) tulipa, but

Rhyssoplax is now considered a valid genus

(Bullock, 1988).

Family ISCHNOCHITONIDAE

Genus Ischnochiton Gray, 1847

Ischnochiton (Lepidozona) asthenes Berry, 1919

Lorquinia 2(6): 47.

= Callistochiton asthenes (Berry, 1919)

Paralectotype: D10404, entire specimen, from

White’s Point, Los Angeles County, California,

United States of America, collected by A. G.

Smith, 14-18 July, 1916.

Note: This specimen was listed by Zeidler and

Gowlett (1986) as a paratype, but as the primary

type selection was made in a later paper (Berry,

1919b) than the original description, it is a

lectotype, and the specimen in SAM is a

paralectotype. Lectotype (34389) and additional

paralectotypes (34390-2) in SBMNH.

Ischnochiton crebristriatus Cochran, 1988

Proc. R. Soc. Vic. 100(1): 1-7, figs 1-4, tables 1,

Holotype: D18405, entire specimen with animal,

from on rocks and pebbles under sand, in 8 m,

inside reef, southwest side of East Franklin Island,

Franklin Islands, Nuyts Archipelago, South

Australia, collected by K. L. Gowlett, 13 April,

1983.

Paratypes: D11774, one entire specimen with

animal (crushed), from Port Willunga, near

Adelaide, South Australia, collected by E. Ashby,

105

date of collection unknown. D14482, one entire

specimen with animal, from Arno Bay, Eyre

Peninsula, South Australia, collected by B. J.

Weeding, date of collection unknown. D16568,

three entire specimens with animal, from on

granite rocks under medium to coarse sand, in 10

m, big break north of islands, Franklin Islands,

Nuyts Archipelago, South Australia, collected by

K. L. Gowlett, 21 July, 1983. D16569, one entire

specimen with animal, from on rock under sand,

in 15 m, rise inshore of big break north of islands,

Franklin Islands, Nuyts Archipelago, South

Australia, collected by K. L. Gowlett, 24 February,

1983. D16570, two specimens, one entire with

animal, one entire specimen and radula, from on

smooth granite and pebbles under sand, in 6-11

m, between West Franklin Island and Seal Island,

Franklin Islands, Nuyts Archipelago, South

Australia, collected by K. L. Gowlett, 25 February,

1983. D16571, one specimen in spirit with

posterior and two median valves disarticulated,

radula and part of girdle on SEM stubs in dry

collection, with same locality data as holotype,

collected by P. Aerfeldt. D16572, three entire

specimens with animals, with same collection

data as holotype. D16573, one specimen with

animal, with anterior, one median and posterior

valves disarticulated, from edge of granite slope

under sand, in 8 m, in cove, northwest point of

East Franklin Island, Franklin Islands, Nuyts

Archipelago, South Australia, collected by K. L.

Gowlett, 20 July, 1983. D16574, two entire

specimens with animals, from on granite rock

under sand, in 6 m, inside cove west end of Seal

Island, Franklin Islands, Nuyts Archipelago,

South Australia, collected by K. L. Gowlett, 17

July, 1983. D18406, one entire specimen with

animal, from near Port Hughes, Moonta Bay,

Yorke Peninsula, South Australia, collected by B.

J. Weeding, Jan. 1932.

Ischnochiton hewitti Ashby, 1931

Ann. S. Afr. Mus. 30(1): 33-4, pl. 5, figs 50-53.

= Ischnochiton bergoti (Velain, 1877)

Syntype: D10997, entire specimen, and radula

mounted on slide, from Table Bay, Cape Town,

South Africa, collector and date of collection

unknown.

Note: Zeidler and Gowlett (1986) list this

specimen as the holotype, but noted that it was

probably only a paratype. However, as Ashby

(1931) did not designate a type and used a series

of specimens in the description, this specimen is a

syntype. Current status according to Kaas and Van

Belle (1990).

106

Ischnochiton (Chartoplax) nubilus Cochran,

1993

Proc. R. Soc. Vic. 105(1): 51-4, figs 1, 2.

Holotype: D11728, entire specimen with animal,

dredged from Gulf St Vincent, South Australia,

collected by J. C. Verco, date of collection

unknown.

Paratypes: D15127, one entire specimen, and

radula on slide, dredged in 20 m, from off

Brighton, near Adelaide, South Australia,

collected by M. J. Tilbrook, 8 June, 1968.

D17592, one entire specimen with animal in

spirit, from on pebble in sand, in Posidonia

seagrass, in 3 m, 150-200 m offshore of northwest

point of Reevesby Island, Sir Joseph Banks Group,

South Australia, collector W. Zeidler, 13 January,

1984. D17593, one specimen with animal in

spirit, anterior, two median and posterior valves

disarticulated, one valve, radula and part of girdle

on SEM stubs in dry collection, from on pebbles

in sand, in sparse Posidonia seagrass, in 3-6 m,

off northwest point of Marum Island, Sir Joseph

Banks Group, South Australia, collected by W.

Zeidler and K. L. Gowlett-Holmes, 23 January,

1986.

Note: The holotype (D11728) is also the

paralectotype of Ischnochiton (Stenochiton)

pallens Ashby, 1900, also listed here. The slide of

paratype D15127 contains two radulae, one of

which is from the paratype, the other is from a

non-type specimen (D14928).

Ischnochiton (Stenochiton) pallens Ashby, 1900

Trans. R. Soc. S. Aust. 24: 86, pl. 1, figs 1a-e, g.

= Ischnochiton (Chartoplax) purus Sykes, 1896

Lectotype: D978, five disarticulated median valves

and anterior valve, from Gulf St Vincent, South

Australia, dredged by J. C. Verco, date of

collection unknown.

Paralectotype: D11728, entire specimen with

animal, with same collection data as lectotype.

Note: Zeidler and Gowlett (1986) incorrectly listed

these specimens as holotype and paratype,

following Cotton and Godfrey (1940) and Cotton

(1964). However, Ashby (1900) did not designate

a type for this species, although he labelled the

specimens ‘type’ and ‘co-type’. Cotton & Godfrey

(1940) by listing D978 as a holotype, actually

designated the lectotype by inference of holotype

(ICZN Article 74(a & b)).

The paralectotype (D11728) is also the holotype of

Ischnochiton (Chartoplax) nubilus Cochran,

1993, also listed here.

K. L.GOWLETT-HOLMES & W. ZEIDLER

Genus Juvenichiton Sirenko, 1975

Juvenichiton komandorensis Sirenko, 1975

Zool. Zh. 54(10): 1445-7, fig. 2.

Paratypes: D18761, two entire specimens with

animals, from on Constantinea subulifera (a red

alga), in 15 m, Cape Fedoskin, Bering Island,

Commander Islands, Bering Sea, Russia, collected

by B. I. Sirenko, 23 September, 1973.

Note: Donated to SAM collections by B. I.

Sirenko, St Petersburg, Russia. The holotype and

additional paratypes are in the Academy of

Sciences, St Petersburg, Russia.

Genus Particulazona Kaas, 1993

Particulazona milnei Kaas, 1993

Basteria 57: 127-30, figs 1-14.

Holotype: D18930, disarticulated and dissected,

most valves and girdle fragments in vials, girdle

and valve fragments on SEM stubs, girdle

fragments and radula on slides, from on mangrove

roots, Frances Bay, Darwin, Northern Territory,

collected by M. J. Tilbrook, 22 July, 1954.

Note: Type unique.

Genus Stenochiton Adams and Angas, 1864

Stenochiton pilsbryanus dilatus Iredale & Hull,

1924

Aust. Zool. 3: 287, pl. 36, fig. 11.

= Stenochiton pilsbryanus (Bednall, 1897)

Syntype: D18766, entire specimen with animal,

from on blades of seagrass, Lucky Bay, 25 miles

east of Esperance, Western Australia, collected by

A. F. B. Hull, date of collection unknown.

Note: From the K. L. Milne Collection. Originally

labelled as a paratype, but Iredale and Hull did

not designate a holotype, so all type specimens

constitute a syntype series. An additional syntype

is in WAM (11658). Present status according to

Kaas and Van Belle (1994).

Family LEPIDOPLEURIDAE

Genus Lepidopleurus Risso, 1826

Lepidopleurus badius Hedley & Hull, 1909

Rec. Aust. Mus. 7: 260, pl. 73, figs 1, 2.

= Leptochiton badius (Hedley & Hull, 1909)

Paratype: D12532, one entire specimen with

MOLLUSC TYPES 6: ADDITIONS AND CORRECTIONS 107

animal, from Long Reef, near Narrabeen, New

South Wales, collector and date of collection

unknown.

Note: The other lot (D10668) listed by Zeidler and

Gowlett (1986) as possible paratypes of this

species, has since been confirmed not to represent

type material.

Family MOPALIIDAE

Genus Placiphorina Kaas and Van Belle,

1994

Placiphorina gowlettholmesae Kaas & Van

Belle, 1994

‘Monograph of Living Chitons 5: 341-3, fig. 138,

map 39.

Holotype: D18837, entire specimen with animal

in spirit, from under stones embedded in silty

black sand, in 6-15 ft (2-4.5 m), Batu Belah

(approximately 4 km west of Amed), Bali,

Indonesia, collected by K. L. Gowlett-Holmes and

W. Runti, 18 September, 1990.

Paratypes: D18838, one disarticulated specimen,

and 9 entire specimens with animals in spirit, from

under rocks embedded in silty black sand, in 6-10

ft (2-3m), at site of MV ‘Liberty’ shipwreck,

Tulamben, Bali, Indonesia, collected by K. L.

Gowlett-Holmes, 8 September, 1990.

Note: Additional paratypes with same data as

D18838 are in the Nationaal Natuurhistorisch

Museum, Leiden, NetherlandsS(RMNH 9357). The

slides of the girdle and radula mentioned by Kaas

and Van Belle (1994) have not been deposited in

SAM.

Family SCHIZOCHITONIDAE

Genus Schizochiton Gray, 1847

Schizochiton polyops Iredale & Hull, 1926.

Aust. Zool. 4: 271, pl. 38, figs 16, 17, 19-21.

= Schizochiton incisus (Sowerby, 1841).

Syntypes: D18767, two entire specimens, from

Howick Island, Queensland, collected by A. F. B.

Hull, date of collection unknown.

Note: From the K. L. Milne Collection. Originally

labelled as paratypes, but Iredale and Hull did not

designate a holotype, so all type specimens

constitute a syntype series. Additional syntypes

should exist in the AM and NMV, but none are

listed by Smith and Robertson (1970) or Boyd and

Phillips (1985), or in the computer printout of

Polyplacophoran types in AM. Present status

according to Kaas and Van Belle (1980).

Family XYLOCHITONIDAE

Genus Xylochiton Gowlett-Holmes and

Jones, 1992

Xylochiton xylophagus Gowlett-Holmes & Jones,

1992

J. Malac. Soc. Aust. 13: 38-43, figs 1-5.

Paratypes: D18770, three specimens, two entire

with animal, one specimen disarticulated, in spirit,

radula on SEM stub in dry collection, dredged in

1075-1100 m, on large waterlogged log of

Coriaria arborea (Tree Tutu), off White Island

[37°23.7'S, 171°39.5-36.6'E], east of North

Island, New Zealand, collected by the U.S.S.R.

FV ‘Kalinovo’, Stn BS 924 (K01/019/81), 23

November, 1981.

Note: The holotype (M.100855) and additional

paratypes (M.74996, M.84251, M.86822,

M.92446) are in NMNZ. Additional paratypes are

in RSM (NMSZ 1991055), USNM (860288),

BMNH (1991145), AM (C168568), MNHN and

LACM (2280).

Class CEPHALOPODA

Family IDIOSEPIIDAE

Genus Idiosepius Steenstrup, 1881

Idiosepius notoides Berry, 1921

Rec. S. Aust. Mus. 1(4): 361-2, chart 11, fig. 67.

Holotype: D17495, complete male specimen in

spirit, from Goolwa, South Australia, collected by

A. Zeitz, date of collection unknown (Berry

Collection No. SSB 719).

Note: Paratype female in USNM (SSB 720).

Family SEPIOLIDAE

Genus Neorossia Boletzky, 1971

Neorossia leptodons Reid, 1991

Bull. Mar. Sci. 49(3): 797-806, figs 14d, 24-28,

tables 23,24, appendix Id.

Paratypes: D18632, one complete female in spirit,

trawled in 130 m, approx. 80 nautical miles

south-southwest of St Francis Island, Nuyts

108 K. L. GOWLETT-HOLMES & W. ZEIDLER

Archipelago, South Australia, [33°42'S, 132°25'E]

FV ‘Merindah Pearl’, collected by M. Jubb, 12

August, 1988. D18724, one complete female in

spirit, trawled in 1000 m, approx. 120 nautical

miles southwest of Cape Adieu, Great Australian

Bight, South Australia, [33°58'S, 131°22'E], FV

‘Saxon Progress’, collected by D. Wheenan, Nov.

1989. D18725, one complete male in spirit, same

collection data as D18724. D18726, three

complete males in spirit, same collection data as

D18724.

Note: Holotype (F57504) and additional paratypes

are held in NMV, and additional paratypes are

held in AM.

Family SEPIADARIIDAE

Genus Sepiadarium Steenstrup, 1881

Sepiadarium austrinum Berry, 1921

Rec. S. Aust. Mus. 1(4): 354-5, chart 10.

Holotype: D17493, complete male specimen in

spirit, from Gulf St Vincent, South Australia,

collected by A. Zeitz, September, 1885 (Berry

Collection No. SSB 716).

Paratype: D17494, complete female specimen in

spirit, same collection data as holotype (Berry

Collection No. SSB 718).

Note: Paratype in USNM (SSB 717).

Sepiadarium nipponianum Berry, 1932

Philippine J. Sci. 47(1): 42-6, pl. 1, figs 2-5.

Paratype: D17496, complete specimen in spirit,

from Sagami Bay, Japan, collector M. Sasaki, 10

April, 1918 (Berry Collection No. SSB 725).

Note: Holotype in USNM (SSB 724).

Family OCTOPODIDAE

Genus Eledone Leach, 1817

Eledone palari Lu & Stranks, 1991

Bull. Mar. Sci. 49(1-2): 73-85, figs 1-6.

Paratypes: D18721, one male and one female, both

complete in spirit, trawled in 157 m, east of North

Stradbroke Island, Queensland, [27°35'S,

153°50'E], FV ‘Iron Humphrey’, collected by M.

Potter, 2 July, 1981, (ex NMV F57531).

Note: Holotype (F57849) and additional paratypes

in NMV, additional paratypes in AM, WAM and

QM.

Genus Octopus Lamarck, 1798

Octopus berrima Stranks & Norman, 1993

Mem. Mus. Vic. 53(2): 355-61, figs 3, 6-11.

Paratype: D18775, dissected male specimen in

spirit, dredged off Mordialloc, Port Phillip Bay,

Victoria, [38°02'S, 145°05'E], FV ‘A. B. Hunter

IP’, 25 September, 1984 (ex NMV F52510).

Note: Holotype (F67132) and additional paratypes

in NMV and an additional paratype in AM.

Octopus bunurong Stranks, 1990

Mem. Mus. Vic. 50(2): 462-4, figs 3a-f.

Paratypes: D17983, one complete male specimen

in spirit, from reef, rubble, sand and Posidonia

seagrass, in 6 m, Partney Shoal, west of Partney

Island, Sir Joseph Banks Group, South Australia,

collected by W. Zeidler and N. J. C. Holmes, 21

January, 1986. D17986, one complete male

specimen in spirit, from 50 m offshore of Marino

Rocks, near Adelaide, South Australia, collected

by R. Browne, 28 January, 1982.

Note: Holotype (F53223) and additional paratypes

in NMV.

Octopus kaurna Stranks, 1990

Mem. Mus. Vic. 50(2): 460-2, figs 2a-f.

Paratypes: D13283, one complete immature male

specimen in spirit, from off Brighton, near

Adelaide, South Australia, collected by W. G.

Hollis, September, 1937. D16195, one female

specimen in spirit, and radula on slide, from off

Glenelg, near Adelaide, South Australia, collected

by A. E. Robb, 29 March, 1949.

Note: Holotype (F24494) and additional paratypes

in NMV.

Octopus warringa Stranks, 1990

Mem. Mus. Vic. 50(2): 457-60, figs 1a-f.

Paratype: D15219, complete female specimen in

spirit, from off Maria Island, Tasmania, [42°40'S,

148°28'E], RV ‘Discovery’, BANZARE Stn 113,

23 March, 1931.

Note: This specimen was originally identified as

Robsonella australis (Hoyle, 1885) by Dell

(1959). Holotype (F57444) and additional

paratypes in NMV, and an additional paratype in

AM.

ACKNOWLEDGMENTS

We are most grateful to Dr. Clyde Roper, USNM for

ensuring the safe return of the cephalopods described by

MOLLUSC TYPES 6: ADDITIONS AND CORRECTIONS

S. S. Berry and to Dr. Boris Sirenko, Academy of

Sciences, St. Petersburg, Russia for donating chiton

paratypes.

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REVISION OF THE AUSTRALASIAN GENERA AGRAPHYDRUS REGIMBART,

CHASMOGENUS SHARP AND HELOCHARES MULSANT

(COLEOPTERA: HYDROPHILIDAE)

C.H.S. WATTS

WATTS, C. H. S. 1995. Revision of the Australasian genera Agraphydrus Régimbart,

Chasmogenus Sharp and Helochares Mulsant (Coleoptera: Hydrophilidae). Records of the

South Australian Museum 28(1) 113-130.

The Australian and New Guinean members of the Hydrophilid genera Agraphydrus,

Chasmogenus and Helochares are revised and redescribed. A key to the genera and species is

given. Fourteen species are recognised of which six are described as new: H. (Hydrobaticus)

marreensis; H. (Hydrobaticus) percyi; H. (Hydrobaticus) anthonyae; H. (Hydrobaticus)

thurmerae; H. (Hydrobaticus) loweryae; and H. (Hydrobaticus) dalhuntyi. H. (Hydrobaticus)

australis Blackburn is synonymised with H. (Hydrobaticus) tristis Macleay. H. (Hydrobaticus)

luridus W. Macleay is resurrected from synonymy with H. tristis W. Macleay.

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

Manuscript received, 25 May 1994.

The three genera under discussion are small

aquatic Hydrophilidae belonging to the tribe

Hydrophilini, subtribe Acidocerina, which are

characterised by lack of a sternal keel, head not

strongly deflexed, middle and hind tibiae without

swimming hairs, maxillary palpi larger than

antennae and with pseudobasal segment, straight

or curved, with convexity on inside, and the

mesosternum without projecting longitudinal

laminae. Until recently they were all included in

the genus Helochares Mulsant, 1844 but were

reinstated as full genera in the extensive revision

of the Hydrophiloids by Hansen (1991).

As well as Australia and New Guinea, species

of Helochares are widespread in both the old and

new world, Chasmogenus Sharp, 1882 in

Neotropical, Oriential and Palearctic regions and

Agraphydrus Régimbart, 1903 in East Africa,

Southern Asia and Japan.

D’Orchymont (1925, 1939, 1943) revised

Chasmogenus and Helochares, including all the

Australian and New Guinean species then known.

The genera and various subgenera of the group are

readily recognised but within them the species are

difficult to distinguish. D’Orchymont noted that in

Chasmogenus the characters of the aedeagus are

the only reliable means of separating species. I

have found this to be true for several Helochares

species also.

Specimens of AH. (Hel.) foveicollis

(Montrouzier, 1860) and C. nitescens Fauvel,

1883 are often mistaken for species of Enochrus

Thomson, 1859 but can be separated by the

maxillary palps having the pseudobasal segment

bent inwards in Helochares and Chasmogenus

rather than outwards as in Enochrus. Helochares

and Chasmogenus also lack the simple

protuberance in the ventral midline in front of the

mesocoxae present in Agraphydrus.

Within Helochares, eleven of the twelve

Australian species belong to the subgenus

Hydrobaticus W. MacLeay, 1871, which is

readily recognised by its strongly punctate surface

with distinct striae or rows of punctures on elytra.

All but one species (H. anchoralis Sharp, 1890)

are endemic to the region and only two are known

from both New Guinea and Australia. In contrast,

the subgenus Helochares lacks strong sculpture

and is represented, in Australasia, by a single

species which is also widespread in South-east

Asia and the Pacific Islands. The single

Australasian species of Agraphydrus and

Chasmogenus are tropical with a wide distribution

in South-east Asia as well as Australasia.

Most species in the group are tropical but one

H. (Hyd.) tristis Macleay, 1871, is common in

southern Australia and another H. (Hyd.)

tenuistriatus Régimbart, 1908 is found only in the

south-west of Western Australia.

All species are inhabitants of shallow still water

or slow moving well-vegetated streams. Females

of H. (Hyd.) tristis are often taken with a large

white coloured eggmass attached to the ventral

surface of the abdomen. The larvae of H. tristis

114 C.H.S. WATTS

and C. nitescens have been described by Anderson

(1976) but I know of no studies of the ecology of

any Australasian species.

Hansen (1991) provides a comprehensive

taxonomic discussion of these genera, including

detailed generic description and phylogenetic

analysis.

The collections from which specimens were

examined are listed under the following

abbreviations:

AM Australian Museum, Sydney

ANIC Australian National Insect Collection

BELG Institut royal des sciences Naturelles de

Belgique, Bruxelles

BM(NH) Natural History Museum, London

CW Private collection of author

FIELD Field Museum of Natural History,

Chicago

MNHN Museum National d’Histoire Naturelle,

Paris

NMV Museum of Victoria

NTM Northern Territory Museum

QDPIM Queensland Department of Primary

Industries, Marreba

QM Queensland Museum, Brisbane

SAMA — South Australian Museum, Adelaide

UQIC University of Queensland Insect

Collection, Brisbane

WAM _ Western Australian Museum, Perth

SYSTEMATICS

The genera Agraphydrus, Chasmogenus and

Helochares can be separated from other

Hydrophilids by the following characters: Head

not strongly deflexed; scutellum not, or not much

longer than its basal width; meso- and metatibia

without swimming hairs. Antennae with eight or

nine segments, maxillary palpi elongate, longer

than antennae, last segment shorter than

penultimate. Mesosternum without projecting

carina in midline. Meso- and metasterna without a

continuous keel. Elytra not strongly carinate.

Curved basal joint of maxillary palpus with

convexity at front.

Generic and subgeneric diagnoses are given in

the key. More extensive generic diagnoses are

given by Hansen (1991).

Key To AUSTRALASIAN SPECIES OF AGRAPHYDRUS

REGIMBART, CHASMOGENUS SHARP AND HELOCHARES

MULSANT

Small size (< 3 mm); head predominantly

black; pronotum and elytra yellow-brown;

elytra sparsely punctured, punctures not in

stria. Male genitalia relatively simple;

aedeagus divided into upper and lower

SECHOnS a(Higse Seca) ae ee

Agraphydrus coomani d’Orchymont

— Not with above combination of characters

Black. Elytral striae virtually lacking, at

most reduced to a few lines of serial

PUNCLUNCS os cesacan seria iene ier cercn cere 3

— Predominately brown. Each elytron with

10 complete striae. Helochares, subgenus

18 OWED Rex coe ocescoccosicbinseoentovecreras! 4

Smaller (< 5.0 mm). Sutural stria strongly

impressed in at least apical half of elytra.

Dorsal surface weakly punctured,

punctures on head smaller than eye facets,

subobsolete on scutellum ............ccccceeee

— Larger (> 6.0 mm). No sutural stria.

Dorsal surface with well marked

punctures, those on head larger than eye

facets. Scutellum with sharply impressed

PUNGC HUES atte oens ees Helochares

sensu stricto H. foveicollis (Montrouzier)

Punctures on pronotum of uniform size

except for small punctures close to front

nok beem tay (denies. TUS) esanseonpccosacebecaeccneecoo: 8

— Punctures on pronotum of two sizes (Fig.

Numerous large punctures on front of

head, particularly laterally, not greatly

different in size and density to those on

TES Of CAC sere anrence nase nn eee 6

— Punctures on centre of head small, with

only a few scattered larger ones, larger and

more numerous laterally and on back half

of head (Fig. 17). Large and small

punctures on pronotum and in elytral

interstriae very different in size (Fig. 19)

ee: H. clypeatus Blackburn

Strial punctures close, often confluent and

tending to form grooves, interstrial areas

raised towards apex accentuating grooves

(Fig. 23). (Punctures on elytron margin

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES 115

strong at front grading to weak apically.)

Mee gemiiaia G8 WO TUE, Sossssoomcrscoscoue:

RP EA eR at H. luridus Blackburn

— Strial punctures for most part not

confluent and not forming grooves except

toward apex to slight degree in some

GDACHIMNEMNS (FZ, YD) s.coordaorsasnrssassassevoroos 7

Male genitalia as in Fig. 4. Most of the

smaller interstrial punctures more than

diameter of larger ones

Sehanobactoetonacrs H. tenuistriatus Régimbart

— Male genitalia as in Fig. 3. Most of the

smaller interstrial punctures less than '/,

diameter of the larger ones (Fig. 20)

bse Mer awa nak ge Ba eee Se H. tristis Macleay

Punctures on head smaller than size of eye

facets. (Serial punctures relatively small,

confluent, forming grooves. Body size

5.5-6.2 mm.) Flanges of elytra with large

punctures much larger than those in

interstriae. Only known from New Guinea

Le an aren H. anchoralis Sharp

— Punctures on head larger than eye facets

(except in H. anthonyae, a species with

serial punctures on elytral disc not

confluent). Punctures on flanges of elytra,

other than in H. tatei, a much smaller

species (body size 3.4-4.8 mm), usually

smaller than those in adjacent interstriae..9

Interstrial punctures at base of elytra (and

often over rest of elytra as well) large,

equal in size to the serial punctures which

AIS USO WEAITVOSY TEBE rcorscrososooncenononneres

HENGE Sol at chaceress H. marreensis sp. nov.

— Interstrial punctures on elytra smaller than

those in elytral stria (Fig. 21). Punctures

in elytral stria also tend to be small (large

i tlae/ 2 Ls eon (211) Oe ar ae Se celia Dili a 10

10 — Punctures on head and pronotum very

large, most about a quarter puncture

width apart or less (Fig. 18). Number of

punctures along midline of pronotum

more than 30. Flattish species with elytra

broadly and weakly flanged, somewhat

truncate apically. Body length relatively

large (> 4.8 mm) ........ H. percyi sp. nov.

— Punctures on head and pronotum small

to medium, many separated by half

puncture width or more (except in some

tatei). Punctures on pronotum larger

laterally. Number of punctures along

midline of pronotum 20-40............... 11

11 — Number of punctures along midline of

pronotum around 25. Punctures in elytral

striae as large or larger than those on

pronotum, interstrial punctures tend to

increase in size laterally even prior to

lateral fringe. Interstrial punctures

between stria 8 and 9 usually in single

line. Body length relatively small (<4.8

IVD) aaoanete ocaetriscsccccoe, ence Cape orerens H. tatei

— Number of punctures along midline of

pronotum around 35. Other characters

WISWAIIKY MOL BS ALYONTE .ccccoscooconasposcoundabo 1)

Strial punctures in inner few rows

diminish in size down apical declivity.

(Distance between stria apically greater

than 2 puncture widths apart. Surface of

elytra towards apex has a very flat

appearance)........ H. anthonyae sp. nov.

— Strial punctures in apical quarter, apart

from last few, do not diminish in size

and may even increase in Size........... 13

13. — Lateral stria with 30 or fewer punctures.

(Strial punctures relatively large.) ........

Se tn re Hi. thurmerae sp. nov.

— Lateral stria with more than 30 punctures

trees ae ett reset H. loweryae sp. nov.

Ey eee ee or H. dalhuntyi sp. nov.

Agraphydrus Régimbart

Type species: Agraphydrus punctatellus

Régimbart, 1903, Madagascar. Designation by

monotypy.

Agraphydrus coomani (d’Orchymont)

Helochares (Agraphydrus) coomani

d’Orchymont, 1927

Description (number examined 51)

Length 1.4—2.6 mm. Narrowly oval, head broad

in front giving it a blunt-nosed appearance.

Yellow-brown. Head black except laterally in

front of eyes, disc of pronotum and elytra variably

darker, tips of maxillary palpi black, underside

dark brown except for yellow-brown appendages.

Front of head broadly and weakly concave,

sparsely covered by weakly impressed small

punctures. Pronotum and elytra weakly reticulate,

sparsely covered with small weakly impressed

punctures, those on elytra more strongly impressed

laterally. A number of barely traceable lines of

serial punctures on each elytron. Maxillary palpi

116 C.H.S. WATTS

relatively short and stout, apical segments a little

larger than penultimate, basal segments a little

larger than apical. Rugose portions of femur

covering all but about apical quarter. Tibia with

several rows of strong spines. Metacoxae and

abdominal segments quite densely covered by well

marked punctures, much stronger and denser than

those on dorsal surface. Apical sternite weakly

notched apically. Midline of metacoxae shining

apically. Midline of metasternum very weakly and

broadly ridged.

Male: Protarsi and claws a little stouter than in

female; claws on protarsi more strongly recurved

than in female. Aedeagus as in Figs 13 and 14.

Types

Holotype: Not located in d’Orchymont

collection BELG. Type locality, Lactho near Hoa

Binh, Tonkin, Vietnam.

Distribution (Australasian only)

Northern Territory

Cooper Creek, ANIC; 14 km NW Cape

Crawford, ANIC; McArthur River, ANIC; Pine

Creek, CW; Yuendumu, CW.

Queensland

16°28'S, 144°46'E, ANIC; Bushy Creek,

Julatten, ANIC; Charters Towers, CW; 75 km

Cooktown, ANIC; Mary Creek, ANIC; Mossman,

Mt Lewis Road, ANIC; Mulgrave, ANIC.

New South Wales

Armadale, CW; Cabbage Tree Creek, ANIC.

Western Australia

Millstream, ANIC; 5 km SE Millstream, ANIC.

Papua New Guinea

Kokoda, BELG

Remarks

This small species is relatively common in

collections from northern Australia. These

specimens agree well with d’Orchymont’s detailed

description and with specimens, I take to be of

this species, from Lenggong Malaya Peninsula in

SAMA. There is a male specimen in BELG

mounted on a card with genitalia extruded labelled

as “Papua: Kokoda 1 200 ft. VIII. 1933. L. E.

Cheesman, B.M. 1933-577’, ‘Paratype’, ‘Borona

laevigata, M. J. Balfour-Browne det’. ‘A.

d’Orchymont, det., Helochares (Agraphydrus)

laevigatus. J. Balf.-Browne’. I can find no

reference in the literature to this name and assume

it is an unused name.

Chasmogenus Sharp, 1882

Type species: Chasmogenus fragilis Sharp, 1882,

Central America; designation by monotypy.

Chasmogenus nitescens (Fauvel)

Philydrus nitescens Fauvel, 1883

Enochrus nitescens (Fauvel, 1883); Knisch 1924

Helochares (Crephelochares) nitescens (Fauvel,

1883); d’Orchymont 1939

Description (number examined 161)

Length 2.5-5.0 mm. Narrowly oval. Black.

Clypeus, edges of pronotum, lateral margins of

elytra often reddish, appendages testaceous. Front

edge of head broadly and shallowly concave, a

weak notch in front margin of head in some

specimens. Dorsal surface with small weakly

impressed well-separated punctures. In addition a

few larger punctures in row across front of head

and between front edges of eyes and pronotum

with two fields of larger punctures on either side

of midline towards front and a field of a few larger

ones on each side in middle; each elytron with

three or four ill-defined rows of well separated

large serial punctures. Each elytron with sharply

impressed sutural stria in apical half to three

quarters, diverging somewhat toward front close

to suture apically. Scutellum lacking punctures or

with a few weak ones. Maxillary palpus long,

slender, apical segment about two-thirds length of

penultimate, first and second segments subequal.

Femur with rugose portion covering all but small

portion near apex. Coxal plates seemingly

impunctate, but rugose and moderately covered

with setae rather as on femurs, bare shiny area in

midline apically. Abdominal sternites similarly

sculptured; a well-developed but relatively low

metasternal keel present. Apex of apical

abdominal segment with small notch. Aedeagus

as in Fig. 15, although there is considerable

variation among the specimens examined.

Types

Lectotype male: ‘Anse Vata — marass d’eau douce

— aotit — savés, ‘Nouvelle Caledonie’, bearing

original det label. ‘Philhydrus nitescens Fvl’, in

BELG, herein designated.

Paralectotypes: 3, — same data as lectotype, in

BELG, herein designated.

Distribution

Northern Territory

Black Point, Coburg Pen., ANIC; Katherine,

ANIC; 8 km N Mt Cahill, ANIC; 19 km NE by E

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES ii 7/

1 2 3 4

FIGURES 1-16. Tip of aedeagus. All are ventral views except for additional dorsal (12) and lateral (14) views of species

as follows; 1 H. anthonyae; 2 H. clypeaus; 3 H. tristis; 4 H. tenuistriatus; 5 H. luridus; 6 H. tatei;7 H. loweryae; 8 H.

dalhuntyi; 9 H. thurmerae; 10 H. percyi; 11 H. anchoralis; 12 H.anchoralis; 13 A. coomani; 14 A. coomani; 15 C.

nitescens; 16 H. foveicollis.

118 ¢€. H.S. WATTS

Mt Cahill, ANIC; Nourlangie Creek, ANIC; 6 km

SW by S, Oenpelli, ANIC.

Queensland

Ayr, ANIC; 5 ml [8 km] N Bloomfield River,

ANIC; Cairns, BELG; Cape Tribulation, ANIC; 3

km S by W Cooktown, Mission Beach, ANIC;

Giru, ANIC; Hann River, N. Laura, QDPIM;

Innisfail, ANIC; 7 km NE Innisfail, ANIC; Mt

Webb National Park, ANIC; Townsville, ANIC; 3

km ENE Mt Tozer, ANIC; 9 km ENE Mt Tozer,

ANIC; 11 km ENE Mt Tozer, ANIC; 5 km W by

N Rounded Hill, ANIC; 9 km SE Yeppoon,

ANIC.

New South Wales

Yuragin, NP, ANIC.

Papua New Guinea

Lae, BMNH; 7 ml [11 km] Lae-Bulolo Road,

BMNH; Gusap Markham Valley, ca 90 ml [145

km] W of Lae, BMNH.

Remarks

I have examined type material of C. abnormalis

(Sharp, 1890), C. simulator (Kuwert, 1922), C.

livicornis (Kuwert, 1889), C. ferrugatus

(Régimbart, 1903) and C. nigritulus (Régimbart,

1903) and consider the Australian specimens to

differ from all of these. This appears to be the only

Chasmogenus in Australia.

Helochares Mulsant, 1844

Hydrobaticus W. Macleay, 1871

Neohydrobius Blackburn, 1898

Type species: Hydrobaticus: Hydrobaticus tristis

W. Macleay, 1871, Gayndah, Queensland:

designation by d’Orchymont (1943). Helochares:

Dytiscus lividus Forster, 1771, Europe.

Neohydrobius: Philhydrus burrundiensis

Blackburn, 1890, Northern Territory, Australia;

designation by Blackburn (1890).

Subgenus Helochares

Helochares (Helochares) foveicollis

(Montrouzier)

Stagnicola foveicollis Montrouzier, 1860

Philhydrus burrundiensis Blackburn, 1890: syn.

nov. Neohydrobius burrundiensis (Blackburn,

1890), Blackburn 1889; syn. nov. ?Helochares

atropiceus Régimbart, 1903

Helochares (Helochares) foveicollis

(Montrouzier, 1860); Knisch 1924: d’Orchmont

1943

Helochares (Helochares) burrundiensis

(Blackburn, 1890); Knisch 1924; syn. nov.

Description (number examined 356)

Length 6.1—8.2 mm. Broadly oval. Black. Front

edge of head, margins of pronotum, lateral

margins of elytra and apical portions of

appendages testaceous. Front edge of head

sinuate. Head and pronotum with small to medium

sized (for genus) punctures, well impressed,

separated by about width of a puncture. Punctures

on side of head a little stronger and denser.

Punctures on elytra same size or somewhat

smaller than those on pronotum and a little

Sparser, particularly towards sides and apex. A

sparse row of serial punctures traceable in about

middle of each elytron and another weaker row

near lateral margin. Maxillary palpus long,

slender, second segment largest, apical segment

about two-thirds length of middle one. Femur with

rugose portion covering all but small portion near

apex. Coxal plates sparsely and weakly punctured,

covered in moderately impressed fine reticulation.

Sternites shiny, covered with relatively sparse,

small setose punctures; apex of apical sternite with

small notch. Weak to well developed metasternal

keel, highest apically.

Male: Aedeagus as in Fig. 16. Protarsi enlarged.

Claws on protarsi strongly recurved, inner one

with large scale-like vertical expansion at its base;

claw on mesotarsi similar in shape but not quite

as developed.

Female: Protarsi about two-thirds size of male’s,

claws simple; mesotarsi a little smaller than in

male, claws simple.

Types

P. burrundiensis: Syntypes: 1 in BM(NH) ‘T

2769’ ‘Philhydrus burrundiensis, Blackb.’ with

BM(NH) Type and Blackburn coll 1910-236

labels. 1 ‘NT, N. Territory’, ~Neohydrobius

burrundiensis Blackb., Co-type’, in SAMA, |

‘Philhydrus burrundiensis, Bl’. Co-type, in

SAMA.

I am uncertain of the status of these types. The

BM(NH) specimen is without locality label but is

mounted on a card with a black ‘T’ and a red

number in typical Blackburn style and carries the

species label in Blackburn’s handwriting. I have

little doubt that this was meant as the holotype.

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES 119

The first presumed syntype in SAMA fits

Blackburn’s style, except for his use of

Neohydrobius as the generic name which was not

described until nine years later by Blackburn

(1898). The second presumed syntype in SAMA

bears a single label not in Blackburn’s hand. At

some stage this specimen was remounted upside

down. In his 1898 publication, Blackburn

mentions original specimens. I am inclined to

believe that all three specimens are part of the

original series and that the two SAMA specimens

were re-examined and relabelled when Blackburn

described Neohydrobius. 1 nominate the BM(NH)

specimen as lectotype and the two SAMA

specimens as paralectotypes.

S. foveicollis: Not located. The type is supposedly

in the Bedel Collection in MNHN but cannot be

found there or in other collections in MNHN (Y.

Cambefort in litt.). A specimen in MNHN from

New Caledonia appears identical to Australian

specimens and it is on this basis that I synonymise

the species. d’Orchymont (1943) came to the

conclusion, based on very limited material, that

the two forms were very close and probably

conspecific.

H. atropiceus Régimbart. Syntypes. 2 in MNHN

seen. This New Guinean species appears to me to

be very similar if not conspecific with H.

foveicollis. According to d’Orchymont (1943) it is

a junior synonym of H. taprobanicus (Sharp,

1890) from Ceylon and differs from H. foveicollis

by its sculpture and male genitalia. I have not,

however, looked closely at these non-Australasian

forms and prefer to leave the question of whether

or not they are conspecific open at the moment.

Distribution

Northern Territory

3.2 km S Adelaide River, ANIC; Alligator

River, MV; Black Point, Coburg Pen., ANIC;

Borroloola, ANIC; 22 km WSW Borroloola,

ANIC; 5 km NNW Cahills Crossing, ANIC; 8

km ESE Cape Crawford, ANIC; Cooper Creek,

ANIC; Darwin, SAMA, CW; 96.5 km S Darwin,

ANIC; Edith River, ANIC; Elcho Island, ANIC;

Holmes Jungle, ANIC; Horn Islet, Pellew Group,

UQIC; Humpty Doo, QDPIM; Katherine, ANIC;

Koongarra, ANIC; Magela Creek, ANIC, SAMA;

Mataranka, ANIC; McArthur River, ANIC;

Melville Island, ANIC; 10 km E by N Mt Cahill,

ANIC; Nabarlek Dam, ANIC; Nourlangie Creek,

SAMA; Pine Creek, CW; Rimbija Island, ANIC;

Roper River, ANIC; South Alligator River, ANIC;

Tindal, ANIC.

Queensland

Ayr, ANIC; Bamaga, ANIC, UQIC; Bentinck

Is., SAMA; Burnett River, ANIC; Cairns, SAMA;

Cape Tribulation, QM; Cardstone, ANIC;

Carnarvon Range, AM; Clerrnont, AM; 19.3 km

WNW Cooktown, ANIC; 40 km N Cooktown,

ANIC; 50 km N Cooktown, QM; Cow Bay,

QDPIM; Darwin, CW; E Alligator, AM; Green

Hills, ANIC; Home Hill, CW; 7 km N Hope Vale

Mission, ANIC; Innisfail, AM, SAM; Iron Range,

ANIC, AM; 15 km W Irvinebark, ANIC;

Kirrama, ANIC; 75 km NW of Laura, QDPIM;

Mackay, MV; 80 km S Mackay, UQIC; Mission

Beach, UQIC; Mornington Is. Mission, SAMA; 5

km ESE Mt Finnigan, ANIC; 9 km ENE Mt

Tozer, ANIC; 3 km NE Mt Webb, ANIC;

Mutchilba, MV; 8 km S of Putty, ANIC;

Renilworth State Forest, UQIC; 5 km W by N

Rounded Hill, ANIC; 40 mile Scrub, ANIC;

Shiptons Flat, ANIC; South Johnstone, QDPIM;

15 km WNW South Johnstone, QDPIM; Stewart

Range, BELG; Strathmore Stn., QDPIM;

Toowoomba, ANIC; Townsville, CW, FIELD,

MV, QDPIM; 9.6 km SSE Yeppoon, ANIC.

New South Wales

2.4 km E Freshwater River, AM; Kyogle, AM;

Valery, ANIC.

Australian Capital Territory

Black Mountain, ANIC.

Western Australia

4 km SSW Cape Bertholet, ANIC; Derby,

WAM; Fitzroy Crossing, ANIC; 161 km E

Kununurra, ANIC; Mitchell Plateau, ANIC;

Wyndham, UQIC.

Papua New Guinea

1.6 km S Morehead, ANIC; Rouku, Western

District, ANIC.

Noumea

Noumea, MNHN.

Remarks

Very distinct from other Australasian

Helochares, by its size, shiny black colour and

virtual lack of elytral stria.

Subgenus Hydrobaticus W. Macleay, 1871

Helochares (Hydrobaticus) anchoralis Sharp

Helochares anchoralis Sharp, 1890

120

Helochares crenatus expansus Knisch, 1921,

1924; d’Orchymont 1943

Helochares (Hydrobaticus) anchoralis expansus

Knisch, 1921; d’Orchymont 1943

Description (number examined 7)

As for H. tristis except as follows. Length 5.5—

6.2 mm. Upper surface shiny, reddish brown with

scattered, ill-defined darker areas near border of

head and Y suture outlined in black. Head

uniformly covered in small-medium sized

moderately dense punctures; pronotal punctures of

uniform size, small for genus, separated from each

other by their own diameter or a little less, a few

larger setiferous setae in one line towards front on

each side; serial punctures of elytra confluent,

small, those at base about same size as those on

pronotum, somewhat larger towards apex and

laterally; interstrial punctures numerous,

moderately dense, smaller than those in stria; area

between elytron edge and most lateral striae

densely covered in large punctures, the more

lateral ones as large or larger than those in

adjacent striae. Elytron flanged. Striae form deep

grooves laterally and apically but interstrial areas

flat. Aedeagus as in Figs 11 and 12.

Distribution

Papua New Guinea

Finisterre Mts, Budemu, ca 1220m, BMNH;

Gusap Markham Vale, 145 km W of Lae, BMNH;

Lae-Bulolo Road, BMNH; 11 km Lae-Bulolo

Road, BMNH; Port Moresby, BMNH.

Remarks

I have not seen the types of either H. anchoralis

Sharp, or H. c. expansus Knisch, and rely on

d’Orchymont (1943) for my identification of this

species. The series of specimens in the BMNH

agree with Sharp and d’Orchymont’s descriptions,

and with the aedeagus illustrated by d’Orchymont.

Among Australasian Hydrobaticus with uniform

punctation, the species is distinctive by being large

and relatively flat with broad elytral flanges and

the punctures on the head smaller than the size of

the eye facets. Outside New Guinea the species is

widespread in South East Asia from Ceylon (type

locality) to the Philippines. The type locality of H.

c. expansus Knisch is given only as New Guinea.

Helochares (Hydrobaticus) anthonyae sp. nov.

Description (number examined 76)

As for H. tristis except as follows. Length 4.5—

C.H.S. WATTS

5.8 mm. Yellow-brown. Front and rear of head, Y

suture on head, portions of pronotum, serial

punctures on elytra and often a quite extensive

area on disc of elytra darker. Punctures on head

moderate in size and density, somewhat smaller

towards front in middle; punctures on pronotum of

uniform size, moderately sized, separated from

each other by their own diameter or slightly less;

strial punctures well impressed, not confluent, not

forming grooves, at least twice size of those on

pronotum, becoming progressively smaller in

apical quarter of elytron; interstrial punctures very

small, much smaller than strial punctures except

towards apex where the serial punctures are much

reduced in size. Sides of pronotum and elytra with

fine reticulation giving mat finish. Edges of elytra

not or only very weakly flanged.

Types

Holotype male: ‘New Guinea: Morobe Dist., ¢ 7

ml [11 km] Lae-Bulolo Rd. 30.xii.1964’, “Stn No.

120’, ‘M.E. Bacchus, BM 1965-120’, in

BM(NH).

Paratypes: 9 in BM(NH); 2 in SAMA; 2 in CW;

all with same data as holotype

Distribution

Northern Territory

Nourlangie Creek, ANIC.

Papua New Guinea

Lae, BMNH; 7 mls [11 km] Lae-Bulolo Road,

BMNH; 90 mls [145 km] W. of Lae; Mt

Lamington, BMNH; Okapa, E. Highlands District,

BMNH.

Remarks

Apart from the distinctive aedeagus (Fig. 1),

separated from other Helochares, particularly H.

loweryae, H. dalhuntyi and H. thurmerae, by

characters given in key. A New Guinean species

known from only one Australian specimen from

the Northern Territory

Helochares (Hydrobaticus) clypeatus

(Blackburn)

Hydrobaticus clypeatus Blackburn, 1891

Helochares (Hydrobaticus) clypeatus (Blackburn,

1890); Knisch 1924; d’Orchymont 1943

Description (number examined 377)

As for H. tristis except as follows. Length 5.1—

7.2 mm. Some ill-defined dark patches on elytra.

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES 121

Punctures on front of head fine, with a few much

larger ones (Fig. 17); punctures on rear of head,

pronotum and elytra of two sizes, the smaller

similar to those on front of head, the larger much

larger, in approximately equal numbers except on

elytra where they are smaller and more numerous;

large punctures on pronotum often separated by

less than their diameter (Fig. 19); serial punctures

a little larger, often confluent and forming shallow

grooves. Interstrial areas virtually flat apically.

Aedeagus as in Fig. 2.

Types

Holotype female: ‘T 3434NT’, ‘Hydrobaticus

clypeatus, Blackb.’” in BM(NH).

Distribution

Northern Territory

1 km NE of Cahills Crossing, East Alligator

River, ANIC; Berry Springs, ANIC; Bessie

Spring, ANIC; Coastal Plains Research Station,

CSIRO, Darwin, ANIC; Cooper Creek, ANIC; 16

km E of Daly River, SAMA; Howard Springs,

ANIC; 10 km N Jabiru, QDPIM; Kakadu NP,

NTM; Kapalga, QM; Koongarra, ANIC; Manton

Reservoir, NIM; 19 km NE by E Mt Cahill,

ANIC; 2 km N of Mudginbarry HS, ANIC;

Nourlangie Creek, ANIC; Oenpelli, ANIC;

Simpson Gap, ANIC.

Queensland

Atherton, ANIC; Ayr, ANIC; 8 km N

Bloomfield River, ANIC; Boar Pkt Road, ANIC;

Brisbane, CW, UQIC; Brookfield, QM;

Caloundra, CW; Cape Tribulation, CW; Cairns,

ANIC, CW, SAMA; Cape Flattery Road, QDPIM;

FIGURE 17. Head of H. clypeatus.

Cardstone, ANIC; Cardwell, ANIC; Carungra

Creek, QM; Cooloola, QM; Cow Bay, QDPIM;

Emerald, QDPIM; Flinders Island, SAMA; Fraser

Island, ANIC; Home Hill, CW; Gayndah, UQIC;

Groomeri, UQIC; Hann River N of Laura,

QDPIM; Hann River, 73 km NW Laura, ANIC;

Helenvale, ANIC; Hope Vale Mission, ANIC;

Innisfail, CW; Kuranda, ANIC, SAMA; Lam.

Nat. Pk, UQIC; 75 km NW Laura, QDPIM;

Mareeba, QDPIM; MclIlwraith ANIC, QDPIM;

16 km S Miriam Vale, UQIC; Mission Beach,

UQIC; Mossman, ANIC; Mourilyan, ANIC; 3 km

NE Mt Webb, ANIC; North Pine River, UQIC;

Nth Stradbroke Island, UQIC; 11 km WSW

Petford, QDPIM; Petrie, UQIC; 15 km NW of

South Johnstone, QDPIM; Tinaroo Dam, QDPIM;

Tolga, QDPIM; 7 km N Tolga, QDPIM;

Townsville, ANIC; Upper Mulgrave River, UQIC;

Walkamin, QDPIM; 40 km S of Weipa, QDPIM;

80 km N Weipa, NMV; Woodford, UQIC.

New South Wales

Coffs Harbour, UQIC; Congo, ANIC; Kyogle,

FIELD; Lansdowne SF, ANIC; 8 km W of Port

Macquarie, ANIC; 18 km W of Uki, ANIC;

Valery, ANIC; Windsor, ANIC; Wootton, ANIC.

Western Australia

Carson Escarpment, ANIC.

Remarks

A distinctive species best recognised by the

form of punctation on the upper surface in which

the large and small punctures are very different in

size (Fig. 19). This character is difficult to

describe in a key but once recognised is readily

used. The sparsity of large punctures on the front

portion of the head (Fig. 17) is usually a reliable

character but does not separate all H. clypeatus

from all H. tristis.

Helochares (Hydrobaticus) dalhuntyi sp. nov.

Description (number examined 30)

As for H. tristis except as follows. Length 4.2—

5.5 mm. Oval. Dark testaceous; patches of lighter

colour on upper surface particularly laterally and

at bases of elytra, palpi and tarsi lighter. Front

margin of head deeply and widely concave.

Punctures on head and pronotum uniformly sized,

small for Helochares, separated by less than own

diameter in most cases, somewhat larger laterally;

punctures in elytral interstriae numerous (Fig. 21),

much smaller than those on pronotum, a row of

larger seta-bearing punctures inwards from striae

12 C.H.S. WATTS

six; punctures in stria variably confluent, deeply

impressed, much larger than those on pronotum.

Aedeagus as in Fig. 8.

Types

Holotype male: right hand specimen of two

mounted on card, ‘Dalhunty River, Qld, 1/10/83,

C. Watts’, in SAMA.

Paratypes: 1, mounted upside down on same card

as holotype in SAMA; 3, mounted on same card,

same data as holotype, in CW; 1, ‘Captain Billy

Creek, CW’, in CW.

Distribution

Queensland

Captain Billy Creek, CW; Dalhunty River, CW;

East Claudie River, UQIC; Iron Range, ANIC,

AM, UQIC; 2 km NNE Mt Tozer, ANIC; 3 km

ENE Mt Tozer, ANIC; 6 km ENE Mt Tozer,

ANIC; 8 km E by N of Mt Tozer, ANIC; 9 km

ENE Mt Tozer, ANIC; 11 km ENE Mt Tozer,

ANIC.

Northern Territory

Darwin, CW; Roderick Creek, Gregory NP,

NIM.

Remarks

Helochares dalhuntyi is difficult to separate

from H. loweryae except by the aedeagus. The

punctation of the upper surface is stronger than in

all but a few H. loweryae, but I have been unable

to reliably quantify this difference.

Helochares (Hydrobaticus) loweryae sp. nov.

Description (number examined 25)

As for H. tristis except as follows. Length 4.4—

5.5mm. Oval. Light testaceous; patches of darker

colour on dorsal surface. Front margin of head

deeply and widely concave. Punctures on head and

pronotum uniform in size, small for genus,

separated by about own diameter or less in most

cases, only slightly larger laterally; punctures in

elytral interstriae numerous, much smaller than

those on pronotum, a row of large setose punctures

inwards from stria 6; interstriae sub-obsolete

apically, on shoulders and between lateral-most

striae; punctures in stria separate, deeply

impressed, much larger than those on pronotum,

larger towards sides than on disc; lateral punctures

on elytra small but well impressed, larger than

those in adjacent interstriae. Aedeagus as in Fig.

Types

Holotype male: ‘Mt Lamington, N.E. Papua, 1300

to 1500 feet, C. J. McNamara’, in SAMA.

Paratypes: 1, ‘New Guinea: Morobe Dist., Lae-

Bulolo Rd, 30.xii.1964’. ‘Stn. No. 131’, in

BM(NH). 1, ‘New Guinea: Morobe Dist., Gusap,

Marthany Valley, c. W. of Lae. 22-30.i. 1965’.

‘M.E. Bacchus, BM 1965-120’. ‘Stn. No. 166’, in

BM(NH). 1, ‘New Guinea: Morobe Dist., c. 7 ml

Lae-Bulolo Rd. 30.x1i.1964’. ‘Stn. No. 120’.

‘M.E. Bacchus, BM 1965-120’, in BM(NH). 1,

‘New Guinea: Morobe Dist. Herzog Mts., Vagau.

c. 4,000 ft 4-17.1.1965’. ‘M. E. Bacchus, BM

1965-120’, in CW.

Distribution

Northern Territory

Darwin River Reservoir, NTM; Holmes Jungle,

ANIC; Howard Springs, CW, NTM.

Papua New Guinea

Bulolo, BM(NH); Kokoda, BM(NH); Morobe

District, BM(NH); Mt Lamington, SAMA; Lae-

Bulolo Road, BM(NH); 5 km Lae-Bulolo,

BM(NH); 11 km Lae-Bulolo, BM(NH); 145 km

W. of Lae, BM(NH); Vagav, Herzog Mts,

BM(NH).

Remarks

The specimens from the Northern Territory

differ from typical specimens by having somewhat

stronger interstrial punctures and. having the

lateral ‘wings’ of the aedeagus (Fig. 8) more

separate from the centre piece than in New Guinea

specimens. Close to H. dalhuntyi and difficult to

separate from it other than by the shape of the

aedeagus. Most specimens can be separated by

the different strength of the dorsal punctation but

this is difficult to use unless specimens of both

species are available. The Northern Territory

specimens appear intermediate in this regard.

Helochares (Hydrobaticus) luridus (W. J.

Macleay)

Hydrobaticus luridus W. J. MacLeay, 1871

Description (number examined 144)

As for H. tristis except as follows. Length 3.5—

5.4 mm. Larger punctures usually smaller than

those of H. tristis, serial punctures usually

confluent over most of elytra forming quite strong

grooves, interstrial areas towards apex of elytra

and laterally weakly ridged, ridging effect

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES 123

accentuated by relatively deeply indented striae;

lateral punctures around elytron get progressively

smaller towards apex where they are usually less

than one quarter diameter of those towards front

corner of elytron (Fig. 23). Aedeagus as in Fig. 5.

Types

Holotype, ?sex. ‘Hydrobaticus luridus McL.W.

Gayndah’ ‘K1924’ with small circular red label

and red Holotype label, in AM.

Distribution

Northern Territory

IG'O2ZS, WOP22 18, INMIME IG OPS, IOP2AS.

NTM; 16°07'S, 130°25'E, NTM; Sth Alligator

River, QM; Cooper Creek, ANIC, SAMA;

Fergusson River, ANIC; Gregory NP, NTM;

Howard Springs, ANIC; Humbert River, NTM;

Humpty Doo, QDPIM; Junction of Arnhem Hwy.

and Oenpelli Road, NTM; Oenpelli Road, NTM;

Kakadu, NTM; Katherine, SAMA; Larrimah,

QM; 13 km S Lawn Hill Stn., QM; McArthur

River, ANIC; 19 km E by N of Mt Cahill, ANIC;

19 km WSW Mt Cahill, ANIC; 19 km NE by E

Mt Cahill, ANIC; 12 km NNW Mt Cahill,

SAMA; 2 km N Mudginbarry HS, ANIC;

Nabarlek Dam, ANIC; 18 km E by N of Oenpelli,

ANIC; Renner Springs, CW; Tennant’s Creek,

NMV; 12 km NNE Victoria River Downs, ANIC.

Queensland

Annan River, ANIC; Ayr, ANIC; Barron Falls,

ANIC; Brookfield, QM; 29 km S Burketown,

QM; Burnett River N of Eidswald, ANIC;

Cardstone, ANIC; Cairns, ANIC, Charleville,

ANIC; 25 km N Coen, CW; Cooktown, ANIC,

CW; 40 km N Cooktown, ANIC; Cunnamulla,

AM, SAMA; Daintree, ANIC; Dalby, SAMA;

Dipperu, QM; Ellery Creek, NMV; Gayndah,

SAMA; 12 km SE Gympie, NMV; Innisfail, QM;

Iron Range, UQIC; Laura, CW; 73 km NW by W

Laura, ANIC; 75 km NW Laura, QDPIM; Little

Laura River, QM; Mary Creek, ANIC; Mcll wraith

Range, CW; 34 km S Mirian Vale, ANIC;

Mossman, ANIC, SAMA; Mt Garnet, CW; 3 km

ENE Mt Tozer, ANIC; 14.5 km W Paluma,

ANIC; 11 km NSW Petford, QDPIM;

Rockhampton, SAMA; Rocklea, QM; 15 km

NNW South Johnstone, QDPIM; Station Creek,

ANIC; Strathmore Station, QM; Tolga, QM; 7 km

NE Tolga, QDPIM; Townsville, AM.

New South Wales

Valery, ANIC.

Western Australia

Carson Escarpment, ANIC; Kalumburu, ANIC;

Mitchell Plateau, ANIC; 5 km SE Pago Mission,

FIELD:

Remarks

The distinctive aedeagus (Fig. 5) is the only

reliable character separating H. luridus from H.

tristis. Over most of its range H. tristis has no

trace of elytral ridging or indented stria, which are

almost always a feature of HA. luridus.

Unfortunately it is in the area of distributional

overlap in southern Queensland that the two

species are hardest to separate since most H.

tristis from this area have quite well marked apical

ridging and grooving on elytra. The character used

in the key—small lateral punctures at apex of

elytron in H. luridus—appears to work, but is

often difficult to see clearly and, at best, is a

subjective character.

H. luridus was synonymised with H. tristis by

Knisch (1924), Zaitzev (1908) and by

d’Orchymont (1943). These species are very

similar, but readily differentiated by the aedeagus.

The holotypes of both species are from Gayndah

in southern Queensland where both species are

known to occur. Both types are fragile and I think

females. I have not attempted to dissect them. The

type of H. luridus is paler and is almost identical

to a male specimen of the northern species from

Gayndah in SAMA. The type of H. tristis is darker

and agrees in diagnostic features with the southern

form.

Helochares (Hydrobaticus) marreensis sp. nov.

Description (number examined 147)

As for H. tristis except as follows. Length 4.0—

5.4 mm. Elongate, reddish-brown including

clypeus. Darker markings on head and pronotum

vague and often absent, elytra rather evenly

speckled brown and reddish-brown or, in some

examples, with linear dark lines. Punctures on

head numerous, dense, large at base, grading to

quite small in front; punctures on pronotum even,

moderately large, most separated by less than their

diameter, particularly towards sides; punctures in

interstriae numerous, strong, those at base near

suture about same size as those on pronotum,

becoming smaller towards rear and sides of elytra;

strial punctures almost confluent, weakly

impressed, same size or slightly smaller than those

on pronotum and base of elytron. I have been

unable to distinguish the aedeagus from that of H.

tatei (Fig. 6).

124 C.H.S. WATTS

FIGURES 18-23. 18 Pronotal disc of H. percyi; 19 Pronotal disc of H.clypeatus, 20 Portion of elytral disc of H. tristis;

21 Portion of elytral disc of H. dalhuntyi; 22 Tip of elytron of H. tristis; 23 Tip of elytron of H. luridus.

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES

Types

Holotype male: ‘12°23'S 132°56'E, 7 km NW by

N of Cahills Crossing, East Alligator River, NT,

27.v.73, E.G. Matthews’, in SAMA.

Paratypes: 3, ‘12°26'S 132°56'E, Cahills Crossing,

NT, East Alligator River, 29.v.73, at light, E.G.

Matthews’, in SAMA; 2, ‘12°48'S 132°42'E,

Nourlangie Creek, NT, 8 km-N of Mt Cahill,

21.v.73, at light, E.G. Matthews’, in SAMA; 2,

*12°25'S 132°58'E, 1 km N of Cahills Crossing,

NT, East Alligator River, 7.vi.73, Upton and

Feeham’, in ANIC; 2, “12°46'S 132°39'E, 12 km

NNW of Mt Cahill, NT, 15.vi.73, Upton and

Res vain; il ANNICS Il, “WA 23S WsvPxalsi, 7 lain

NW by N of Mt Cahills Crossing, East Alligator

River, NT, 9.vi.73, Upton and Feeham’, in ANIC;

3, ‘Forbes, NSW, 15.iii.63, E.W.’, in CW.

Distribution

Northern Territory

16°02'S, 130°23'E, NTM; Adelaide River,

ANIC; Baroalba Creek Springs, ANIC; 13 km

SW Borroloola, ANIC; 30 km NE by E

Borroloola, ANIC; 1 km N Cahills Crossing,

SAMA, ANIC; 5 km NNW Cahills Crossing,

ANIC; 7 km NW by N Cahills Crossing, SAMA,

ANIC; Cahills Crossing, SAMA, ANIC; Cooper

Creek, SAMA, ANIC; Daly River, ANIC, NMV;

Fogg Dam, NTM; Howard Springs, CW; 10 km

N Jabiru, QDPIM; Junction of Arnhem Hwy and

Oenpelli, NIM; Katherine, ANIC; Koongarra,

ANIC; McArthur River, ANIC; 13 km N Mt

Cahill, SAMA; 9 km N by E Mudginbarry HS,

ANIC; 6 km SW by S Oenpelli, ANIC, SAMA;

Renner Springs, SAMA; Tindal, ANIC; U.D.P.

Falls, NTM; 30 km N Wauchope, ANIC.

Queensland

Bunya Mountains, ANIC; 16 km S Miriam

Vale, UQIC; Crystal Creek, ANIC; Cunnamulla,

QM, SAMA; Dalby, QM, SAMA; 68 km East

Roma, QM; 96 km E Hughenden, UQIC; Julia

Creek, CW; Marina Plains, QDPIM; 16 km S

Miriam Vale, UQIC; Musgrave HS, ANIC;

Nocundra, ANIC; Normanton, SAMA; Silver

Plains, UQIC.

| New South Wales

| Canowindra, VM; Forbes, CW; Gilgandra,

CW; Moree, SAMA; Mt Kaputar, ANIC.

Victoria

Lake Hattah, ANIC, QDPIM; Wyperfield

} National Park, ANIC.

125)

Western Australia

3 km S Coulomb Pt, ANIC; Fitzroy Crossing,

ANIC; Kununurra, ANIC; Lennard River Xing,

WAM.

South Australia

Marree, SAMA.

Remarks

An elongate, parallel sided species with strong

interstrial punctures which, over most of elytra,

are as large as the strial punctures. Uniquely,

among Australasian species at least, the aedeagi

of H. marreensis and H. tatei are so similar that I

have not been able to distinguish them. The two

species are readily separated by the punctation of

the upper surface which is much stronger in H.

marreensis, and the presence of more than one

row of punctures in interstriae 7-8 and 9-10 in H.

marreensis compared to a single row in H. tatei.

Helochares (Hydrobaticus) percyi sp. nov.

Description (number examined 101)

As for H. tristis except as follows. Length 4.8-

6.0 mm. Oval, rather flattened. Yellow-brown.

Clypeus darker, extreme sides of pronotum and

elytra lighter. Punctures on head and pronotum

large, dense, of uniform size (Fig. 18), those at

sides of pronotum and head slightly larger and

closer together; interstrial punctures small to

medium, dense, uniform in size; serial punctures

nearly confluent, deep, larger than punctures on

pronotum. Interstrial areas weakly ridged near

apex and laterally. Aedeagus as in Fig. 10.

Types

Holotype male: ‘Boar Pkt.Rd., N.Q. 2 12.70 J.G.

Brooks’, in ANIC.

Paratypes: 1, ‘Boar Pkt.Rd., N.Q. 1/70 J.B.’

‘J.B.B. 27’ J.G. Brooks. Bequest, 1976, in ANIC;

1, ‘Boar Pocket Road, ca 8 km N of Gillies Hwy,

Qld 21.11.70, at light, J. G. Brooks’, ANIC; 2,

‘Mcllwraith Rng., Weather Stn., N. Qld, 23/7/82,

C. Watts’, in SAMA.

Distribution

Northern Territory

Adelaide River, ANIC.

Queensland

Atherton, QDPIM; Brisbane, UQIC; Cardstone,

ANIC; Cairns, CW; Eidsvold, ANIC; Emu Vale,

UQIC; 8 km N of Gillies Highway, ANIC; 12 km

126

SE Gympie, NMV; 37 km SSE Ingham, ANIC;

Iron Range, UQIC; Jimboomba, ANIC; Julatten,

UQIC; Kenilworth State Forest, AM;

Koombaloomba, BM(NH); Kuranda, ANIC; 18

km W Mareeba, QDPIM; Mcllwraith Range,

Weather Stn., CW; 117 km NW by W Laura,

ANIC: Mossman, ANIC; Mt Finnigan, ANIC; Mt

Spec, ANIC; Oxley Creek, QM; Palmerston

National Park, ANIC, UQIC; 24 km W of

Paluma, ANIC; Nth Pine River, QM; Sth Pine

River, QM: 32 km S of Ravenshoe, ANIC; 15 km

WNW South Johnstone, ANIC; Tolga, QDPIM;

Wilson’s Peak, UQIC.

New South Wales

Armadale, CW; Blue Mountains, ANIC;

Canterbury, SAMA; Coffs Harbour, UQIC; 12 km

N Eccleston, NMV; 9 km SW Gloucester, NMV;

Hastings River, ANIC; Nepean River, ANIC;

Salisbury, UQIC; Taree, ANIC; 18 km W of Uki,

ANIC; Wahroonga, ANIC.

Australian Capital Territory

Black Mountain, ANIC.

Western Australia

Carson Escarpment, ANIC.

Remarks

A broad flat species with large close-packed

punctures on pronotum (Fig. 18). Many specimens

have broad, weakly-flanged elytra often distinctly

truncated at apex. A wet area species common in

north Queensland but present as far south as the

Blue Mountains and the Australian Capital

Ternitory.

Helochares (Hydrobaticus) tatei (Blackburn)

Hydrobaticus tatei Blackburn, 1896

Helochares (Hydrobaticus) tatei (Blackburn,

1896), Knisch 1924; d’Orchymont 1943

Description (number examined 483)

As for H. tristis except as follows. Length 3.4—

4.8 mm. Elongate oval, elytra variegated light and

dark-brown. Head and pronotum quite densely

covered with large well impressed punctures, a

little larger laterally, most punctures on pronotum

separated by less than half their widths; interstrial

punctures on elytra fine, numerous, those in

interstriae 7-8 and 9-10 (ignoring short innermost

stria) arranged in single loose row; strial punctures

large, much larger than punctures on pronotum,

mostly confluent, deeply impressed, forming strial

C.H.S. WATTS

grooves, interstrial area flat. Aedeagus as in Fig.

Types

Syntypes: One, in NMV mounted on a card

labelled ‘Reedy Ck.’ and bearing labels, ‘Cent.

Aust. Coll. Horn. Exp. Pres. 7.94’, ‘Hydrobaticus

tatei, Blackb’, in addition it has a modern red label

with ‘Type’ printed on it; two specimens on the

same card on which is written “T Co-type 5484

Palm Ck.’ in Blackburn’s hand and labelled

‘Hydrobaticus tatei Blackb.’ in SAMA; One

specimen labelled ‘5484 Palm Ck, Hydrobaticus

tatei Blackb. co-type’ in SAMA; ten specimens in

NMV mounted two to a card on each of which is

written ‘Palm Ck.’ in Blackburn’s hand; two of

these are labelled ‘Cent. Aust. Coll. Horn. Exp.

Pres. 7.97’ and ‘Hydrobaticus tatei Blackb. det. by

Blackb.’, one pair bears only the locality label,

another only the species label and the last pair has

no label. All are clearly from the same series as

the other syntypes.

Since the use of ‘T’ to denote his Holotypes was

a normal practice of Blackburn’s. I consider the

specimen in SAMA so labelled to be the specimen

intended by Blackburn as the Holotype. I nominate

it as the lectotype and all the other specimens

mentioned above as paralectotypes.

Distribution

Northern Territory

Adelaide River, BELG, ANIC; Berry Springs,

ANIC; 33 km SW Borroloola, ANIC; 7 km NW

by N of Cahills Crossing, East Alligator River,

ANIC; Cahills Crossing, East Alligator River,

ANIC; Coastal Plains Station, ANIC; Cooper

Creek, ANIC; Darwin, BELG, CW; Fogg Dam,

ANIC, NTM; Howard Springs, CW, NTM;

Huckitta, ANIC, SAMA; Humpty Doo, ANIC;

Jim Jim Creek, ANIC, SAMA; Junction Arnhem

Hwy and Oenpelli Road, NTM; Kakadu N P,

NTM; Katherine, ANIC; Koongarra, ANIC;

Magela Creek, ANIC; Manton Reservoir, NTM;

McArthur River, ANIC; 16 km E by N of Mt

Cahill, ANIC; Mt Gilruth, QM; 9 km N by E of

Mudginbarry HS, ANIC, SAMA; Nourlangie

Creek, ANIC; 18 km E by N. of Oenpelli, ANIC;

4,8 km S Renner Springs, SAMA; South Alligator

R., ANIC, QM; 6.5 km W Timber Creek, SAMA.

Queensland

Archer River, ANIC; Archer Bend, CW; Boggy

Creek, ANIC; 25 km N, Coen, CW; NW by W.

Laura, East Claudie River, UQIC; 60 km S, Coen,

CW: Colosseum Creek, 16 km S Miriam Vale,

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES

UQIC; Cooktown, ANIC; Dalhunty River, CW;

Hann River, 73 km S of Coen N. CW; Hann

River, 110 km S of Coen N. NMYV; Iron Range,

UQIC; Mornington Island Mission, SAMA; 3 km

ENE Mt Tozer, ANIC; Musgrave, ANIC;

Normanton, SAMA; Rockhampton, ANIC.

New South Wales :

32 km SSW Bourke, SAMA; Whitton, SAMA.

Western Australia

21°31.55 S., 119°06.57 E, WAM; Beverley

Springs, WAM; Behn River, SAMA; 172 km SSE

of Carnarvon, ANIC; Carson escarpment, ANIC;

Charnely River, 40 km N, Beverley Springs,

WAM; Dampier Island, FIELD; Drysdale River,

ANIC; Milstream, ANIC, WAM; Mitchell

Plateau, ANIC, FIELD; 8 km NNE Mt Broome,

WAM; Murchinson River, ANIC; 5 km S, Pago

Mission, FIELD; Woodstock Station, WAM.

South Australia

Arkaroola Creek, SAMA; Brachina Creek,

SAMA; Mt Chambers Gorge, CW, SAMA; 16

km E Curdimurka, SAMA; Eringunda Valley,

SAMA; Salt Creek, SAMA; Wilpena Pound,

SAMA.

New Caledonia

Grotte de Ninain Rev Poya, SAMA.

Remarks

Resembles H. (H.) marreensis in its narrow

shape and similar male genitalia. H. tatei can be

separated from this species by having interstrial

punctures much weaker than those in adjacent

striae and having only one row of punctures in

interstriae 7-8 and 9-10. As far as I know this is

the first record of this species from New

Caledonia.

Helochares (Hydrobaticus) tenuistriatus

Régimbart

Helochares tenuistriatus Régimbart, 1908

Helochares (Hydrobaticus) tenuistriatus

Régimbart, 1908; Knisch 1924; d’Orchymont

1943.

Description (number examined 9)

As for H. tristis except as follows. Length 5.2—

6.5 mm. Oval. Distinctly reddish, upper surface

with darker markings. Front margin of head

broadly concave. Punctures on head and pronotum

of two sizes, smaller %4—% diameter of larger

127

which are separated by about their own diameter,

slightly larger laterally; punctures on elytron rather

similar in size, with strial punctures relatively

small and weak and not much larger than larger

interstrial punctures, which in turn are often not

much larger than smaller interstrial punctures;

interstrial punctures not confluent, not forming

grooves or ridges. Aedeagus as in Fig. 4.

Types

Syntypes, in MNHN. Type locality, Mongers

Lake, North Subiaco, Perth, Western Australia; 1,

from same series in WAM. Lectotype not

designated.

Distribution

Western Australia

Busselton, CW; Bunbury, AM; Camel Lake,

WAM; King George Sound, AM; Mogumber,

WAM; North Lake, Fremantle, FIELD;

Ravensthorpe, ANIC; Swan River, ANIC;

SAMA; Swan View, ANIC.

Remarks

Separated from H. tristis by the distinctive

reddish hue and the comparative uniformity of

punctures on the elytra. The aedeagus is

distinctive. In the relatively few specimens I have

seen there is no hint of elytral grooves or ridges as

in some H. tristis and virtually all H. luridus.

Although I saw the types many years ago they are

temporarily unavailable and I have relied on my

notes and descriptions by Régimbart and

d’Orchymont and the illustration of the aedeagus

given by d’Orchymont to identify this species. I

have not designated a lectotype due to the

unavailability of the Paris specimens.

Helochares (Hydrobaticus) thurmerae sp. nov.

Description (number examined 15)

As for H. tristis except as follows. Length 3.8—

5.0 mm. Reddish-yellow; rear of head, front of

pronotum, serial punctures and other areas of

upper surface darker. Punctures on head subequal,

small for subgenus, separated by half to one times

their diameters, somewhat smaller towards front;

punctures on pronotum a little denser and stronger

than on head; strial punctures on elytra well

impressed, much larger than those on pronotum,

separate but close to each other, those on disc

somewhat smaller than elsewhere including apical

declivity; interstrial punctures of uniform size,

much smaller than those on pronotum, moderately

128 C.H.S. WATTS

impressed, those of lateral fringe larger. Elytra

weakly flanged. Aedeagus as in Fig. 9.

Male: Protarsi and claws a little stouter than in

female.

Types

Holotype male: ‘New Guinea: Morobe Dist.,

Gusap, Markham, Valley c. 90 ml W. of Lae,

1,000 ft 27-30.i.1965’. ‘M.E. Bacchus, BM,

1965-120’. ‘Stn. No. 166’, in BM(NH).

Paratypes: 4, in BM(NH); 1, in CW; 1, in SAMA,

same data as Holotype.

Distribution

Papua New Guinea

Gusap Markham Valley, 145 km W of Lae,

BM(NH); Lae, BMNH; 11 km Lae-Bulolo Rd.,

BM(NH).

Remarks

Known only from a relatively small area of New

Guinea. Apart from the aedeagus, H. thurmerae

can be separated from the rather similar H.

loweryae and H. dalhuntyi by the strial punctures

being larger and fewer than in those species.

Helochares (Hydrobaticus) tristis Macleay

Helochares tristis Macleay, 1871

Hydrobaticus australis Blackburn, 1888; syn.

nov.

Description (number examined 250)

Length 3.8-6.1 mm. Oval, widest behind centre

of elytra. Yellow-brown. Clypeus, back of head,

markings in central panel of pronotum and much

of underside darker. Front margin of head

shallowly and widely concave. Head and

pronotum densely covered with punctures of two

sizes, in approximately equal numbers, larger

much larger than smaller; punctures denser, larger

and more numerous laterally. Elytra similarly

punctured (Fig. 20), with, in addition, ten more or

less distinct lines of serial punctures; lateral band

of punctures on elytron continues strongly to apex.

Serial punctures same size or larger than larger

interstrial punctures. Pronotum and elytra weakly

flanged. Maxillary palpi elongate, first and second

segments subequal in length, apical somewhat

shorter. Femur with rugose portion covered with

short setae, covering all but small portion of apex

of femora. Underside rugose punctate. Sternites

with covering of short setae; apex of apical

sternite broadly notched, notch with row of stout

setae along margin practically filling up area of

notch.

Male: Maxillary palpi, tarsi and tarsal claws

slightly stouter than in female. Aedeagus as in

Fig. 3.

Types

H. tristis: Holotype, ?sex ‘Hydrobaticus tristis

Mcl. W. Gayndah’ ‘K19621’ with small circular

red label and red Holotype label, in AM.

H. australis: Syntypes: two male specimens

mounted on same card, under the left hand

specimen is the number 410 & “T’. The pin has

the following labels, BM(NH) Type label,

‘Blackburn coll. 1910-236’, ‘Hydrobaticus

australis Blackb.’, in Blackburn’s hand, in

BM(NH); one labelled ‘410, Port Lincoln

Blackburn, Hydrobaticus australis Blackb., co-

type’ in SAMA; two mounted on same card

labelled ‘410, Port Lincoln Blackburn, Australis

Blackb.’ in SAMA (there is an empty card

labelled ‘410’ on same pin). I nominate the left

hand specimen in the BM(NH) below which is

written ‘410 T’ as the lectotype, and the remaining

syntypes as paralectotypes. Synonymy is based on

examination of types.

Distribution

Northern Territory

Ayers Rock, UQIC; 80 km W Mt Olga, WAM,

160 km W Mt Olga, WAM; Vaughan Springs,

CW.

Queensland

Brisbane, UQIC; Brookfield, QM; Cunnamulla,

QM; Dalby, QM; Goomeri, UQIC; Green Island,

WAM: Jimboomba, ANIC; Malanda, CW; 16 km

S Miriam Vale, UQIC; Stanthorpe, BELG, UQIC;

Tambourine Mt., QM.

New South Wales

Blowering Dam, ANIC; Cooma, ANIC; 23 km

NNE Coonabarabran, ANIC; 15 km S

Condobolin, ANIC; Culcairn, ANIC; Deniliquin,

ANIC: Forbes, CW; Forest Reefs, SAMA; Kitty’s

Creek, ANIC; Macleay River, ANIC;

Muswellbrook, ANIC; Native Dog, ANIC;

Sydney, BELG, NMV; Yanco Creek, ANIC.

Australian Capital Territory

Acton, ANIC; Canberra, ANIC; Gungahlin,

ANIC; 3 km NW Tharwa, ANIC.

AUSTRALASIAN AGRAPHYDRUS, CHASMOGENUS AND HELOCHARES 19)

Victoria

Alexandra, NMV; Anakie, NMV; Avoca,

NMV; 11 km NNW Ballan, ANIC; Ballarat,

ANIC; Birchip, NMV, UQIC; Dandenong Range,

NMV, 34 km E Echuca, ANIC; Ferntree Gully,

NMV; Grampians, UQIC; Healesville, CW,

NMV; Inglewood, NMV; Iona, ANIC; L. Colac,

NMV; L. Punrumbeti, NMV; Lake Hattah, NMV;

Latrobe River, NMV; Lilydale, SAMA; Maffra,

NMV; 8 km SW Maffra, ANIC; Melbourne,

NMV, UQIC; Mitta Mitta River, NMV;

Mordialloc, NMV; Mornington, UQIC; Natya,

NMV; Preston, NMV; Sea Lake, UQIC; Goudie,

UQIC; 13 km SE Shepparton, ANIC; Stawell,

CW; Suggar Buggar, NMV; Tynong, NMV;

Morwell, ANIC; 4 km W Violet Town, ANIC;

Wartook, NMV; Whittlesea, NMV.

Tasmania

Asbestos Range NP, ANIC; East Tamar,

SAMA; Launceston, SAMA; Hobart, BELG,

NMV, SAMA.

South Australia

Adelaide, CW; Blanchetown, SAMA; Chain of

Ponds, CW; 13km N Ernabella, FIELD; Everard

Ranges, SAMA; Flinders Ranges, CW; Gawler,

SAMA; Gawler Ranges, SAMA; Mannum, CW;

Mt Barker, SAMA; Murray River, NMV; Olary,

ANIC; Scorpion Springs C.P., SAMA.

Western Australia

8 km S Giles, WAM; Gill’s Pinnacle, WAM.

Remarks

Elytra are usually smooth (except for punctures)

(Fig. 22) but in some specimens, particularly from

more northern localities, the strial punctures are

confluent towards apex forming grooves, and in

some the interstrial areas are ridged accentuating

the grooving in a way very similar to H. luridus.

This species appears to be replaced in south-west

Western Australia by H. tenuistriatus, a slightly

larger species recognisable by its uniform reddish

colour and strong interstrial punctures. H. tristis

is sympatric with H. luridus in southern

Queensland. The difference between these two

species is discussed under H. luridus.

ACKNOWLEDGMENTS

Without the support of a professional infrastructure

this work would have been impossible. I thank the

curators of the collections listed earlier for allowing me

to examine specimens in their care; Ms D. Churches, Ms

R. Cherrington and Mrs V. Wade typed the manuscript;

Ms J. Thurmer drew the figures of the male genitalia; Ms

J. Forrest took the SEM photographs; Mrs M. Anthony

ferreted out obscure references and Dr E. Matthews

much improved the manuscript. I thank them for their

help in what are perhaps the less exciting aspects of

scientific endeavours but nevertheless necessary ones.

REFERENCES

ANDERSON, J. M. E. 1976. Aquatic Hydrophilidae

(Coleoptera): The biology of some Australian species

with descriptions of immature stages reared in the

laboratory. Journal of the Australian Entomological

Society 15: 219-228.

BLACKBURN, T. 1888. Notes on Australian

Coleoptera, with Description of New Species.

Proceedings of the Linnean Society of New South

Wales Series 2 3: 805-875.

BLACKBURN, T. 1890. Notes on Australian

Coleoptera, with Description of New Species. Part III.

Proceedings of the Linnean Society of New South

Wales 4(1889): 445-482.

BLACKBURN, T. 1891. Notes on Australian

Coleoptera, with Description of New Species.

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420.

THE KAMPTOZOAN PEDICELLINA WHITELEGGI JOHNSTON & WALKER, 1917 AND

OTHER PEDICELLINIDS IN AUSTRALIA AND NEW ZEALAND

K. WASSON

WASSON, K. 1995. The kamptozoan Pedicellina whiteleggii Johnston & Walker, 1917 and

other pedicellinids in Australia and New Zealand. Records of the South Australian Museum

28(2): 131-141.

Colonies of the kamptozoan (entoproct) Pedicellina whiteleggii are ubiquitous in the low

intertidal and shallow subtidal zones of southern Australia and New Zealand. This species has

been known under a variety of names, but can be distinguished from other pedicellinids by a

suite of traits including longitudinal rows of conspicuous oblong cells on each tentacle, and a

tall, narrow, particle-covered larva. Pedicellina whiteleggii is re-described and illustrated, and

compared to all other pedicellinids reported from the waters around Australia and New Zealand.

P. whiteleggii is a senior synonym of P. hispida and appears to be very similar to a number of

other pedicellinids reported from these waters. However, P. whiteleggii is clearly distinct from

three other species (P. cernua, P. compacta and P. pyriformis) known from Australia and New

Zealand. The distribution of all these pedicellinids is discussed, and the importance of larval

traits in kamptozoan taxonomy is emphasised.

K. Wasson, Biology Department, University of California, Santa Cruz, CA 95064, USA.

Manuscript received, 30 May 1995.

Members of the phylum Kamptozoa

(Entoprocta) have rarely been studied in Australia

and New Zealand. There are about twenty

published reports on the kamptozoan fauna of

these regions, and only a few of these have

involved detailed taxonomic investigations

(Hastings 1932, Johnston & Angel 1940, Ryland

1965).

Examination of museum collections revealed

that only one pedicellinid species is common in

the shallow waters of Australia and New Zealand.

This species, Pedicellina whiteleggii Johnston &

Walker, 1917, accounts for most of the museum

specimens of Pedicellina from these waters.

Published records and museum labels refer to

these specimens under various names, but a suite

of distinctive traits unites them. The ubiquitous

nature of this species on these southern seashores

was reinforced by field collections which indicated

that colonies were present on many low intertidal

and shallow subtidal substrata at all sites

surveyed.

The purpose of this report is to unify these

various specimens under one name, Pedicellina

whiteleggii, and to justify this unification by

describing the configuration of traits which

characterises members of this species. This re—

description of P. whiteleggii is followed by a brief

review of all other pedicellinids known from

Australia and New Zealand.

MATERIALS AND METHODS

I examined all pedicellinid kamptozoans from

localities around Australia and New Zealand

deposited in the collections of the South

Australian Museum (SAM), the Australian

Museum (AM), the Museum of Victoria (NMV),

the Museum of Tropical Queensland (MTQ), the

Otago Museum (OM), the Portobello Marine

Laboratory of the University of Otago (PML), and

the British Natural History Museum (BMNH).

Apparently no other museums contain collections

of pedicellinids from Australia or New Zealand.

I collected live pedicellinids from the low

intertidal or shallow subtidal zone at Fairlight,

Port Jackson, New South Wales; Aldinga Reef,

Gulf St. Vincent, South Australia; Pukerua Bay,

North Island, New Zealand; and Aquarium Point,

Otago Harbour, South Island, New Zealand. At

each site, about one hour was devoted to collecting

various living and non-living substrata from

sheltered habitats under rocks or overhangs. These

substrata were then examined under a

stereomicroscope to detect pedicellinid colonies.

This two step method of searching revealed that at

each site pedicellinids were present on about 5—

10% of the substrata collected.

182 kK. WASSON

SYSTEMATICS

Phylum KAMPTOZOA Cori, 1929

(= Entoprocta Nitsche, 1870)

Order COLONIALES Emschermann, 1972

Sub-Order STOLONATA Emschermann, 1972

Family PEDICELLINIDAE Hincks, 1880

Genus Pedicellina Sars, 1835

Pedicellina whiteleggii Johnston & Walker, 1917

(Figs. 1-4).

? Pedicellina sp. MacGillivray, 1887: 221, no fig.

Pedicellina cernua (Pallas, 1774)

Whitelegge, 1889: 293, no fig.

Pedicellina whiteleggii Johnston & Walker, 1917:

60, fig. 14; Stach 1937: 374, no fig.

Pedicellina hirsuta Jullien, 1891 sensu Johnston

& Angel, 1940: 227, figs. 38-42.

Pedicellina hispida (from New Zealand) Ryland,

rose IO, sis, 5,

Gordon, 1972: 510, fig. 3.

SENSU

non Pedicellina cernua (Pallas, 1774):57, fig. 10

in plate 4.

non Pedicellina hirsuta Jullien, 1891:13, no fig.

non Pedicellina hispida (from Europe) Ryland,

1965:200, fig. 8.

Etymology

Johnston & Walker (1917) named this species

Pedicellina whiteleggii to honour Thomas

Whitelegge, who had earlier collected colonies of

the same species from the Port Jackson area.

Whitelegge in 1883 became the first cataloguer of

marine invertebrates at the Australian Musuem,

and held that position until 1908. Whitelegge

contributed greatly to the characterisation of the

invertebrate fauna of the region, and was one of

few Australians to report the presence of

kamptozoan species on these seashores.

Type

Johnston & Walker provided two syntypes

(SAM E942 and E943) of this species. I now

designate SAM E942 as the lectotype; SAM E943

thereby becomes a paralectotype.

Type Locality

Johnston & Walker (1917) indicate that the

types of Pedicellina whiteleggii were collected

under stones in the intertidal zone at Port Jackson,

NSW. They list both Middle Harbour and

Watson’s Bay as collection sites, and do not

specify at which of these two localities the types

were taken.

Material Examined

South Australia: Port Willunga, intertidal,

25.111.1944, SAM L709; Outer Harbour, 0.5 m,

R.G. Chittleborough, 2.v.1951, SAM L710; Point

Turton Jetty, Yorke Peninsula, 3-4 m, K. Gowlett-

Holmes, 2.iv.1994, SAM L711; Aldinga Reef,

Gulf St. Vincent, 1-2 m, S.A. Shepherd & K.

Wasson, 18.11.1995, SAM L712; Aldinga Reef,

Gulf St. Vincent, 1-2 m, S.A. Shepherd & K.

Wasson, 18.ii.1995, personal collection of K.

Wasson.

New South Wales: Port Jackson, intertidal, T.

H. Johnston & M. J. Walker, SAM E942 & E943

(syntypes); Port Jackson, intertidal, T. H. Johnston

& L. M. Angel, SAM BANZARE collection; Port

Stephens, intertidal, T. H. Johnston, SAM L708;

Port Stephens, intertidal, AM U672 & U673;

Rose Bay, Port Jackson, AM U880; Long Reef,

Sydney, intertidal, P.A. Hutchings & W.F.

Ponder, 16.xi.1970, AM W22254; Port Kembla

Harbour, 1 m, J. Watson, NMV_ F77075;

Fairlight, Port Jackson, intertidal, K. Wasson &

Aust. Mus. Party, 15.ii.1995, AM W2225S.

Victoria: Little Henty Reef, near Apollo Bay,

2.5—8.0 m, C. Handreck, 5.ii.1994, NMV F76892.

North Island, New Zealand: Pukerua Bay,

intertidal, S. O’Shea & K. Wasson, 28.i1.1995,

PML reference collection.

South Island, New Zealand: Cemetery Bay,

Otago Peninsula, intertidal, E. Batham, 4.v.1961,

PML reference collection (P. hispida holotype);

Cemetery Bay, Otago Peninsula, intertidal, E.

Batham, 4.v.1961, BMNH 1964.2.8.5 (P. hispida

holotype); Cemetery Bay, Otago Peninsula,

intertidal, E. Batham, 4.v.1961, BMNH

1964.2.8.25; Cemetery Bay, Otago Peninsula,

intertidal, E. Batham, 10.xii.1962, OM Iv2010/

A.64:9 (P. hispida paratype); Cemetery Bay,

Otago Peninsula, intertidal, E. Batham,

10.x11.1962, BMNH 1964.2.8.6; Aquarium Point,

Otago Peninsula, intertidal, M. Barker & K.

Wasson, 4.ii.1995, PML reference collection;

Aquarium Point, Otago Peninsula, intertidal, M.

Barker & K. Wasson, 4.ii.1995, personal

collection of K. Wasson.

Diagnosis

Stolon narrower than stalk; stolonic septa of

variable thickness, but often very delicate; stalk

quite thick, about three times as long as calyx;

PEDICELLINA WHITELEGGII 133

stalk and calyx hispid; calyx wide and somewhat

asymmetrical in side view; 16-36 tentacles,

usually 20-24; tentacular membrane high; axial

rows of large cells on abfrontal surface of

tentacles, conspicuous as glistening bands in

living zooids; larva small, tall and narrow, often

densely coated with particles.

Description

Colony: A colony of Pedicellina whiteleggii

resembles those of most other pedicellinids, with

a network of stolons creeping on the substratum,

from which the zooids arise at regular intervals

(Fig. la). The stolons, stalks and calyces are

translucent beige. New buds are formed at the

FIGURE 1. Pedicellina whiteleggii. a. part of a colony. Scale bar=1000 um. b. zooid with a contracted male calyx in

frontal view. Scale bar=500 pm. c. expanded female calyx in side view, with brooded larvae. The conspicuous rows

of cells are shown on two tentacles only. Scale bar=100 um. d. particle-coated larva. Scale bar=20 um.

134 K. WASSON

base of older zooids, and remain connected by a

basal stolon. Calyces are deciduous, as in other

stolonate kamptozoans, and at any time about

10% of the zooids in a colony are in the process of

regenerating calyces that have been shed.

FIGURE 2. Two Pedicellina whiteleggii zooids (from

Long Reef, NSW). Calyces are contracted and fixed, in

side view. scale bar=200 yum.

Stolon: The stolon is always considerably

narrower than the upright stalk (Table 1). The

inter-zooidal stolonic distance varies within

populations, but is often quite short, resulting in a

high density of zooids (20-30 zooids/cm?) on the

substratum.

The septa which delineate stolon segments

bearing zooids (“‘fertile” segments) from segments

without zooids (“sterile” segments) are often

unusually delicate and are sometimes so

inconspicuous that they cannot be distinguished

even by careful examination of the appropriate

portion of the stolon. The stolonic septa in P.

whiteleggii appear to vary in thickness even

within the same colony, ranging from distinct,

fairly strong septa to extremely faint or even

absent septa. This sort of septal variation is not

typical of other pedicellinid species, in which the

septa are either always present and conspicuous,

as in P. cernua, or always absent, as in P.

pyriformis.

Stalk: The stalk in P. whiteleggii is thicker and

sturdier than those of many other pedicellinid

species (Figs. 1b; 2). As in other pedicellinids, the

stalk is highly muscular, and living colonies are

characterised by the active bending motions of the

zooids’ stalks. Contracted zooids, which have

been preserved unrelaxed, sometimes appear to

have annulate stalks.

Stalk length varies within a population, but

typically the stalk is about 1.2-1.6 mm long,

which is about three times as long as the calyx is

high (Table 1). The stalk tapers in width from

base to apex (Table 1). The stalk is invariably

hispid, ornamented everywhere with cuticular

spines (Figs. 1b; 2). The spines are often hook-

shaped, down-curved with broad bases and

narrow tips. Spine size and density vary within a

population; some stalks may be sparsely covered

with small spines while others are densely covered

with long spines.

Calyx: The calyx is laterally compressed, being

much wider in side view (Fig. Ic) than in anterior

(Fig. 1b) or posterior view (Table 1). In side view,

the calyx is broad and somewhat asymmetrical

due to a slight aboral bulge. This bulge and the

resulting asymmetry are variable within a

population, and occasionally are rather

pronounced.

The calyx is invariably hispid. While the size

and density of calycal spines vary within

populations, they are generally similar to those on

the zooid’s stalk.

The tentacles are extended directly above the

calyx parallel to the stalk (Fig. lc) rather than

PEDICELLINA WHITELEGGII 3S)

FIGURE 3. Pedicellina whiteleggii calyces. a. contracted, fixed, female calyx (from Rose Bay, NSW) in side view.

b. contracted, live, immature calyx (from Aldinga Reef, SA) in side view. ¢. semi-contracted, live, male calyx (from

Portobello, NZ) in frontal view. d. semi-contracted, live, immature calyx (from Fairlight, NSW) in side view. Scale

bar=100 pm in a—d.

136 K. WASSON

TABLE 1. Dimensions of zooidal components of Pedicellina whiteleggii. All measurements are given in micrometers.

Dimension Average (adult) Observed Range

CALYX HEIGHT 600 400-760

(calyx base to tentacular membrane)

CALYX DEPTH 500 300-600

(in side view)

CALYX WIDTH 350 220-450

(in anterior or posterior view)

TENTACULAR MEMBRANE HEIGHT i 60-85

(region of conspicuous circular musculature)

TENTACLE NUMBER 20-24 16-36

CALYX HEIGHT/STALK LENGTH sles) 1:2-1:4

STALK LENGTH 1400 720-2600

(stolon to base of calyx)

APICAL STALK WIDTH 130 110-180

(at calyx)

MID-STALK WIDTH 160 120-280

(in middle)

BASAL STALK WIDTH 240 160-340

(just above stolon)

STOLON WIDTH 90 60-120

FERTILE SEGMENT LENGTH 400 240-600

STERILE SEGMENT LENGTH 300 100-600

LARVAL HEIGHT 150 130-200

(prototroch to apical organ)

LARVAL WIDTH 16) 60-100

(in side view)

tilted anteriorly as in some other pedicellinid

species. The tentacular membrane is high, and in

contracted calyces the tentacles appear to be

deeply infolded. When living colonies are

examined under reflected light, conspicuous pale

yellow axial bands glisten on the tentacles. At

higher magnification these bands appear to consist

of two longitudinal rows of large, oval cells, on

the abfrontal surface of each tentacle. (Figs. Ic;

3b; 3c; 3d). These large cells may have a

glandular function. This tentacular organisation is

an unusual and distinctive feature of P. whiteleggii

which has not been previously recorded.

The shape and size of the digestive tract and its

components vary somewhat with state of

anaesthetisation. The mouth leads into a typical

wide oral funnel which narrows into a tubular

esophagus. The stomach is large, often somewhat

triangular in appearance, wider on top than near

the base (Fig. 3a). The intestine is a broad cylinder

which narrows abruptly at the rectum. The rectum

is quite broad, and sometimes widens towards the

anus. The anal cone often extends obliquely, but

may be folded down horizontally in a contracted

calyx or extended up vertically in a fully relaxed

animal.

Sexual Reproduction: Sexual reproduction

probably occurs year-round. In very mature

colonies, 80-90% of calyces have testes or

ovaries. Since colony boundaries could not be

determined, it was impossible to determine

whether colonies are gonochoric or

hermaphroditic. Calyces contain gonads of only

one sex, and thus appear to be gonochoric,

although the possibility of sequential

hermaphroditism cannot be excluded. Testes are

similar to those of other kamptozoans in

appearance and location: large white sacs packed

with sperm, lying on either side above the

stomach (Fig. 1b). Females have relatively large

PEDICELLINA WHITELEGGII 137

ovaries for a kamptozoan, located in the same

position as the testes (Fig. Ic). About 12-16

embryos are clustered in a thin-walled, flexible

brood chamber. The embryos seem unusually

small and numerous.

The larva is relatively small (Table 1), tall and

very narrow in frontal and side view (Figs. Id; 4a;

4b). The surface of the swimming larva is densely

coated with particles of apparently external origin

which appear beige under reflected light and dark

under transmitted light. Particles sometimes cover

even larvae which are still retained in the brood

chamber. Perhaps sticky secrections coat the larval

surface, attracting and retaining particles. The

outline of the prototroch is roughly circular when

viewed from below. The larval foot and frontal

organ are highly reduced or absent. As in other

FIGURE 4. Pedicellina whiteleggii larvae. a. larva (from

Portobello, NZ), probably in frontal view. b. larva (from

Aldinga, SA), probably in side view. Scale bar=100 pm

in a and b.

kamptozoans, the larval body is quite contractile.

The form of the larva in Pedicellina whiteleggii

is very distinctive and differs from that of other

pedicellinids in which the larva is knowa. In both

P. cernua and P. nutans the larva is much larger

and much wider in side view, and has a well-

developed foot and frontal organ (Nielsen 1971).

While the larva of P. whiteleggii differs markedly

from those of some of its congeners, it bears a

striking resemblance in its proportions and in the

dense particulate covering to the ‘type A’ larva

described for the barentsiid Barentsia gracilis

from European waters (Nielsen 1971). The B.

gracilis ‘type A’ larva also lacks a foot and has a

reduced frontal organ.

Newly settled larvae are found on or near adult

colonies, suggesting that in this species, as in

many or all other kamptozoans, larval settlement

is gregarious.

Habitat

Colonies of Pedicellina whiteleggii generally

occur on living substrata, including algal

holdfasts, sponges, hydroids, serpulid polychaete

tubes, oyster shells, erect and encrusting bryozoa,

and ascidians. The most common hosts seem to

be serpulids, encrusting bryozoans, and solitary

ascidians. Occasionally, colonies may also grow

directly on protected rock surfaces.

This pedicellinid occurs in sheltered places,

such as under stones or in deep overhangs, and

grows in fouling communities of hydroids,

ascidians, and bryozoans. The material examined

for this study was collected primarily from the low

intertidal zone, with only a few specimens

collected subtidally at a few meters depth. This

species’ distribution may well extend into far

deeper water; it should be sought by examination

of appropriate substrata collected by SCUBA or

dredging.

Distribution

South Australia (Gulf St. Vincent; Yorke

Peninsula), New South Wales (Port Jackson; Port

Stephens; Port Kembla), Victoria (Lady Julia

Percy Island; Little Henty Reef), and New Zealand

(Goat Island Bay; Pukerua Bay; Otago Peninsula).

Pedicellina whiteleggii and its synonyms

Three reports of pedicellinids from New South

Wales and Victoria roughly match the description

of P. whiteleggii, but the animals are not well

described or figured in these texts. MacGillivray

(1887) merely noted Pedicellina sp. from Port

Phillip Heads without providing descriptive

information or a figure; but since P. whiteleggii is

138 K. WASSON

common at that locality, his report may well be of

this species. Whitelegge (1889) listed Pedicellina

cernua from Sydney Harbour, again without a

description or figure, but Johnston & Walker

(1917) synonymised his material with P.

whiteleggii. Stach (1937) identified P. whiteleggii

from Lady Julia Percy Island off the Victorian

coast, and since Johnston & Angel (1940)

examined other collections he had made and found

his identifications of P. whiteleggii correct, I also

infer that his material was indeed P. whiteleggit.

Ryland (1965) described a new species,

Pedicellina hispida, from New Zealand, based on

three colonies growing on brachiopods in Otago

Harbour, New Zealand. Examination of the types

of P. hispida revealed that it closely matches P.

whiteleggii: a narrow stolon with inconspicuous

septa, hispid stalk and calyx, somewhat

asymmetrical calyx with a high tentacular

membrane. The sizes and proportions of the

zooids also agree with those of P. whiteleggii.

Live colonies I collected from Otago Harbour were

clearly the same as Ryland’s P. hispida based on

zooid structure and calyx morphology. In the fresh

material, I noted the rows of large tentacular cells

glistening yellow under reflected light, and the

tall, particle-covered larva, both characteristic of

P. whiteleggii. These traits, taken together, justify

the synonymy of Ryland’s P. hispida from New

Zealand with P. whiteleggii. Gordon (1972)

identified a kamptozoan from the North Island of

New Zealand as P. hispida Ryland. Based on the

figure, this kamptozoan was probably also P.

whiteleggii.

Ryland (1965) also identified colonies from

Europe as P. hispida, but the calyces in these

colonies are glabrous, while those of P.

whiteleggii are always hispid. The tentacle and

larval structure of the European colonies is not

known, and so their status is uncertain. Because

of this uncertainty and because their localities are

so distant from Australia and New Zealand, I have

excluded the European specimens of P. hispida

from the list of synonyms of P. whiteleggii.

In 1940, Johnston & Angel synonymised P.

whiteleggii Johnston & Walker with P. hirsuta

Jullien. This decision reflected a general trend

then to consider all the world’s pedicellinids as

variations of one species, Pedicellina cernua

Pallas. Cori (1936) suggested that P. whiteleggii

Johnston & Walker might be P. cernua var.

hirsuta, described by Jullien (1891) as P. hirsuta

from Tierra del Fuego. Although Cori had

apparently never seen either Jullien’s or

Johnston’s material, Johnston & Angel (1940)

were nevertheless swayed by Cori’s authoritative

opinion. While they did not believe P. whiteleggii

was a variety of P. cernua, they compromised by

synonymizing it with P. hirsuta. Jullien’s (1891)

description of P. hirsuta is sketchy and there are

no illustrations. The type material (Muséum

National d’Histoire Naturelle, Paris) consists of

five dried zooids on natural substratum (Bry—

2177) and a slide mount of five other zooids (Bry—

39), of which four are recently budded, tiny zooids

at the stolon tip. My examination of this type

material suggested that this species, while

resembling P. whiteleggii in general shape and in

the hispid stalk and calyx, is distinct from it.

Based on these ten poorly preserved zooids, P.

hirsuta has a wider stolon that is hispid in places,

which contrasts with the narrow, glabrous stolon

of P. whiteleggii. The stalk tapers less in P.

hirsuta than in P. whiteleggii, and the spines on

the P. hirsuta types are longer and denser than is

typical for P. whiteleggii. Most conclusively, the

tentacles of the type specimens did not contain the

conspicuous large cells found in P. whiteleggii.

The larval form of P. hirsuta remains unknown,

because the only sexual mature zooids in the type

specimens were male. Since there is no evidence

for synonymy with P. hirsuta Jullien, P.

whiteleggii now reverts back to its original 1917

name.

Other pedicellinids from Australia and New

Zealand

Pedicellina compacta Harmer, 1915: This

species was reported from the Great Barrier Reef

by Hastings (1932). Her material (BM

1932.4.20.93) and additional colonies I collected

in Rowes Bay, Townsville, Queensland (MTQ

G21264) are certainly referable to Harmer’s P.

compacta (BM 1916.8.23.33-34) from the Aru

Islands (Indonesia). P. compacta apparently

replaces P. whiteleggii as the ubiquitous coastal

species in tropical and sub-tropical waters. P.

compacta has much smaller zooids than P.

whiteleggii, with total zooid height (stalk and

calyx together) averaging 600-800 pum. The stolon

is extremely narrow (30-40 pm) and the stolonic

septa are much more conspicuous than in P.

whiteleggii. The stalk tapers much less between

base and apex, and is proportionately shorter,

typically one to two times as long as the calyx is

high. The stalk is very muscular and mobile. The

proportionately big calyx (but still only about 300

um high!) is narrower in side view than in P.

whiteleggii; it is not as compressed laterally. The

calyx and tentacles show a slight anterior tilt

PEDICELLINA WHITELEGGII 139

absent in P. whiteleggii, and there are only about

12-16 tentacles. The calyx and stalk are covered

by filiform, cylindrical, extremely long spines that

are completely different from the shorter, thicker,

hook-like spines of P. whiteleggii.

Pedicellina pyriformis Ryland, 1965: This

exquisite species forms the tallest and densest

colonies known for any pedicellinid. It was

described by Ryland (1965) from the Otago

Peninsula. I have examined his material (PML

reference collection; OM Iv2008—9/A.64:7-8) as

well as a colony collected by K. Gowlett-Holmes

(SAM PH 0008) from Tasmania. The stolon is

very much wider than in P. whiteleggii and is

completely non-septate. The stalk is listed by

Ryland as being about 2 mm long, but a colony

from Otago deposited at PML after his study was

completed has stalks reaching an astonishing 5-6

mm. The stalk tapers only very slightly from base

to apex, and both calyx and stalk are glabrous.

The triangular calyx is much less laterally

compressed than in P. whiteleggii, and is not

asymmetrical in side view. The calyx is typically

600-700 um high, but in some cases attains a

height of 1 mm. The gut is also distinctive: the

stomach is compact and triangular, and the

intestine and the rectum are long and narrow, the

latter extending within an anal cone high into the

tentacular crown. The gut has empty spaces

around it rather than filling the calyx as in most

other pedicellinid species.

Pedicellina grandis Ryland, 1965: This

species was described from one colony from

Otago Harbour. This colony (PML reference

collection; OM A.64:6) resembles P. whiteleggii

in general form. The stolon is narrow and the

septa inconspicuous; the stalk and calyx are

hispid. The main difference between P. grandis

and P. whiteleggii appears to be quantitative:

some P. grandis zooids have considerably longer

stalks than do P. whiteleggii zooids. Ryland

(1965) also noted that the tentacles in semi—

contracted calyces form a distinctive conical cap,

but this trait appears to vary with the state of

zooidal relaxation. Further examination of the

tentacle structure and larval form of this species

may reveal further distinguishing features which

separate the two species, or, on the other hand,

that P. grandis is synonymous with P. whiteleggii,

and that its longer stalks can be attributed to

habitat or age differences.

Pedicellina pernae Ryland, 1965: This species

from Otago is distinguished from P. whiteleggii

by its somewhat smaller size and a glabrous or

only sparsely hispid calyx. Ryland (1965) noted

that the calyx was more asymmetrical than that of

P. hispida (=P. whiteleggii), but my examination

of the types (PML reference collection; OM

Iv2011/A.64:10) revealed that the calyx shape

falls within the range of variation of P.

whiteleggii. Ryland (1965) himself noted that this

species lacks really distinctive features. Collection

of living material and examination of tentacle

structure and larval form could reveal whether this

is indeed a different species than P. whiteleggii.

Pedicellina cernua (Pallas, 1774): This

cosmopolitan species occurs mainly in bays and

harbours. Kirkpatrick (1890) found this species in

Port Phillip, Victoria. Chittleborough (1952)

reported P. cernua from a community of primarily

introduced species at Port Adelaide and Outer

Harbour, South Australia, and it is probably

present in other harbours in Australia and New

Zealand. Kirkpatrick’s (BM 1888.5.17.24) and

especially Chittleborough’s material (SAM L713—

718) have many features which distinguish the

cosmopolitan P. cernua from the indigent P.

whiteleggii. The stolonic septa in P. cernua are

more clearly visible. The stalk is narrower and

tapers more gradually from base to apex. The stalk

is hispid, and the spines longer than in P.

whiteleggii, although this feature is variable. The

calyx of P. cernua is wider in side view, almost as

wide as high, and its aboral bulge and consequent

asymmetry are more pronounced. The tentacles in

P. cernua are tilted anteriorly. The tentacles lack

axial rows of large cells, although there is dark

green granular pigmentation in the tentacles of

some zooids. The tentacular membrane is not so

high as in P. whiteleggii. A more striking

difference is the consistent absence of spines from

the calyx. The rectum in P. cernua generally

appears narrower and extends higher into the

tentacular crown than in P. whiteleggii. The brood

chamber of P. cernua is lobulate, while that of P.

whiteleggii is not. As already discussed, the larva

of P. cernua differs from that of P. whiteleggii in

being larger, wider in side view, in having a well—

developed foot and frontal organ, and in lacking a

dense particulate covering (Nielsen 1971).

DISCUSSION

Knowledge of pedicellinids from Australia and

New Zealand is rather limited. This is certainly

not because the animals are rare or hard to find;

only a few hours in the field turned up many

colonies at every site I visited.

The diversity of Australian and New Zealand

pedicellinids is low. There appears to be one

140 K. WASSON

ubiquitous coastal pedicellinid (P. whiteleggii ) in

colder waters and another (P. compacta) in

warmer regions, one cosmopolitan species in

harbours (P. cernua), and one other distinctive

species (P. pyriformis) in southern areas. Other

unreported and perhaps undescribed species will

surely be found in unexplored habitats (e.g. deeper

water) or unsurveyed regions (e.g. Western

Australia). In Northern Europe, which has been

much better surveyed for kamptozoans, there is

also a low diversity of pedicellinids; only one or

two common species and a few rare ones. It is

difficult to compare the species diversity of

pedicellinids from Australia with other regions,

since the pedicellinid fauna of most parts of the

world has not been characterised.

The distributions of the pedicellinids found in

Australia and New Zealand seem fairly large.

Pedicellina pyriformis, known only from Otago

and Tasmania, appears to have the most limited

range, but this rare species may yet be found in

other areas. The distribution of P. whiteleggii

includes much of New Zealand and southern

Australia, but its western and southern limits are

not known. P. compacta is found from Indonesia

to Queensland. And P. cernua has been reported

from all over the world. Only more thorough

taxonomic surveys will delineate these ranges

more adequately.

The taxonomy of pedicellinids (and of tiny,

soft—bodied creatures in general) always poses

challenges because of the apparent paucity of

morphological characters. Historically,

kamptozoan taxonomy has been based on adult

traits. Larval form has been largely ignored (but

see Nielsen 1971), although since all kamptozoans

brood, and most are reproductive year—round,

larvae are usually easy to obtain. Larvae can also

be examined in well-preserved museum

specimens. In this study, a distinctive larval form

enabled museum and the field material in

Australia and New Zealand to be united under

one name, and helped to separate Pedicellina

whiteleggii from P. cernua. In the future,

pedicellinid taxonomy would be strengthened by

inclusion of larval traits in species descriptions.

ACKNOWLEDGMENTS

I am grateful to P. Arnold, E. & M. Barker, P.

Berents, K. L. Gowlett-Holmes, J.-L. d’Hondt, S.

O’Shea, S. A. Shepherd, M. Spencer Jones and W.

Zeidler for their generous assistance in accessing

museum collections and in carrying out field work. J. S.

Ryland’s excellent study of pedicellinids from New

Zealand was a resource and an inspiration for this work.

I am indebted to C. Nielsen for drawing my attention to

the usefulness of tentacle structure and larval form in

kamptozoan taxonomy. A. T. Newberry and D. Potts

provided invaluable assistance in the preparation of the

manuscript. Thanks also to C. Glasby and the Australian

Nature Conservation Agency for support of this research.

REFERENCES

CHITTLEBOROUGH, R. G. 1952. Marine Fouling at

Port Adelaide. Master of Science Thesis, University

of Adelaide.

CORI, C. I. 1929. Kamptozoa. Pp. 1-64 in ‘Handbuch

der Zoologie’, Volume 2(5). Ed. W. Kiikenthal & T.

Krumbach. W. de Gruyter: Berlin.

CORI, C. I. 1936. Kamptozoa. Pp. 1-119 in ‘Klassen

und Ordnungen des Tierreichs’, Volume 4, Part 2,

Book 4. Ed. H. G. Bronn. Akademische

Verlagsgesellschaft: Leipzig.

EMSCHERMANN, P. 1972. Loxokalypus socialis gen.

et sp. nov. (Kamptozoa, Loxokalypodidae fam. nov.),

ein neuer Kamptozoentyp aus dem nérdlichen

Pazifischen Ozean. Ein Vorschlag zur Neufassung der

Kamptozoensystematik. Marine Biology 12(3): 237-

254.

GORDON, D. P. 1972. Biological relationships of an

intertidal bryozoan population. Journal of Natural

History 6: 503-514.

HARMER, S.F. 1915. The Polyzoa of the Siboga

Expedition. Part I. Entoprocta, Ctenostomata and

Cyclostomata. Siboga Expedition Reports 28A: 1-

180.

HASTINGS, A. 1932. The Polyzoa. Great Barrier Reef

Expedition, 1928-1929, Scientific Reports 4: 399-

458.

HINCKS, T. 1880. ‘A History of the British Marine

Polyzoa’. Van Voorst: London.

JOHNSTON, T. H. & WALKER, M. J. 1917. A new

species of Pedicellina from Sydney Harbour.

Proceedings of the Royal Society of Queensland

29(5): 60-63.

JOHNSTON, T. H. & ANGEL, L. M. 1940. Endoprocta.

B.A.N.Z. Antarctic Research Expedition 1929-1931.

Reports (Series B; Zoology and Botany) 4(7): 215-

Bil.

JULLIEN, J. 1891. Bryozoaires. Mission Scientifique du

Cap Horn (Zoologie) 6(3): 1-92.

KIRKPATRICK, R. 1890. Polyzoa from Port Phillip.

Annals and Magazine of Natural History 6(2): 12—

PEDICELLINA WHITELEGGII 141

MACGILLIVRAY, P. H. 1887. A catalogue of the

marine Polyzoa of Victoria. Proceedings of the Royal

Society of Victoria 23: 187-224.

NIELSEN, C. 1971. Entoproct life-cycles and the

entoproct/ectoproct relationship. Ophelia 9(2): 209-

341.

NITSCHE, H. 1870. Beitrage zur Kenntniss der

Bryozoen. Zeitschrift fiir wissenschaftliche Zoologie

20: 1-36. i

PALLAS, P. S. 1774. ‘Naturgeschichte merkwiirdiger

Thiere’. Zoophyta, pp. 52-63 and plate 4. G. A.

Lange: Berlin.

RYLAND, J. S. 1965. Some New Zealand Pedicellinidae

(Entoprocta), and a species new to Europe.

Transactions of the Royal Society of New Zealand

(Zoology) 6(19): 189-205.

SARS, M. 1835. ‘Beskrivelser og Iagttagelser over nogle

maerkelige eller nye i Havet ved den Bergenske Kyst

levende Dyr af Polypernes, Acalephernes,

Radiaternes, Annelidernes og Molluskernes Classer’.

T. Hallager: Bergen.

STACH, L. W. 1937. Bryozoa. Lady Julia Percy Island.

Proceedings of the Royal Society of Victoria 49(2):

374-385.

WHITELEGGE, T. 1889. List of the marine and fresh-

water invertebrate fauna of Port Jackson and the

neighbourhood. Journal and Proceedings of the

Royal Society of New South Wales. 23: 163-323.

MYTH AS HISTORY? THE NGURUNDERI DREAMING OF THE LOWER MURRAY,

SOUTH AUSTRALIA

PHILIP A. CLARKE

CLARKE, P. A. 1995. Myth as History? The Ngurunderi Dreaming of the Lower Murray,

South Australia. Records of the South Australian Museum 28(2): 143-157.

The ethnographic record of Aboriginal mythology in the Lower Murray cultural region of

South Australia provides a number of accounts which describe the activities of the main

‘Dreaming’ ancestor, Ngurunderi. Rather than attempting to standardise the cultural data, this

diversity is used here to examine Aboriginal perceptions of the landscape that reflect differing

world views. Although the literature acknowledges the important religious dimension of these

creation myths, the socio-political dynamics have hitherto generally been dismissed. This paper

demonstrates that it is part of the essential nature of mythology, and its associated site-related

data, to be flexible and constantly altered and appended. Here is a study of cultural geography,

which considers both the material and non-material aspects of Aboriginal cultural construction

of the landscape.

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

South Australia 5000. Manuscript received 8 November, 1994.

INTRODUCTION

In pre-European Australia, Aboriginal people

perceived the social and physical aspects of their

world to be closely interwoven. They believed that

their spirit ancestors had imbued the landscape

with social relevance, and had thereby humanised

it. The southern Australian ethnographic literature

generally distinguishes between myths that

focused on events said to have occurred during the

‘Dreaming’ or ‘Dreamtime’, when the main

‘creation’ took place, and those that concern the

period afterwards. The ‘Dreaming’ in the Lower

Murray region was referred to as the Kulhal or

Gulal (Berndt 1940: 170; Berndt & Berndt 1981:

229). Ronald and Catherine Berndt record that the

Kalalwu or Kalalal reportedly meant ‘long ago’

(Berndt & Berndt 1993: 75, 242). Mythology

gained from enlightenment through actual dreams

was termed pekeri (Berndt & Berndt 1993: 213,

214). The Dreamtime represents an Aboriginal

English gloss of a range of meanings. The

‘Dreaming’ can loosely be defined as the whole

body of mythology in Aboriginal Australia that

provides some insight into significant cultural

events. The form of ‘history’ that the Dreaming

provides is generally portrayed in the popular

literature as a monolithic entity, without the

possibility of revision or reinterpretation. Under

this model, adhered to by both Aboriginal people

and some early scholars, such history (or

‘tradition’) is not changed, only forgotten. Here I

follow the work of Kolig (1984 [1981]), Myers

(1986), and Sutton (1988), and take a more

flexible view of Aboriginal Dreaming, showing it

to be a text that reflects the dynamism of culture.

This paper follows the argument put forward in

a previous work that mythology provides an image

of the dynamic aspects of cultural relationships to

landscape (Clarke 1991a: 66-69). Here, the

development of tradition is treated as a constant

process of re-evaluation of the links between

present and past. In this context, I begin by

considering the variations occurring within a

major creation myth, that of Ngurunderi. As

discussed by Maddock (1976) in reference to

myths in northern Australia, the local variations

form a universe of discourse in which their owners

collectively determine what is in common to them

all, and through which new possibilities are

introduced. The differing versions are not just

answers to varied life situations, but are put forth

by Aboriginal people as responses to each other. I

demonstrate how this process continued in the

Lower Murray, under the new socio-political

dynamic of post-European colonisation, creating a

proliferation of ‘new’ versions of important myths.

The cultural geography concept of the ‘cultural

landscape’ considers the physical and perceptual

elements of landscape that are absorbed by other,

P. A. CLARKE

S ES A F

S “gMuRRAY BRIDGE

Oo ~

= oe

WS S

A S TAILEM BEND

c \

oe RAPID BAY, a \

—_ — “S \

CAPE Le

JERVIS MENINGIE

KANGAROO Hi dec 7 \

En MURRAY

ISLAND are

Uy, MOUTH

SOUTHERN

OCEAN

KINGSTON

\ |

| NORTHERN !

| TERRITORY |

| QUEENSLAND

WESTERN |

AUSTRALIA | ee

South: “oie ee sane

| AUSTRALIA

Above map | VICTORIA

Figure 1. The Lower Murray cultural region.

MYTH AS HISTORY?

usually successive, cultural groups. It is useful

here to provide a framework of evaluating

continuity.

The Lower Murray cultural region may be

defined as the area bounded by Rapid Bay in the

west, across to the southern side of Murray

Bridge, and south to Kingston (Fig.1). It takes in

the southern part of the Fleurieu Peninsula, all of

Encounter Bay, Lake Alexandrina and Lake

Albert, and the Coorong. In the early years of

European settlement this region was noted by

several observers to be relatively culturally

homogeneous in relation to neighbouring groups

that differed in language, custom and material

culture (Meyer 1843: vi; 1846 [1879: 185];

Cawthorne 1844 [1926: 1]; Mathews 1898: 336—

343). Rather than comprising a few large units

that past scholars have labelled as ‘tribes’, it is

better to treat the Lower Murray people in the

early years of European settlement as a more

numerous series of extended family units, headed

by several politically active people who led by

their coercive powers over others (Clarke 1994,

Chapter 2). These land-owning units were descent

groups tracing their origins to particular locations

of mythological significance in their territory. In

most parts of Australia, these places are usually

termed ‘sacred sites’ (Berndt 1970; Myers 1986:

50, 51, 134-136). An individual’s connection to

the Dreaming provided the basis of his or her

identity. Marriage and trade relationships helped

the Lower Murray groups maintain a high degree

of social and cultural cohesion. Because of a

number of factors, the Ngarrindjeri people have

retained their regional and cultural distinctiveness

up to the present (Jenkin 1979; Hemming 1988;

Hemming et al 1989; Clarke 1994). Due in part to

the close proximity of this region to Adelaide, the

| Aboriginal people from the Lower Murray

continue to have a major role in Aboriginal affairs

in South Australia.

CONTACT BETWEEN EUROPEANS AND ABORIGINAL

PEOPLE

Aboriginal groups in South Australia were

influenced by Europeans long before official

settlement in 1836. The first significant effect of

) European settlers on South Australian Aboriginal

| populations was probably the introduction and

} spread of smallpox from eastern Australia,

reaching the Lower Murray along the river system

in two waves sometime between 1814 and 1820,

and between 1829 and 1831.' In the mythology of

the Lower Murray, smallpox is linked with the

145

ancestral spirit, Kulda, who came from the

Southern Cross constellation foretelling death

(Tindale 1931-34: 251, 252; 1937: 111, 112;

1941: 233, 234). The devastation this brought to

indigenous populations, and the effect upon their

culture, can only be guessed at.

The activities of whalers and sealers, operating

chiefly out of Bass Strait, and their effect upon the

southern Aboriginal populations, warrant a

mention here. These men, some of whom were

dwelling on Kangaroo Island as early as 1819,

lived outside colonial control then based in

Tasmania (Clarke 1990 MS; 1994: 192-203).

They periodically raided the mainland for

Aboriginal women to serve as labourers and

wives. Some of these Kangaroo Island men had

trading relationships with mainland Aboriginal

groups, in particular those in the Lower Murray

region. Therefore it is likely that most southern

South Australian Aboriginal people had some

knowledge of Europeans before official settlement

in 1836, even if only indirectly, and had

rationalised this incursion within their own

cosmological views. With the range of contacts

discussed above, both indirect and direct, we must

acknowledge that the early southern South

Australian ethnographies, however skilfully

obtained, do not provide strictly pre-European

accounts of Aboriginal society.

Since official settlement by Europeans in South

Australia in 1836, Aboriginal people in the

southern agricultural districts have gradually

become marginalised in respect to land. During

the nineteenth century, Aboriginal people in the

Lower Murray worked as fishermen, shepherds,

and as labourers to bring in the harvest (Clarke

1994, Chapter 6). By the early twentieth century,

most were forced into living at mission stations

governed under restrictive legislation. Due

primarily to the break-up of the pastoral stations

in the region at this time, Aboriginal hunting and

gathering activities were chiefly restricted to the

Coorong area where Aboriginal reserves and

vacant crown land still existed (Clarke 1994,

Chapter 8). For this reason, contemporary

knowledge of the landscape retained by

Aboriginal people relates mainly to the southern

parts of the Lower Murray region. A broader

Aboriginal cultural identity formed from the social

environment of enforced interaction between

formerly separate groups from throughout South

Australia. Aboriginal people started identifying

themselves more according to mission settlements,

rather than as territorial-based descent groups.

Since the 1960s, when many of the legal

146 P. A. CLARKE

restrictions were removed, Aboriginal people have

moved into rural town centres in the Lower

Murray, with a large number of Ngarrindjeri

people living in Adelaide (Inglis 1961; Gale 1969,

1972; Gale & Wundersitz 1982; Schwab 1988).

The detailed recording of Aboriginal culture in

the Lower Murray starts with the German

missionary Meyer who was active in the

Encounter Bay region during the late 1830s and

early 1840s (1843; 1846). During the 1870s the

writings of Taplin, based at the mission of Point

McLeay on the southern shore of Lake

Alexandrina, focused the attention of northern

hemisphere scholars upon the Ngarrindjeri.

Nineteenth century missionary records are to some

extent balanced by the recordings of Penney

(1840-43 [1991: 1-107]), Cawthorne (1844

[1926]), Moorhouse (1843 [1990]; 1846), Angas

(1847a; 1847b), and Wyatt (1879). During this

century, various anthropologists such as Brown

(1918), Tindale (1930-52; 1931-34; 1934-37;

1935; 1937; 1938; 1938-56; 1941; 1981; 1987;

Tindale & Mountford 1936; Tindale & Pretty

1980), Harvey (1939; 1943), and the Berndts

(Berndt 1940; Berndt & Berndt 1993) have

studied this Aboriginal culture. The attention that

these scholars have directed towards the Lower

Murray has meant that it is ethnographically one

of the best described regions in southern

Australia.

NGURUNDERI: AN ABORIGINAL DREAMING OF THE

LoweR MurRAY

The mythology of Ngurunderi touches upon

most aspects of Aboriginal life. Preeminent in the

anthropological record of the Lower Murray, he is

a major ‘spirit creator’ credited with shaping the

region’s topography and distinctive culture. The

myth provides an explanation of the landscape,

not only the formation of features such as The

Bluff at Victor Harbor and the course of the

Murray River, but the distribution and

characteristics of particular plant and animal

species. For example, Ngurunderi’s actions were

thought to have resulted in the present distribution

of the pigface plant, an important food resource

(Tindale 1931-1934, vol.1: 186; Berndt 1940:

179; Berndt & Berndt 1993: 226). Another

illustration of his perceived creative powers was

to give the bony bream fish its numerous fine

bones (Education Department of South Australia

1990: 56-58). His body parts are represented in

the landscape; his legs form the Sir Richard and

Younghusband Peninsulas at the Murray Mouth

for instance (Berndt & Berndt 1993: 13; Clarke

1994: 114, 115). Thunder was regarded as

Ngurunderi’s voice, rainbows showed him

urinating (Taplin 1874 [1879: 58]). Customs

attributed to Ngurunderi’s law include the

prohibition of young male initiates eating certain

types of Murray cod, considered to be

‘Ngurunderi’s fish’ (Tindale 1934-1937, vol.2:

39). He and his son Matamai are also credited

with introducing mortuary rites to the Lower

Murray people, involving smoking the dead

(Tindale 1934-1937, vol.2: 51; Berndt 1974: 26,

27; Berndt & Berndt 1993: 227, 228). Through

association with Ngurunderi and Matamai,

desiccated bodies were considered to be sacred

objects, at least until the rites were completed.

Ngurunderi was perceived to have power over

both the living and the dead. Taplin records a

discussion with an Aboriginal man, Captain Jack:

He thinks death to arise from sorcery when caused

by sickness, but to be by the special interposition of

Nurundere when caused by spear or waddy wounds.

He says Nurundere invented the waddy and spear

and boomerang, and wimmera [sic.] and plonggee

{small club] long ago. And it appears that they regard

him as a sort of war-god (Taplin Journals, 20

October 1859).

To die as the result of injuries due to fighting

was considered by the Lower Murray to please

Ngurunderi (Taplin Journals, 19 January 1860). It

was believed that after death, Ngurunderi guided

the souls of Lower Murray people to the ‘Lands to

the West’. Here they would live in huts around

him. The Raminyerar considered that they would

reside in Ngurunderi’s hut, with other descent

groups in neighbouring shelters (Meyer 1846

[1879: 206]).

Episodes of the Ngurunderi creation epic were

acted out during dances. Taplin recorded that one

night:

Two of their songs in particular attracted my

attention. One was called “The Nurundere”’, and is

about God ... [It] began with a low chant as if they

were chanting Latin. However, all through the piece

they say the same words over and over again, then

the chant rose higher and higher with beat of the

tartengk a native drum, then it sank again and the

men’s voices broke in shouting in time to the chant

and brandishing the weapons with tartengk. Then

the shrill treble of the women broke in like an

imploring vociferation in answer to the shouts of the

men. These ceased, and the whole concluded with a

loud chant to the beat of the tartengk and drum. The

latter piece was to slower time, and was very

plaintive and wild (Taplin Journals, 30 June 1859).

The passage of Ngurunderi through the

MYTH AS HISTORY? 147

territories of certain descent groups during the

Dreaming had perceived implications for Lower

Murray marriage relationships (Turner 1980: 6).

This mythology was so much a part of the Lower

Murray cultural system that to a large degree an

individual’s possession of detailed knowledge

concerning it helped locate that person within

particular parts of the region.

From descriptions of the myth that stress

warfare, Ngurunderi appears to have had special

significance to men. Nevertheless, he was of much

wider importance as the creator of Lower Murray

culture. In contrast to more northerly regions the

available accounts of Aboriginal ceremonial life

in the Lower Murray indicate that there was a

large degree of sharing of knowledge between

men and women. Tindale (1934-1937, vol.2: 223)

states that in the Lower Murray some old women

were allowed at the Narambi secret initiation

ceremonies. Women were given a say at major

meetings, and were considered to have had a miwi

(spirit) as strong as a man’s (Berndt & Berndt

1993: 71, 285). The extreme separation of the

male and female realms appears to be a distinctive

characteristic of the Central Australian region.’

Elsewhere, such as in parts of the Lake Eyre

Basin, women participated in many of the same

ceremonies as men.* Similarly, in the Lower

Murray region Aboriginal women had traditional

authority, being the partners of men in tribal lore

(Berndt 1981: 181, 182; 1982: 50). This is not to

say that women and men did not have differing

perspectives on some aspects of cultural

knowledge. Generally, only younger people would

have been excluded from secret-sacred categories

of information.

The southern Aboriginal ethnographies show

considerable variation in mythology, even from

within linguistically and culturally similar areas.

In the South Australian region of the Murray

Basin, there are creation accounts for the river that

variously involve the different mythic heroes

Nurelli, Ngurunderi, Korna and Thukabi.° There

are even major variations within the records of

Nurelli and Ngurunderi, particularly in relation to

sites. In the case of the complex of mythology

associated with Ngurunderi, it has been argued

that the variations were simply accounted for by

each Ngarrindjeri group only knowing in most

detail that segment of the Ngurunderi epic relating

to their own area (Hemming 1988). Contrary to

this approach, which attempts to reconcile cultural

differences, I argue that the diversity of these

beliefs, many in direct opposition to others, can be

explained in terms of the dynamic relationship

people had with the landscape. Local knowledge

generates alternative sites where certain events

were perceived to have taken place, therefore

producing distinct versions.

The existence of many versions of the

Ngurunderi myth provides further evidence that

the ‘tribe’ model of Aboriginal social structure as

used by Tindale in his southern South Australian

ethnographic work, is too simplistic.’ For

example, in his treatment of the Tjilbruke

mythology of the Fleurieu Peninsula, Tindale

selectively garnered segments of myth from

various informants from the Lower Murray and

Adelaide regions into one form recognised by the

“Kaurna tribe’ (Tindale 1987; Clarke 1991a). But

by doing this he effectively produced yet another

variation, one that would never have been elicited

from a single Aboriginal source. It is more

productive to treat this variation in the

ethnography as a pre-European characteristic of

the body of knowledge that ‘explained’ the world.

In spite of the apparently confusing number of

creation accounts, there are some common

elements of the mythology within the Lower

Murray region that distinguish it culturally from

surrounding regions.

For the Aboriginal groups of the Lower Murray,

the main creator of the landscape is generally

recorded in the ethnographic literature as

Ngurunderi, or one of the many linguistic

variations of the term.* The different versions of

the Ngurunderi mythology generally fall into two

categories — those that have a coastal bias, and

those that emphasise the inland aspect of the

Murray River.’ The former versions were chiefly

recorded from coastal groups away from the river

in the western and south-western side of the

Lower Murray, the latter from Lower Lakes people

on the north-eastern end near the entrance of the

Murray River into Lake Alexandrina. I will now

describe this variation.

Amongst the earliest recorded myths in

southern South Australia was a Ngurunderi

account recorded by Penney in 1844 from the

Encounter Bay people, concerning the formation

of the Lower Murray landscape.'° In this version,

Ooroondooil (= Ngurunderi) was the first great

spirit to wake. Three or four other beings later

woke, some complaining that they were hungry

and cold. Ooroondooil told one spirit to make a

fish, and he taught them how to cook it. He made

the lesser spirits go off to collect firewood, water

and other necessary items. Ooroondooil then sent

the spirits away to lands he was creating.

Ooroondooil himself went westwards where he

148 P. A. CLARKE

first made the ‘Big Murray people’ (Coorong

groups), and then the groups further west. After

teaching the Lower Murray people their customs,

he left, swimming to other lands in the west. Two

of his wives drowned while trying to follow him,

becoming the Pages Islands of the Backstairs

Passage. Ooroondooil created Kangaroo Island,

and then went further west where he was believed

to have still lived. In a variation of this account,

Ooroondooil had three wives who drowned when,

due to their curiosity, they tried to reached

Kangaroo Island from the mainland."

In another Encounter Bay version of the

Ngurunderi mythology recorded by Meyer from

the Raminyerar descent group, several important

mythological events had already taken place

before his arrival (Meyer 1846 [1879: 205, 206]).

In this rendering Ngurunderi was a large and

powerful man with two wives and several

children. On one occasion his wives fled from

him. He chased them along the southern Fleurieu

Peninsula coastline and while doing so created

many of the geographic features there. He finally

caught the women and beat them, but they

escaped again. This time he tired of chasing them

and ordered the sea to flow and drown them. They

became rocks a few metres from the shore that

can be seen at low tide.'? After he had transformed

the landscape, Ngurunderi was said to have gone

west. One of Ngurunderi’s sons who was

accidentally left behind found his way to the land

‘towards the west’ by catching hold of a line

thrown by Ngurunderi, which was attached to his

testicles. In this account, the creation of new

rivers, hills and other features ceased after

Ngurunderi left the Lower Murray.

The versions of Ngurunderi recorded from the

Lake Alexandrina descent groups vary in much of

the detail from those of Encounter Bay. In one

account from the Lake area obtained by George

Taplin, there were once three great hunters,

Ngurunderi, Nepeli and Waiyungari.'3 As

evidence for their hunting prowess, numerous salt

lagoons around Lake Alexandrina were considered

to have been created by Nepeli and Waiyungari

while pegging out fresh kangaroo skins, thus

denuding these places of grass.'* On one occasion,

Ngurunderi and his sons drove an enormous

Murray cod, Pondi, down the Darling and Murray

Rivers to Piltangk, on the southern shore of Lake

Alexandrina. Here they obtained the assistance of

Nepeli. They eventually caught Pondi near

Raukkan. Ngurunderi tore it into pieces, throwing

each fragment back into the water, and thus

creating different species of fish. The fish-making

episode differs from the Encounter Bay

mythology, where this is performed at a salt water

locality by another spirit named Pungngane

(Meyer, 1846 [1879: 202]). Ngurunderi had four

children by two wives. Once, while camping at

Tulurrug (Pelican Point), two of his children

strayed into the eastern scrub and were lost.

Ngurunderi’s two wives later fled and Ngurunderi

followed them to Encounter Bay where, seeing

them in the distance, he drowned them by making

the waters rise. He then searched up the Coorong

for his two lost children and came across them

after he fought and killed a sorcerer at Salt Creek.

Ngurunderi later left the lower landscape for

Wyirrewarre, the Sky World, taking his children

with him. Although recorded as a myth, Taplin

clearly considered the possibility that ‘creators’

such as Ngurunderi were once actual people. For

example, he refers to Ngurunderi as a ‘deified

chief’ (1874 [1879: 58]). To Taplin, myths were

corrupted versions of history.

In the Lower Lakes district, away from the sea,

the creation of the Lakes was given greater

emphasis. Angas provides a version in which the

two wives of Oorundoo (= Ngurunderi) ‘proved

untractable [sic], and ran away from their lord;

and to punish this unwarrantable behaviour on

their part, Oorundoo very properly made two lakes

to drown them, which correspond with the lakes

Alexandrina and Albert’ (Angas 1847a: 96, 97;

see Fig. 2 of this paper). Another version, possibly

derived from Angas and Europeanised, states:

The Murray dragon, Oorundoo, first caused that

great river to flow. Having fallen out with his two

wives, who must have been dragonesses of a huge

size, and not accustomed to water exercise, that Blue

Beard of New Holland constructed two lakes, at

present known as lakes Victoria [= Alexandrina] and

Albert, so that he might effectually drown his

partners, who had actually attempted to elope from

him with somebody else.!°

Thus there are accounts of the creation of Lake

Albert and Lake Alexandrina that contain

elements similar to those of the Ngurunderi

mythology as recorded at Encounter Bay.

However, these versions makes the mythology

more relevant to descent groups such as the

Piltinyerar, living in the Lower Lakes and Murray

River region, by giving prominence to places

contained within their own mythic landscape.

The most detailed published description of the

Ngurunderi myth is provided by Ronald Berndt,

chiefly from his main Lower Murray informant,

Albert Karloan, a Yaraldi-speaker of the

Manangki descent group.'® Like other accounts

MYTH AS HISTORY? 149

VICTOR

W- Drowned wife

ALEXANDRINA

Figure 2. The wife-drowning sites of the Ngurunderi myth, showing inland and coastal variation.

published by anthropologists, he focuses on the

cosmological and creation aspects of the

mythology. In this version, Ngurunderi pursued

the giant Murray cod Pondi down the previously

narrow bed of the Murray River. As the cod was

chased, it widened the river to its present width.

With each sweep of the tail Pondi created a

swamp. When Pondi escaped into Lake

Alexandrina Ngurunderi called out to his brother

in-law, Nepeli, to spear the cod. Nepeli speared it

near Raukkan (Point McLeay), dragging the fish

to a submerged sandbank to wait for Ngurunderi.

Upon reaching Nepeli, Ngurunderi cut up the fish

into many small pieces. Each portion became a

different species of fish as it was thrown back into

the water. In a later episode of this long account,

Ngurunderi’s two wives broke a food taboo and

fled across the Lake Albert country, heading down

the Coorong. Ngurunderi followed and near

Blackford, which is inland from Kingston, he

came across a sorcerer with whom he quarrelled.!”

Ngurunderi killed him and burnt his body which

formed the Granites near Kingston. Ngurunderi

went back along the Coorong, eventually crossing

the Murray Mouth and moving along the southern

Fleurieu Peninsula coast. He created many of the

landscape features along the way. Ngurunderi

finally found his wives crossing to Kangaroo

Island and drowned them by making the seas rise.

The bodies of the wives became The Pages

islands. Ngurunderi then ascended to the Sky

World, Waieruwar, via Kangaroo Island.'*

Tindale provided other accounts of the

Ngurunderi mythology.'? His main published

version was distilled from recordings he obtained

from many Aboriginal people with varying

backgrounds, including the Maraura of the Upper

Murray cultural region, the Ngaralta and

Nganguruku of the Mid-Murray, and down-river

to the Lower Murray groups such as the

Portaulun, Jarildekald (= Yaraldi), Ramindjeri

and the Tanganekald (= Tangani). It is interesting

150

to view Tindale’s printed version of this

mythology compiled this century, in the light of

the considerable variation in the earlier records.

Not only did Tindale standardise the name of

Ngurunderi, he appears to have chosen particular

accounts of episodes over others with which they

would have conflicted. Although very similar in

its general outline to Berndt’s recorded version,

Tindale’s explanation of some events, such as

what happened to Ngurunderi’s canoes, does not

fit Berndt’s account. However, both Berndt and

Tindale shared the same Yaraldi-speaking

informant, Albert Karloan.

The discussion indicates that there were many

versions of the Dreamtime events within the

Lower Murray region which reflected local biases.

This suggests that the sites Aboriginal people

insert into their accounts of the Dreaming are

‘places’ upon which are built both local and

broader regional identities. The term ‘place’ refers

to a humanistic concept within human geography

that focuses upon the culturally determined values

of particular parts of the landscape (Relph 1976;

Buttimer 1980). Places are fusions of human and

natural order and are the significant centres of the

experiences people have of the world. They can

evoke a sense of belonging to a social group and

provide a sense of group identity. It is likely that

some Lower Murray people, as elsewhere in

Australia, even inserted themselves into the

‘Dreaming’ accounts of the landscape they

experienced.

Age and perhaps gender were other factors in

determining what mythological details were

known by individuals, and what emphasis they

were given. The Dreaming therefore reflects some

socio-political aspects of Aboriginal life. This is

consistent with the Dreaming epics recorded from

other parts of Australia (Myers 1986: 60; Berndt

& Berndt 1989: xxvi). Although on one level this

appears to contradict the cultural homogeneity of

the region, the early mythology is nevertheless

unified to some extent through being organised

according to salt water and fresh water accounts,

with some elements linking all. The proliferation

of ‘creation’ versions are not the result of recorder

or informant error, but the product of different

versions of the ‘cultural landscape’ itself.

NGURUNDERI AND LINKS WITH THE BROADER

LANDSCAPE

The incorporation of new elements from an

altered socio-political environment after European

colonisation is reflected in recorded Aboriginal

MYTH AS HISTORY?

mythology. In the Lower Murray, Aboriginal

mythology was modified as people embraced

Christianity. For instance, Ngurunderi was taken

by the Lower Murray people as a suitable

interpretation of the concept of ‘God’ being taught

to them by early missionaries (Taplin Journals, 25

June 1859; 22 September 1859). For some

contemporary Ngarrindjeri people, this linkage is

once again perceived and articulated by them. One

young adult told me in 1988 ‘Ngurunderi is like

Jesus. God gave Ngurunderi to the people. God

spoke to the people through Ngurunderi.’ Several

older informants claim that the reason that the

graves in the Point McLeay cemetery are arranged

with head towards the west is so that the dead

person’s spirit can go in the direction of

Ngurunderi. In the South East region in 1934, a

Meintangk Aboriginal man, Alf Watson, claimed

that Ngurunderi had warned the Aboriginal people

about the coming of Europeans and their

destruction of the environment when he said

‘Beware of puruki (ants)’ (Tindale, 1934-1937,

vol.2: 57; 1938: 20). The syncretism of tradition is

illustrated in an account of Aboriginal people on

the Point McLeay Mission in the 1880s who

reportedly laid bodies on a Christian-style cross

for a short time before being placed on a pre-

European-type burial platform.” It is clear that a

blend of their own practices with new elements

has been integral to the constant development of

tradition. Old beliefs and customs are made more

relevant to contemporary situations through this

process.

The Ngurunderi myth complex contains

references to areas outside the Lower Murray.

Many of these connections may have been made

after European settlement, when Aboriginal

people from diverse cultural areas were forced into

living at the same locations, such as fringe-camps

and mission stations. For instance, Taplin noted

the belief that two warriors from Ngurunderi’s

group returned to the Upper Murray, but were

never heard of again (1874 [1879: 61]).

Furthermore, in the Ngaiawang mythology of the

Upper Murray, the two wives of Ngurunderi were

said to be the Bakindji sisters involved in the

eagle and crow myth epic of the Upper Murray/

Darling district (Tindale 1939: 259). In the Mid-

Murray region there is mention of Wurranderra—

an ‘Aboriginal Moses’, who reportedly came later

than the main creative ancestor, Noreela, to lead

forth many northern tribes to the rich waters of the

Murray, in the process giving them law and

customs.2! Although Wurranderra is given a

secondary creative role in this account, it is

MYTH AS HISTORY? 151

possible that the name is a poor European

rendering of the word Ngurunderi. Due to the

similarity of this description with that of the

Lower Murray ancestor, Ngurunderi, and the

likely linguistic similarity, it seems probable that

this was an aspect of Mid-Murray mythology

given greater prominence by the Lower Murray

people. Whether knowledge of the Ngurunderi

mythology was widespread in Mid-Murray

regions before European colonisation is not

known. It is possible that Aboriginal people there

incorporated this Ngurunderi-type account during

historic times on the basis of having heard it from

Lower Murray people they had met.

According to Taplin’s informants two other

young men from Ngurunderi’s group had taken a

party south along the Coorong, establishing

themselves in the Lower South East near Mount

Gambier (Taplin 1874 [1879: 62]). Another

version, obtained from Alf Watson by Tindale in

1934, acknowledged Ngurunderi and his two

nephews as ‘creators’ for areas inland from

Kingston, involving places such as Mount Benson

and Cape Jaffa (Tindale 1934-1937, vol.2: 58-

60). All these places are outside the Lower Murray

cultural region. Therefore, although particular

episodes of the Ngurunderi myth, as told by the

Lower Murray people, were chiefly confined to

their own territories, this mythology did provide

links to other cultural regions. In the last half of

the nineteenth century, the large ceremonial

gatherings where initiations were held involved

| Aboriginal people from throughout the Lower

Murray, as well as people from places outside the

cultural region, such as from Murrundi

(Moorunde) on the Murray River, and the Rufus

and Darling River area (Taplin Journals, 24

November 1859; 12 May 1860; 6-10 & 27

August 1860; 17 March 1862). The last recorded

initiation sequence of ceremonies which included

Lower Murray people took place from 1879 to

1882 at Matanga (Metang) south of Mannum, and

involved people from the Victorian and New

South Wales part of the Murray River, as far away

as the Darling River Junction at Wentworth

(Berndt & Berndt 1993: 166, 167). At these large

heterogenous gatherings, revised explanations of

the cosmos could have been achieved to make the

pre-existing mythology more relevant to all

regions.

Some recorded versions of myth reveal

innovations which, because of their wide areal

coverage across major cultural divisions, suggest

post-European origins. A version of the

Ngurunderi epic recorded in 1936 from Clarence

Long, a Tangani-speaker from the Coorong,

involves range of localities from Mount Gambier

in the lower South East to the Adelaide district.”

In this example one of Ngurunderi’s sons travelled

south and was chased by a ‘big devil’ named

Mirka, who emerged from the Blue Lake. The son

fled north along the Coorong and across the

Mount Lofty Ranges to Willunga, forty kilometres

south of Adelaide. Ngurunderi, who saw them

approach, attacked and wounded Mirka at Red

Ochre Cove.” The congealing blood of the ‘devil’

formed a rich red ochre deposit at this place, now

part of an Adelaide outer suburb. Mirka fled back

to the Mount Gambier district, his dripping blood

creating other ochre outcrops.

The existence of this broader account is

probably best explained in terms of post-colonial

Aboriginal people gaining extensive geographical

knowledge through participation in early

agricultural activities throughout the colony, such

as shearing and harvesting.When Aboriginal

people from the Lower Murray region encountered

mythologies concerning a different landscape, it is

likely that they drew close comparisons with their

own creation beliefs. Through this process,

Ngurunderi may have assumed the identity of

mythical ancestors in new areas. The possession

by Aboriginal people of knowledge concerning

key points in the landscape, however short the

experience with them, imparts a sense of authority

over their environment. Mythological sites, as

‘places’, reflect both the political and social

dimensions of the relationship Aboriginal people

have with the landscape.

In 1935 Tindale recorded an unpublished

version of Ngurunderi from an Aboriginal man

named Spender, who had descent connections to

both the Adelaide and Lower Murray regions. In

his journal, Tindale wrote:

Ngurunderi came from east to Cape Jervis; he chased

his two wives who were sisters; they swam towards

Kangaroo Island; he killed them and turned them

into the Sisters islands (the Pages). Ngurunderi

landed on Kangaroo Island beside a sheoak tree. He

went to the western end of the island and threw his

spear out into the water. It made some islands there

(he [Spender] had only heard about them).

Ngurunderi then crossed the sea, manner not defined,

and came to Corny Point. From Corny Point he

entered the sea and swam to “Cotton Island” which

is off Tumby Bay. He landed on the island and, at

low tide, walked over the shallows to the mainland.

He changed his name (name forgotten [by Spender])

and travelled to the west (Tindale 1935 MS: 45).

This version links several distinct cultural

regions across southern South Australia.

Sy P. A. CLARKE

Another example of adaptation was provided by

Lola Cameron-Bonney, who defined herself as a

Coorong Aboriginal person with family

connections to western Victoria, and by Ronald

Bonney, who had links from the Lower Murray to

the South East and Victorian regions.* They gave

a variation of the Ngurunderi epic that appears to

have been influenced by the New Testament. In

this version, Ngurunderi travelled to an Aboriginal

camp in western Victoria and found that people

there were starving. He left, returning half a day

later with fish and bread for the camp. The fish

were of a type not found there. From here,

Ngurunderi then went back into South Australia.

Accounts such as these indicate that Aboriginal

people often make links, through the ‘discovery’

of myth, between their own cultural area and the

new places they find themselves, as part of

extending their ‘cultural landscape’.

For most of this century knowledge of

Ngurunderi within the Aboriginal community was

retained almost exclusively by a few elderly

people who still had some understanding of pre-

European beliefs and customs. It was part of what

has been defined as ‘memory culture’ (Berndt

1974: 22, 25; Tonkinson in Berndt & Berndt

1993: xix). Portrayals of Aboriginal mythology in

the popular literature have helped keep

Ngurunderi at the forefront of public knowledge

of Aboriginal beliefs concerning the ‘Dreaming’.”®

During the early 1980s much site-related

information could still be gathered concerning

historic Aboriginal sites, but not so with the

creation myth information, as it had increasingly

become less relevant to Aboriginal people who

were experiencing the landscape in new ways

(Clarke 1994, Chapter 7). Many localities of

previous mythological significance were no longer

visited by Aboriginal people, who increasingly

lived at missions, working-class urban areas of

Adelaide, and fringe-camps, from the early

twentieth century to the 1960s. The emergence of

Point McLeay (or Raukkan as Aboriginal people

call it) as a ‘place’ that connected all Ngarrindjeri

people through a shared mission history became a

key element of Aboriginal identity in Lower

Murray.

The Ngurunderi mythology was rejuvenated in

the late 1980s, partly through the South Australian

Museum adopting this mythology as the basis of

its Lower Murray cultural exhibition, and also

through the coverage which the Education

Department of South Australia gave it in its

Aboriginal studies curriculum (Hemming 1988;

Hemming & Jones 1989; Education Department

1990; 1991). The video, ‘Ngurunderi: a

Ngarrindjeri Dreaming’, was produced as an

introduction to the Ngurunderi exhibition in the

South Australian Museum.*° This relatively recent

activity has had a major impact upon the portrayal

of Lower Murray culture. The modern promotion

of Ngurunderi has therefore gained much impetus

from forces external to the Aboriginal community.

With the increasing voice that Aboriginal people

are having in heritage issues, coupled with a

renewed focus by them on the past culture,

Ngurunderi is once again a prominent figure,

albeit a standardised one from the many accounts

available.

DREAMING As REALITY

As a number of researchers have pointed out,

myth is defined by Western tradition in two main

contradictory ways (Kirk 1973: 8; Berndt &

Berndt 1989: 1). Myth is firstly construed as an

invented or fictitious story. In this sense, myth is a

false and trivial belief. This definition is often

used by those, possessing the ideology of a

dominant culture, who seek to define what they

consider to be the ‘superstitions’ of historically

subordinate cultures. The second concept of myth

describes it as a ‘traditional’ belief or a reflection

of a culture, and allows for less emphasis on

evaluating its true/false aspects. As an expression

of a culture myth has been well explored by social

science. Durkheim (1915) considered myth as one

of the essential elements of religious life,

representing the way in which society portrays

humanity and the world. Malinowski (1948)

developed this further, seeing the prime function

of myth to be the recording and validating of

cultural institutions. Kirk has pointed out that a

problem with these interpretations is that not all

myths are closely associated with ritual or

religious practices (1973: 11, 12). Some myths

appear quite secular although they still reflect

deeply rooted cultural values.

Although Aboriginal myth _ provides

explanations of the world, the analogy of

Aboriginal myth with a European notion of history

as a systematic and linear record of past events is

confusing. The Aboriginal English use of ‘history’

to mean the ‘Dreaming’ in some regions of

Australia provides non-Aboriginal people a further

source of misunderstanding. Myth can, in some

contexts, be regarded as an Aboriginal version of

history, but it is much more. Levi-Strauss (1966;

1977; 1978) considered all myths, when correctly

understood, to be speculative, or problem

MYTH AS HISTORY?

reflecting. He recognised that myth provided a

view of the world that can be constantly explored

and modified by culture. It follows that a

particular myth can mean different things to

different people, with many equally valid versions.

Many myths have varying layers of meaning

(Berndt & Berndt 1989: 3). They are not static

constructs that withstand empirical testing, but, as

the Lower Murray evidence suggests, are able to

incorporate new elements with ease. Baker

maintains that the Dreaming ‘straddles European

distinctions between politics and religion’ (1989:

110). Following Hiatt (1975) and Sutton (1988), I

argue that history, as it is perceived by Western

Europeans, and Dreaming mythology as perceived

by Australian Aboriginal people, may have similar

functions in their respective cultures, but are

nevertheless not synonymous.

The “places” highlighted in the myths are

tangible points that linked people with the

Dreaming. These focal points in the mythic

landscape helped provide Aboriginal people with

a local identity. Berndt claims that “there is no

religion without sites or without a concept of

“Jandedness” (1970: 53). As the nature of the

cultural relationship with the land changed, such

as through European intrusion, new sequences of

myth were “discovered” or developed so as to

validate it. Geographic information concerning

other parts of southern Australia were

appropriated by myth-makers. Ngurunderi’s tracks

coincided with the spread of Ngarrindjeri people

across the land. When Aboriginal movement

patterns became more restricted by government

agencies in the early twentieth century, Aboriginal

cultural identity became more closely aligned to

mission and fringe-camp sites. Both myth and

tradition adapted to fit new cultural landscapes.

CONCLUSION

The variation described for the Lower Murray

mythology does not represent the poor recording

of a standard account. Rather, as a whole it

153

represents a symbolic text through which differing

relationships that people have with the landscape

are articulated. In the early years of European

settlement, there were many distinctive accounts

of Ngurunderi throughout the Lower Murray

region, reflecting its importance in explaining the

country. Thus the myths reflect the existence of

differing perceptions of the cultural landscape

itself. The perceptual world reflects the perceived

relationships between people. Places featured in

the mythology are tangible elements which help

give authority to particular world views.

Aboriginal people, like other cultural groups,

draw meaning from the landscape.

Through the partial collapse of pre-European

systems, the spread of Lower Murray people in

southern South Australia resulted in the

Ngurunderi mythology becoming significant for a

much broader landscape. When these people

moved outside their region, the feats of the

different ancestors they encountered when

socialising with other groups were sometimes

reassigned to Ngurunderi. As a dynamic record of

the past, in relation to the present, the Ngurunderi

myth complex has been a vehicle for Aboriginal

people to investigate and restate their own

identity. Today, this spirit ancestor remains as a

symbol of the Lower Murray as a cultural region.

This myth is symbolic of a regaining of the past,

albeit through the agency of museum displays and

videos rather than through the earlier context of

ceremonial gatherings at particular important

sites. The construction of mythology has not been

restricted to a time long before European invasion,

but has existed as a continuing process of the

development of tradition up until the present.

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. Drafts of this article were

commented on by Peter Sutton, Tom Gara, and Philip

Jones.

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ENDNOTES

1 Smallpox was introduced into Australia on several occasions,

probably initially in the north by Malays (Cleland 1966:

155, 156). One outbreak reached Port Jackson in 1789. The

occurrence of the disease in the Lower Murray in about 1830

was part of a later wave. See accounts provided by

Teichelmann & Schurmann (1840, pt 2: 34) and Gell (1842

[1904: 99]) for the Adelaide region, and by Hahn (1838-

1839 [1964: 129]) for Hahndorf, Mount Lofty Ranges area.

In the Lower Murray and the South East regions, the disease

is documented by Angas (1847a: 123), Taplin (1879: 45),

Taplin (Journals, 25 June 1861), McCourt & Mincham

(1977: 68, 69; 1987: 148), and Jenkin (1979: 28-30).

Similar records exist for the Darling area (Newland 1889: 1)

and for Victoria (Dawson 1881: 60; Bulmer 1887: 31).

Stirling (1911), Berndt (1989: 64), and Berndt & Berndt

(1993: 292) comment on the timing of these waves.

2 Taplin (1874 [1879]; 1879) uses the term ‘Narrinyeri’ (=

Ngarrindjeri). See Darwin (1890), Ratzel (1896), and

Durkheim (1915) as examples of scholarly interest in

‘primitive’ Aboriginal culture from the Lower Murray

region.

3 For accounts of the distinct separation of ceremonial

knowledge between the sexes in Central Australia see Bell

(1983: 182-228), and Payne (1989: 47-49). These authors

refute that either gender possessed ‘inferior’ sacred

knowledge — just that they were somewhat different and

expressed within their own realms.

4 Hercus (1989: 105, 106), Jacobs (1989: 90) blames an

MYTH AS HISTORY?

external system controlled by non-Aboriginal people for

having constructed a lesser role for women in land-related

issues in relatively recent times.

The same variation is found in other parts of Australia, such

as Arnhem Land (Berndt & Berndt 1989: 389-407).

Nurelli has also been recorded as Noreele (Eyre, 1845, vol.2:

356). Berndt (1974: 27) equates Nurelli from the Mid—

Murray with Nepeli of the Lower Murray. Korna (= Corna)

was a Creation being recorded by Penney from the Encounter

Bay region (Clarke 1991b: 96-98). Thukabi was an ancestral

turtle who created the river and lakes when leaving the land

for the sea (Clarke 1994: 114).

Peterson (1976) summarises the ‘tribe’ debate. Berndt

(1959) provides a critique of the application of ‘tribe’ theory

in the Western Desert region.

Many recorders of the name Ngurunderi, apparently did not

hear the initial ‘Ng’. For example, he is recorded as

“Runderudie’ (Register, 1 June 1885) and ‘Narrundi’

(Register, 21 June 1909). It is also possible that the ‘Ng’ is a

later introduction. In 1894, an Aboriginal man, Jacob Harris,

spelled Ngurunderi as ‘Noorondourrie’ (J. Harris letters,

D6510(L)13, Mortlock Library, Adelaide). Similarly, Parker

(1936) recorded this as ‘Nar-oong-owie’. Nevertheless,

Ngurunderi is the standard orthography now in general use

(Berndt 1940; Tindale 1974: 64, 133; Hemming 1988;

Hemming & Jones 1989; Education Department of South

Australia 1990; Berndt & Berndt 1993). Unless the

rendering of Ngurunderi is greatly altered by the sources

cited, I will use Ngurunderi as the standard term.

See Meyer (1846 [1879: 205, 296]), and Tindale (1934-37:

51, 293, 294; 1930-32: 102-105, 117-121) for Encounter

Bay interpretations of the Ngurunderi mythology. Accounts

by G. Taplin (Register, 30 January 1862; 1874 [1879: 55-

62]; 1879: 38, 39), F. Taplin (Register, 24 April 1889),

Ramsay Smith (1930: 17, 173, 182-184, 216, 250, 317-

331), Tindale (1934-37: 284-292, 295, 296; 1930-52:

120), and Berndt (1940) are concerned more with the inland

areas of the Lower Murray. Versions of Ngurunderi from the

Coorong/South East region are recorded by Tindale (1931—

34: 182, 183, 185; 1934-37: 30-37, 57-60). An unlocalised

version appeared in the Register, 2 December 1893.

Generalised treatments of this myth are given by Tindale &

Pretty (1980: 48-50), Hemming (1988), and Hemming &

Jones (1989).

This version is given by R. Penney (as ‘Cuique’) (South

Australian Magazine, November 1843, vol.2: 331-336).

Cawthorne (1844 [1926: 25, 26]), and Gill (1909: 98, 99)

cite Penney without reference.

Account from the ‘Herald of Kapunda’, in the Register, 4

September 1872.

In a photograph taken by Ransay Smith (c.1920), these rocks

are shown to be the two largest boulders in the shallow water

near the site of the Bluff Whaling Station (AA263, Acc.no:

2126, AP2971, Anthropology Archives, S.A. Museum).

Taplin (1874 [1879: 55-62]). The Taplin Journals (18

December 1861, 10 November 1862) have an outline of the

Ngurunderi epic involving ‘Noorunderee’ and ‘Neppelle’

chasing a ‘giant fish’ off from Tipping (Point Sturt). The

standard spelling of ‘Nepeli’ and ‘Waiyungari’ used here is

taken from Berndt & Berndt (1993).

Tindale (1930-52: 188-191; 1935) recorded the feats of

Waiyungari and Nepeli from a Yaraldi-speaker named Frank

Blackmoor. Other versions are given by G. Taplin (Register,

30 January 1862; 1879, p.56). F. Taplin (Register, 24 April

157

1889), and Berndt & Berndt (1993: 228-230). A similar

account by Laurie (1917, citing Hackett) appears to have in

error replaced ‘Nepeli’ with ‘Nurundie’. The original

manuscript (in a notebook titled ‘Narrung Alpha’, August

1915—personal collection of L. Padman) from which Laurie

probably copied his version refers only to a ‘mighty man’,

with no reference to ‘Nurundie’. Meyer (1846 [1879: 201,

202]) gives a related version that has ‘Pungngane’ instead of

Nepeli.

Bonwick (1870: 204). The account of the wives attempting

to elope with someone else bears more relation to the

Waiyungari account, than that of Ngurunderi (see endnote

14). The description of the main characters as ‘dragons’

appears to be a poetic introduction by Bonwick.

Berndt (1940), and Berndt et al (1993: 13-16, 229-223,

433-441). Tindale’s work with Karloan remains largely

unpublished (Tindale, 1934-37, vol.3: 284-292, 295, 296).

The inclusion of Blackford was probably a recent

introduction. Blackford is a farming property where several

Aboriginal families in the South East region have lived. It is

now managed by the Aboriginal Lands Trust, and leased out

by local Aboriginal people.

Before Aboriginal people were able to perceive Kangaroo

Island as an isolated land mass, it is likely that this place

itself represented the ‘Land of the West’ where souls go. Its

role as a stepping stone to the Sky World is possibly a

modification brought about by Aboriginal experience

commencing during the occupation of the island by European

sealers during the early nineteenth century. One of the

Ngarrindjeri names for this previously uninhabited island was

‘Ngurungaui’, reportedly meaning the dead spirit ‘is

departing to travel along the track of Ngurunderi’ (Berndt

1940: 181).

Tindale published the ‘Story of Ngurunderi’ in the

Advertiser, 16 May 1936. His main Coorong informant used

in this account was Clarence Long (Tindale 1931-34: 182,

183, 185; 1934-37: 30-36, 223). Tindale & Pretty (1980:

48-50) give an outline of the composite account earlier

developed by Tindale.

Bellchambers (1931: 112, 125) recorded the Noreela myth

from a Mid-Murray Aboriginal man named Natone (Nettoon).

Although Noreela is female, there appears to be a close

linguistic similarity of the name with the male spirit, Nurelli.

Tindale (1981-34: 182, 183, 185; Advertiser, 16 May

1936). See discussion of this myth by Clarke (1991a: 68).

The importance of Red Ochre Cove, in the post-contact

situation, is indicated by the record of Lower Murray people

from Point McLeay Mission, travelling to Noarlunga to

obtain initiation ochre in 1860 (Taplin Journals, 12

September 1861). Tindale calls this place Putawatang

(Tindale, 1934-37: 154). Berndt & Berndt (1993: 20, 23,

117, 129, 234, 312, 446) record this site as Mulgali or

Putatang.

Account received during South East fieldwork by S.

Hemming and P. Clarke in the mid to late 1980s.

Ramsay Smith (1930: 182-184, 317-331), Parker (1936),

Mountford & Roberts (1969: 20-25; 1971: 34, 35), and

Reed (1980: 64-72).

“Ngurunderi: a Ngarrindjeri Dreaming’ was produced by the

South Australian Film Corporation and the South Australian

Museum. It has appeared several times on national television.

Hemming (1988) discusses the making of this video.

OBITUARY

NORMAN B. TINDALE

12 October 1900 — 19 November 1993

In 1967 at the age of sixty-six, and after a

professional career of forty-nine years spent in the

service of the South Australian Museum, Norman

Barnett Tindale received an honorary doctorate

from the University of Colorado. Among the

voluminous manuscripts bequeathed by Tindale to

the South Australian Museum is a collection of

thirty-nine letters written by colleagues and peers

from around the world in support of this award.

As more than one letter observed, that such an

award was being contemplated by an American

university did not reflect well upon the lack of

initiative of Australian institutions in this respect.

Tindale was eventually honoured with a doctorate

by the Australian National University in 1980. But

none of those letter writers, assessing the

contribution of an anthropologist and scientist in

the twilight of his career, could have predicted

that Tindale would continue to publish and

undertake research for another quarter of a

century.

Tindale was an early starter as well as a late

finisher. He had already published thirty-one

papers on entomological, ornithological and

anthropological subjects before receiving his

Bachelor of Science degree at the University of

Adelaide in March 1933. These papers joined a

further 100 papers published during his

employment at the South Australian Museum, an

average output of three papers each year, mostly

published in refereed journals. In this sense he

was an outstanding product of the British

institutional science tradition—trained on the job,

self-educated and judged by scientific

contributions rather than degrees. That the

Australian expression of this tradition also

reflected a perennially meagre budget commitment

to science on the part of state governments and a

general lack of public support in most intellectual

areas may have been apparent to Tindale, but was

rarely dwelt upon. There was work to do.

Tindale was born in Perth on 12 October, 1900,

the eldest of four children. His parents were

committed members of the Salvation Army and in

1907 the family travelled to Tokyo, Japan, where

his father took up a position as an accountant with

the Salvation Army mission operating in China.

Tindale’s personal and professional life was

marked by turning points; this was the first. He

grew up with a good knowledge of German and

French, as for several years these were the only

languages taught in the small private school

which he attended with the children of diplomats.

One of Tindale’s close school friends was Gordon

160

Bowles, later to become Professor of

Anthropology at Syracuse University as well as a

colleague in wartime Intelligence work. But

Tindale spent most time with the children of

Japanese neighbours, speaking street Japanese,

playing in the semi-rural suburbs of Tokyo (still a

largely feudal city), and exploring the countryside

nearby. It was these rambles, and resultant trips to

Tokyo’s Imperial Museum, which introduced

Tindale to the world of natural history and to

entomology in particular. Through the Museum,

his father’s library, and his own experience of

Japanese customs, Tindale gained a taste for

anthropology.

But by the time that the Tindales left Japan

during August 1915 to settle first in Perth and by

February 1917, in Adelaide, Norman had no doubt

that he would pursue a career as a natural

scientist. Butterfly and moth collecting had

become his passion and he explored the possibility

of gaining a job at the South Australian Museum.

Aware of a possible impending vacancy there, he

took up a position as a library cadet at the

Adelaide Public Library in May 1917 working

alongside another young cadet, the future nuclear

scientist Mark Oliphant. More than thirty years

later, Tindale encountered Oliphant again, in the

Top Secret area of the Washington’s Pentagon,

emerging from a section labelled ‘Manhattan

Project’.

A few months after taking up his cadet’s

position Tindale lost the sight of one eye in an

acetylene gas explosion while assisting his father

with ‘limelight’ photographic work. The accident

dulled none of Tindale’s enthusiasm or ambition.

Just before the explosion he had begun to read

Alfred Wallace’s ‘Travels on the Amazon and Rio

Negro’; a few days after the explosion he took it

up once more and wrote in his diary: ‘My mind

seems made up about following such a life as his.

I hope to take him as my model’ (Tindale ms.). In

January 1919 he finally secured a Museum

position as Entomologist’s Assistant under the

mercurial Arthur M. Lea. He later recalled that

Lea told him, ‘Tindale, you’ll never make a blind

entomologist, but you might make a blind

anthropologist!’. Both seemed possible to the

young scientist.

The next turning point in Tindale’s career came

when he received permission in 1921 to undertake

an extended field trip to Groote Eylandt in the

Gulf of Carpentaria. The opportunity arose

through Tindale’s family background in

missionary activity. This had brought him into

contact with the Church Missionary Society of

P. G. JONES

Australia and Tasmania, which was extending its

mission work from a base at Roper River to

Groote Eylandt. Tindale was engaged by the

Society for twelve months to assist in the

establishment of a home on Groote Eylandt for

half-caste children from the mainland. He was to

be given time to collect for the South Australian

Museum which would purchase his specimens at

the completion of the trip. At this stage no

Aboriginal objects from the island were preserved

in any museum. Tindale’s Director, Edgar Waite,

recognised the ethnographic potential of the

expedition and directed the young entomologist to

visit the doyen of Australian anthropologists,

Baldwin Spencer, at Melbourne’s National

Museum, for advice. Spencer’s advice was

simple; to follow the directions for field

observation laid out in ‘Notes and Queries in

Anthropology’ (he gave Tindale his own copy)

and to keep a field journal with a daily record

under every circumstance, even if the following

day’s events invalidated a previous entry. Spencer

also introduced Tindale to the Geographic I

method of language transcription, the basis for

Tindale’s unique collection of more than 150

parallel vocabularies across Aboriginal Australia.

Tindale followed Spencer’s advice to the letter

and gathered a remarkable collection of

ethnographic data and more than 500 artefacts

from Groote Eylandt and Roper River during his

twelve months in the field. This was the longest

period to that date spent by a scientist in the

company of Aboriginal people. During that

expedition Tindale’s main informant, a Ngandi

man named Maroadunei, introduced the young

scientist to the concept of bounded tribal

territories, ‘beyond which it was dangerous to

move without adequate recognition’ (Tindale

1974: 3). Yet Tindale went to Groote Eylandt and

the Roper as a naturalist, and returned as one. The

crucial shift in his career took place well after his

return to Adelaide, when his synthesis of

anthropological data for publication made him

aware of the openings and challenges which the

new field offered. In particular, when Edgar Waite

insisted that Tindale remove tribal boundaries

from a map of Groote Eylandt and the adjacent

mainland being prepared for publication in the

Museum’s Records, maintaining that nomadic

Aborigines could not occupy defined territories,

Tindale realised that a new paradigm in ways of

regarding and describing Aboriginal Australia

was sorely needed.

The Groote Eylandt expedition revealed

Tindale’s remarkable appetite for fieldwork.

OBITUARY — NORMAN BARNETT TINDALE 161

Taken together, his dozens of field trips amounted

to more than seven years of his professional career

spent in the field, an average ratio of nearly two

months of every year. A colleague later observed

that Tindale was never quite himself back at the

office, and it required very little to entice him out

once more. But it was in the field, exposed to the

nuances and implications of the natural and

cultural environment which he regarded as the

unrestricted object of his study, that Tindale came

into his own. His broad-based training enabled

him to undertake this task confidently and to

weave together the diverse strands of natural and

human science. Trained in geology (a Pleistocene

Stratigraphy course at the University of Adelaide)

under Douglas Mawson, geography under

Grenfell Price and heavily influenced by the

publications of Wallace, it was axiomatic that

Tindale would adopt a strongly ecological

approach to his field observation and collecting.

This approach was greatly reinforced by his

contact with Aboriginal people for whom the

distribution and habits of plant and animal species

were crucial data. Tindale’s bibliography reflects

this great diversity of interest and its

complementary character, particularly in the case

of his geological papers which overlap with those

discussing Pleistocene archaeology, or his

entomological or botanical papers which overlap

with interests in linguistics or material culture.

Looking back on Tindale’s career it is possible

to discern half a dozen research paths which he

followed, converging and diverging but persisting

across several decades until his death. Few

specialists would attempt to emulate such a course

today; in Tindale’s time, as his colleague and

friend Joseph Birdsell put it later, it represented

the ‘proper breadth of interest’.'! In entomology,

his first love, Tindale selected the study of the

Hepialidae, one of the most primitive of the moth

families; in geology his particular interest became

the study of Pleistocene shore-lines and Tindale

was to become recognised as one of the ‘foremost

workers on the Pleistocene geology of Australia’

(Daily 1966). In linguistics as in broader

anthropological studies his object was to gather

sufficient data to scientifically describe variation

in Aboriginal culture and society across the

country. The same applied to his physical

' This phrase forms the title for the Masters thesis written by

Tindale’s grand-daughter Karen Walter, dealing with

Tindale’s early career (Walter 1988). I am grateful to her

for the opportunity to use data contained in this thesis for

this survey of Tindale’s life.

anthropological surveys. More focused studies,

such as an investigation of initiation practice,

Western Desert art and mythology, or the detailed

description of a coastal and riverine society,

followed from this survey data. In archaeology,

informed by his geological and ecological training,

Tindale’s object was to establish the broad canvas

on which more specific applied or theoretical

investigations could be painted. Tindale’s field

trips became the testing ground for this tumult of

ideas and theories against a background of wide

reading in each area and constant rapport with

colleagues, nationally and internationally.

The tracks connecting these paths were often of

equal interest. For example, Tindale’s interest in

the primitive Hepialidae led him into the

palaeontological field, linking with his geological

and anthropological interests. He eventually

discovered and described Eoses, considered to be

the most ancient fossil lepidopteran (of Triassic

age), through a careful examination of the Mt

Crosby fossil beds in Queensland. So much for a

blind entomologist. Likewise, Tindale’s expertise

in the Lepidoptera field brought him into contact

with scientists and administrators charged with

eradicating or controlling insect pests, and with

the issue of satisfactorily managing Australia’s

national parks. Tindale was appointed in 1958 as

a member of the national Committee on National

Parks and Reserves of the Australian Academy of

Science and made active contributions to the

growing debate about conservation issues. His

input to South Australian legislation on National

Parks, enacted during 1966, was also

considerable. Through such connections he also

became a founding member of South Australia’s

National Trust committee. As chairman of the

Trust’s Nature Preservation Committee, Tindale

could take much credit for the preservation of the

internationally known glacial pavements at Hallett

Cove. He was able to doubly underline their

significance through his geological expertise with

ancient shorelines and through his careful

documentation of the massive ‘Kartan’ Aboriginal

stone tools associated with this particular locality.

Tindale’s life was full of such connections. The

most striking, yet least known, was the use which

he made of his intimate knowledge of the

Japanese language. At the outbreak of the Second

World War Tindale’s attempt to enlist in the

Australian army was thwarted by his damaged

eyesight. With Japan’s later entry into the war his

value to military intelligence operations was soon

recognised; he and his brothers, together with his

old friend Gordon Bowles, were among the few

162 P. G. JONES

Australians fluent in Japanese. During 1942

Tindale joined the R.A.A.F. and was assigned the

rank of Wing Commander in England before

being transferred to the Pentagon to advise on

strategic bombing. There he headed the military

intelligence unit charged with deciphering Japan’s

military codes and with ascertaining the origin

and volume of production of munitions and spare

parts. Tindale and his small unit spent time

combing through the wreckage of crashed

Japanese aircraft. They intensively analysed this

debris in a laboratory established at his initiative

near Brisbane. Through metallurgical and serial

number analysis and by deciphering the company

marks found on different components Tindale

obtained remarkably accurate data on production

figures and Japanese shortages of critical alloys.

Professor W. V. MacFarlane later wrote: ‘this

somewhat esoteric complex of knowledge of

language, ability to associate minute and

apparently unrelated fragments of information to

induce patterns of understanding, and to deduce

consequences, has been characteristic of his work

amongst Aboriginals from every part of the

continent and its surrounding islands’

(MacFarlane 1966).

Tindale eventually achieved two breakthroughs

which altered the course of the war in the Pacific.

Both are still unknown to the wider public. He

was instrumental in cracking the Japanese aircraft

production code system, which gave the Allies

reliable information as to Japanese air power.

More importantly, he and his unit deciphered the

Japanese master naval code. Another commander

in U.S. military intelligence later wrote that the

success of the attack ‘upon the homeland of Japan

was more effectively measured through the work

of Tindale and his group than through any other

source of intelligence we had available at the time’

(Brown 1966). This fact was established through

the Strategic Bombing survey undertaken in Japan

after the war.

Just as extraordinary was Tindale’s application

of his special skills of detection in halting the only

enemy attack on the continental United States—

the fire-bombing of the Pacific North-West which

occurred for a twelve-month period during the

war. The attacks caused many forest fires and

killed several civilians. Tindale examined the

shattered remains of the balloon carriages which

transported these bombs from Japan and

established not only the rate of their production,

but their points of manufacture and release. With

this information those sites were bombed and

destroyed, ending this form of attack.

Tindale was reluctant to talk about this

momentous phase in his career, believing himself,

even as late as 1989, to be bound by the wartime

British Official Secrets Act. He continued his

practice of keeping a daily journal throughout the

war period but restricted his observations to

natural history and anthropology. He found time

to record anthropological detail and collect

artefacts during his ‘special duties’ in New

Guinea and the Solomon Islands. During spare

moments in America he studied and reorganised

the Australian osteological collection at the

Smithsonian Institution and even discovered a

new species of Lepidoptera (Sthenopis darwinii)

in Tennessee.

Tindale’s entomological career may have been

overshadowed by his anthropological

achievements but he never lost contact with this

first love. This was so in a direct sense; his desk

invariably contained a jar of insects or an open file

on the subject. On field trips his days usually

ended with an examination of the evening’s haul

of moths and insects caught in his portable light-

trap. The same routine was observed, particularly

during summer months, at his Blackwood home,

overlooking the lights of Adelaide. Both in

Adelaide and in retirement in Palo Alto he made

his own carefully chamfered wooden boxes for

Lepidoptera specimens. But Tindale’s entomology

did not simply consist of collection and

description. As with his anthropological studies,

he became interested not only in questions of

geographical variation but in origins and early

stages, as well as in specific issues of ecology.

Tindale had begun collecting and classifying

butterflies in Japan at the age of ten. His

professional entomological studies began during

his Groote Eylandt expedition of 1921-22.

Concentrating upon the Lepidoptera and the

Orthoptera on his return, Tindale undertook

revisions during the later 1920s of the Australian

Mantidae (mantids) and Gryllotalpidae (mole

crickets), regarded for decades afterwards as

standard works in this field. But it was his interest

in the more primitive Lepidoptera, specifically the

Hepialidae, which established Tindale’s long-

standing international reputation as an

entomologist. I. F. B. Common observed that

Tindale’s revision of the Hepialidae family

‘represented a new critical phase in the study of

Australian Lepidoptera’ (Common 1966). Until

the Second World War it was the only major

revision of the fauna which had taken the male

genitalia into account, and thereafter placed the

classification of the Hepialidae on a firm footing.

OBITUARY — NORMAN BARNETT TINDALE 163

This detailed revision, published in several papers

over a period of thirty-two years from 1932 to

1964, provided the basis for ecological studies

leading to more effective control of several

Oncopera species, for example, which are serious

pests of high-yielding sown pastures.

Tindale became a world authority on the

hepialid moths, a notable achievement considering

the difficulties which they present to researchers.

In acknowledging this, H. K. Clench of the

Carnegie Museum observed that the moths are

often rare, and with brief flight periods, difficult

to collect in adequate numbers for study; their

characters are cryptic and, because of their ancient

origins, their distribution poses additional

problems for the investigator who requires an

intimate knowledge of them across their entire

world distribution (Clench 1966). Patiently

amassing material and data over several decades,

visiting museums throughout the world and

collecting in as many regions and environments as

possible, Tindale acquired this knowledge and

earned the respect of his entomological colleagues.

He discovered and described many geographical

races of moths, some species, and many life

histories, carefully analysing the events which he

considered were responsible for each situation. In

John Calaby’s words, his entomological studies

became ‘much more than mere descriptions of

animals’ (Calaby 1966). His attention to the

evolutionary implications of his data was of great

importance to the much broader fields such as

speciation patterns and the general evolutionary

history of the Australian fauna as a whole.

As noted, Tindale’s entomological work often

connected with his anthropological studies. His

interest in the Hepialidae and Cossidae was

particularly apposite here, since many of them

have wood-boring larvae used as food by

Aborigines. Several of his papers addressed this

subject, providing a vivid example of his attention

to the ecological contexts which sustained human

and animal life throughout Australia. Already

inclined to an ecological approach, Tindale was

given great encouragement through his association

with the University of Adelaide-based Board for

Anthropological Research, and in particular its

chairman, J. B. Cleland. The Board undertook

annual expeditions during the university’s August

vacations of the late 1920s and the 1930s with the

primary object of recording series of physiological

and sociological data relating to Aboriginal

groups which had experienced little sustained

contact with Europeans. Cleland ensured that

these data were recorded within an ecological

frame, encouraging Board members to note

aspects of geology, flora and fauna. Tindale

applied this general approach to the study of

Aboriginal territoriality, relating particular groups

to specific environments through a range of

careful observation, backed up by ethnographic

and archaeological evidence. He developed this

approach thirty-five years before territoriality and

ecology (or even Australian anthropology itself)

became voguish fields.

As the Board expeditions progressed during the

1930s Tindale began to correlate his data in ways

which few anthropologists had previously done.

He collected and documented artefacts with a

strong awareness of how their manufacture

reflected necessities dictated by the environment

and opportunities to manipulate or exploit that

environment. Over the years the careful

accumulation of such detail resulted in important

papers on material culture and the Australian

environment, such as his 1976 publication on the

Panara or seed-milling technology of the Central

Australian grasslands. During the 1930s he paid

particular attention to the documentation of social,

as well as technological, processes, and was

encouraged by other Board members to master the

arts of sound and film recording. Tindale first

attempted ethnographic film-making during his

expedition with Herbert Hale to Queensland’s

Princess Charlotte Bay in 1926-27. Films made at

Hermannsburg (1929), MacDonald Downs

(1930), Cockatoo Creek (1931), Mt Liebig (1932),

Mann Range (1933), the Diamantina south of

Birdsville (1934, assisted by H.K. Fry) and

Warburton Range (1935, assisting Stocker)

followed. Of these, his Mann Range sequence,

four reels comprising ‘A Day in the Life of the

Pitjandjara [sic]’ remains the most compelling.

Tindale cleverly edited sequence from dramatic

footage shot over a period of several weeks to

construct a ‘typical’ day in the nomadic existence

of the Pitjantjatjara as they travelled from water to

water through the Mann Ranges. Wax cylinder

recordings of ceremonies and song cycles were

made separately by Tindale on each of these

expeditions; in many cases they represent the only

surviving record of their kind.

Tindale’s concern was not to preserve an

account of culture for its own sake, but to

document in sufficient detail to enable further

analysis by others. ‘Making a useful record’ was a

phrase he often used, applying it equally to his

compilation of 150 parallel vocabularies as to his

descriptions of manufacture and use of spears,

spear-throwers, dishes, stone tools, resin, hair

164 P. G. JONES

string, and other essential items of desert life. An

artefact could be collected, together with examples

of raw materials used in its manufacture, and the

process could be filmed. All processes and

observations and linguistic terms were recorded

meticulously in note form, together with the

genealogies and backgrounds of the makers or

participants, from whom further mythological

detail could be elicited through crayon drawings

on brown paper which he distributed and later

carefully annotated.

The sound recordings made by Tindale during

the 1930s have still not been properly studied. As

with film, he continued to make these vital records

of Aboriginal life well into the 1960s, in central

and northern Australia. On his return to active

fieldwork after the war Tindale adopted reel-to-

reel tape recorders instead of the wax cylinder

machine and began experimenting with colour

film. Willing to adopt any worthwhile advance in

technology during his working life, Tindale

nevertheless stopped short of the computer age.

His voluminous shoe-box card files on Aboriginal

place names and language terms and his own

indexed journals provided a ready entree into most

of his research areas.

Tindale’s particular duty during the Board

expeditions was to note the identities, social and

totemic background and genealogical relationships

of Aboriginal people. This responsibility provided

him with the structure and discipline required to

complete his apprenticeship as an anthropologist.

In the first place, it brought him directly into

contact with Aboriginal people as individuals on a

one to one basis. His calm and straightforward

manner, leavened with an easy humour but

sharpened by an incisive and enquiring approach,

elicited data with a minimum of fuss. Aboriginal

people remembered Tindale with respect and

affection years after his visits. The genealogical

exercise also gave Tindale an appreciation of the

kinship network as the generative basis of

Aboriginal society, in all its different forms across

the continent. But in contrast to the social

anthropologists trained in the British tradition

who were beginning to graduate under Radcliffe

Brown and Elkin during the 1930s, genealogies

represented far more to Tindale than ‘samples

upon which to hang kinship terminology’ (Birdsell

1966). In the detail recorded by him, they provided

concrete evidence of the relationship between

individuals, sites and Dreaming sequences.

Additionally, and with major implications for

his later research, Tindale’s genealogies enabled

him to develop a demographic record of

Aboriginal Australia as a whole. As with his

ambitious work on documenting tribal boundaries

and territories, the continental scale of this

undertaking was rarely even considered as a

possibility by his contemporaries. Tindale first

undertook applied demographic studies through

his analysis of the Tasmanian Aboriginal

descendants of Bass Strait. Published in 1953, the

study remained for many years the best total

community demographic study in Australia.

The field of population dynamics was almost

unknown within Australia when Tindale

undertook his Tasmanian study. He was to make a

major contribution to it, particularly through his

diachronic analysis of the Bentinck Island Kaiadilt

people. In a series of papers Tindale demonstrated

the effective isolation for approximately 3 500

years of this self-contained population of about

120 people. Through meticulous use of the

genealogical method he documented a pre-contact

population crash which reduced the island’s

population by more than 40%. In collaboration

with the serologist Roy T. Simmons, and the

micro-evolutionist Joseph Birdsell, Tindale

demonstrated that the Bentinck Islanders

represented a classic case of founder effect,

described by Birdsell as the ‘most important

remaining portion. of the Sewell Wright Effect’

(Birdsell 1966). Due to the small numbers in the

original emigrant ancestors of the Kaiadilt, their

descendants differ from mainland Australian

Aborigines in some genetic properties as much as

do major racial groups from each other in other

parts of the world.

Tindale’s awareness of the potential of this field

was stimulated through his time in the United

States during 1936-1937. Following his award of

a 1936 Carnegie Research Fellowship (to be

followed by a second in 1959), Tindale spent

several months studying Australian ethnographic

material and lecturing in Europe and the United

States. Apart from meeting Birdsell, who became

a lifelong friend, Tindale made other friends and

contacts in America to the extent that, following

the death of his first wife Dorothy during 1969, he

was readily able to adjust to life there. Meetings

with the environmentally-oriented anthropologists

Alfred Kroeber and Earnest A. Hooton, the

serologist Carl Sauer, and the geographer T.

Griffith Taylor were especially influential,

confirming Tindale’s anthropological development

along lines increasingly distinct from those

promoted by social anthropologists emerging from

the University of Sydney. In particular, Tindale’s

meeting with Kroeber at Berkeley refocused his

OBITUARY — NORMAN BARNETT TINDALE 165

commitment to mapping the tribal distribution

throughout Australia, in order to provide a firm

basis for the study of culture traits. At Harvard

Earnest Hooton further supported this approach,

seeing the importance of a well-documented

distributional template for the anthropometric and

serological studies which he proposed as a project

to Tindale and his favoured student, Joseph

Birdsell. The result of these meetings shifted the

course of South Australian, and Australian,

anthropology.

Hooton’s influence enabled one of the most

ambitious anthropological surveys every

undertaken in Australia, jointly funded by the

University of Adelaide and Harvard University.

During an eighteen month period through 1938—

39, Tindale led a data-gathering expedition

supported by Birdsell and accompanied by their

wives as secretaries and research assistants. The

team travelled by road to almost every Aboriginal

settlement and mission throughout eastern,

southern and south-western Australia. The object

was to undertake a comprehensive survey of the

Aboriginal population in its interaction with the

non-Aboriginal population, several generations

after first contact had occurred. As on the previous

Board expeditions, the pair gathered an immense

range of physiological and sociological data,

measuring, blood-sampling, photographing and

interviewing more than 3 000 individuals. The

project was completed under Birdsell’s leadership

and with Tindale’s active participation during the

years 1952-54, when north-western Western

Australia and parts of the Northern Territory were

surveyed and some earlier ground retraced,

resulting in the survey of a further 2 000

individuals. For Tindale and Birdsell, the 1938—

39 expedition represented the beginning of a firm

friendship and a working partnership which

spanned the next half-century.

The photographic and genealogical records was

also to serve another, unforeseen purpose, making

Tindale’s name familiar to many thousands of

Aboriginal people across the country decades

later. The establishment of the Aboriginal Family

History Project at the South Australian Museum

during the 1980s made these records accessible to

the descendants of those contacted by Tindale and

Birdsell during the 1938-39 and 1952-54

expeditions. Tindale lived long enough to

appreciate the impact which this historical record

was to make upon the lives of Aboriginal people.

Birdsell’s detailed genetic studies of Australian

Aborigines (discussed elsewhere in this volume)

derived from data first gathered during the 1938-

39 expedition. Much of this work was directed

towards establishing the thesis, still controversial,

that Aborigines had arrived in Australia in three

chronologically, and physically, distinct groups. It

was Tindale and Birdsell’s hypothesis that the

first of these groups, ultimately stranded in

Tasmania by rising sea levels, were represented

on the mainland by the ‘negrito’ population in the

‘ecological refuge zone’ of the Queensland

rainforest near Cairns. This group was contacted

and studied by the pair during the 1938-39

expedition. Tindale furthered his interest in

genetics, eventually publishing the first systematic

study of gene flow for any simple human

population. His 1953 paper in ‘Human Biology’,

based upon his massive genealogical data for the

Australian continent, remains a classic.

For Tindale, the 1938-39 expedition enabled

him to interview many of the last Aboriginal

individuals with knowledge of the group

structures and territories of those regions of

Australia overtaken by settlement and pastoralism.

Combining this primary data with that drawn from

manuscripts and secondary sources, he was able

to realise his long-standing ambition to prove that

Aboriginal groups did relate territorially to distinct

regions that could be successfully mapped. His

tribal map of Australia, first published in 1940

and revised in 1974 together with his

encyclopaedic catalogue of Aboriginal tribal

groups, was radical in its fundamental implication

that Australia was not terra nullius—decades

before the Mabo judgement made it a national

issue. As importantly, both the 1938-39 and

1952-54 expeditions (the latter accompanied by

Tindale only until 1953), resulted in several

hundred sheets of genealogical data and more than

6 000 well-documented photographs of Aboriginal

people.

If Tindale warmed to any particular Aboriginal

group among the hundreds encountered by him

during his career it was undoubtedly the

Pitjantjatjara. He and the physical anthropologist

Cecil Hackett spent almost three months in their

company during 1933, observing initiation

ceremonies and the minutae of daily life as family

groups split and reformed during their travels

through the Mann Ranges. Using camels, Hackett,

Tindale and their European guide Alan Brumby

accompanied the group as ‘virtual parasites’,

making a detailed and unique record of a society

on the brink of decisive change. As well as film

and sound records, closely documented artefacts

and genealogies, Tindale prepared the first

detailed vocabulary of the Pitjantjatjara language.

166

He had further opportunities to learn from these

people during field trips to the region during the

1950s and 1960s.

If Tindale warmed to any particular Aboriginal

individual during his career it was undoubtedly

the Tangane man, Milerum (Clarence Long).

Tindale subsequently wrote an entry on Milerum

in the ‘Australian Dictionary of Biography’

(Tindale 1986), detailing the coincidence of

Milerum’s first contact with Europeans, at the age

of six, with Tindale’s mother’s family, inland from

South Australia’s Coorong. After 1931 Milerum

assisted Tindale by recording a substantial corpus

of song in his native Tangane and related

languages, and participated in intensive site

recording throughout the Coorong and Lakes

region. In this Tindale was assisted by the social

anthropologist H. K. Fry, who had already given

Tindale informal training in anthropological

theory. At the time of his death, perhaps Tindale’s

greatest unfinished work was an ethnographic

study of the Coorong region as seen through the

eyes of Milerum. Much of this data was gathered

during Milerum’s extended visits to the South

Australian Museum, during which he became a

‘resident’ artefact maker until his death in 1941.

By the late 1930s Tindale’s fieldwork had taken

him to every geographic and cultural zone of

Aboriginal Australia. His 1921-22 fieldwork in

tropical Australia was complemented by a ten

week expedition to Flinders Island and Princess

Charlotte Bay on Cape York during early 1927 in

the company of a museum colleague, Herbert Hale

(later Director). Tindale’s first, successful

experiments with ethnographic film-making

occurred during this expedition. He made further

expeditions to the tropics during the 1938-39

expedition, and in 1960 and 1963, visiting

Mornington and Bentinck Islands and the

government settlement at Palm Island. In

temperate Australia, Tindale undertook numerous

trips to the Coorong, the Lower Murray, the south-

east of South Australia and Yorke Peninsula,

particularly during the 1930s, and to south-

western Western Australia during 1968. His arid

zone studies began in late 1924 when he and

Herbert Hale worked among the Wailpi and

Adnjamathanha people of the Flinders Ranges.

This expedition also provided Tindale’s first

introduction to Aboriginal rock carvings and to

the iconography of Central Australia. It also

provided a solid basis of comparison for his

analyses of rock art of southern South Australia,

also undertaken during these early decades. From

1928 Tindale’s experience of Central Australia

P. G. JONES

grew rapidly. He was a participant on the Board

for Anthropological Research Expeditons to

Koonibba (1928), Hermannsburg (1929),

MacDonald Downs (1930), Cockatoo Creek

(1931), Mt Liebig (1932), Mann Ranges and

Ernabella (1933), Diamantina River (1934), and

the Warburton Range (1935). During 1934 and

1951 he made individual expeditions to Ooldea. A

further Board for Anthropological Research

expedition to Yuendumu during 1951 signalled

the end of these large, team surveys. Tindale’s

Central Australian fieldwork was completed with

expeditions to the north-west of South Australia

in 1957 and 1966, Haast Bluff (also 1957) and the

Rawlinson Ranges in Western Australia during

1963.

Tindale’s participation in the 1929

Hermannsburg expedition provided him with his

first close encounter with desert people and their

social system. The eight members of the

expedition party travelled on the first train to

arrive in Alice Springs from Adelaide and reached

Hermannsburg at the height of a scurvy epidemic

precipitated by the worst Central Australian

drought this century. The medical members of the

expedition diagnosed the condition and treated it

successfully. Perhaps through this solicitude, and

the obvious numerical congruence, each of the

eight men was assigned a ‘skin’ name

corresponding to the eight sub-class terms used by

the northern Aranda groups. Tindale was given

the brown hawk totemic affiliation, erukalandja,

and was designated as ‘Mbitjana’ (7Zjampatjimpa)

skin, a classification which he was able to apply

to himself in all subsequent dealings with Central

Australian Aborigines.

On the day of Tindale’s official retirement from

the South Australian Museum he set out with his

old friend, the American folklorist, amateur

archaeologist and accomplished bibliographer

John Greenway, on an expedition to Koonalda

Cave on the Nullarbor Plain. It was fitting that

this expedition, colourfully chronicled by

Greenway (1973), had archaeology as its theme.

Tindale’s career had been characterised by the

relentless search for origins which archaeology

expresses as a discipline. From the 1920s

Tindale’s geological studies under Mawson had

trained him to interpret the stratigraphy of an

archaeological site and he was the first in

Australia to use sea-level changes for dating

purposes. As Edmund Gill expressed it, Tindale’s

archaeological work was remarkable for the fact

that he studied sites from ‘a number of points of

view—anthropologic, geologic and palaeologic’

OBITUARY — NORMAN BARNETT TINDALE 167

(Gill 1966). This form of analysis at

archaeological sites near Adelaide such as

Fulham, Pedler’s Creek (Moana) and the South

Para River had prepared him for his crucial role in

Australia’s first truly scientific archaeological

excavation, undertaken during 1929 at Devon

Downs near Swan Reach on the Murray River.

Tindale’s emphasis on stratigraphy led him to

posit a model of culture succession in Australia,

based entirely upon local critieria rather than

adopting European models as had been accepted

by most other Australian archaeologists. Before

Devon Downs, Australian archaeology did not

exist as a discipline, largely because it was

assumed that Aboriginal people were relatively

recent arrivals. Tindale’s meticulous excavation

established not only that Aboriginal people had

lived for several millennia in the Murray valley,

but demonstrated that their strategies for

subsistence had altered in response to

environmental change. He showed how stone

tools, animal bones and cultural remains could be

used to piece together a previously untold story

about Australia’s past. His foresight in preserving

charcoal samples against the predicted

development of C'* dating has received scant

recognition. Nevertheless, critics of Tindale’s

construction of an Australian cultural chronology

based on his Devon Downs, Tartanga and Noola

Rockshelter excavations, together with his

classification of the large ‘Kartan’ implements,

acknowledge the precision of his work and the

quality of his data.

Tindale’s precise work habits, particularly when

applied to the documentation of field collections,

have ensured that several thousand Aboriginal

artefacts, whether archaeological or ethnographic,

may be interrogated by future researchers. Struck

by the apocryphal anecdote related to him in the

British Museum of a worker who had become

unhinged and burnt a large number of collection

records, Tindale determined that this would never

befall his institution. His careful practice of

inscribing each artefact with inked locality data

was followed for many years. Many of his

specimens are also figured and described in his

journals over a collecting period of almost half a

century.

These journals, Tindale’s collected specimens,

and the entire range of his publications, sound

recordings, films, photographs, genealogies,

crayon drawings, maps and other illustrations will

remain as this tireless worker’s legacy. The lasting

significance of this data lies neither in its bulk nor

its scope, but in the fact that it was gathered with

a focused goal in mind, to describe the diversity of

an entire people before transformation by

European contact. Tindale was well aware that his

attempt to do this, as symbolised by his 1974 map

of ‘Aboriginal Tribes of Australia’ and its

accompanying compendium, would never be fully

acceptable and that, indeed, its chances of

acceptance would diminish as more and more

scholars entered the field. Nevertheless, when he

began his task in the 1920s, the number of

practising anthropologists in Australia could be

counted on the fingers of one hand, and he

believed in his ability to complete the task,

although he once confided in this author that he

didn’t have time to die until about 2020.

A side of Tindale’s career which his many

achievements tend to obscure is that concerned

with his role as a public servant. Tindale

represented the public face of anthropology in

South Australia for almost half a century. He

pronounced upon controverial issues and

developments in the ‘Advertiser’ and other

Adelaide newspapers, gave lectures and radio

talks, and answered innumerable public enquiries

about Aboriginal place names, archaeological

finds and so on. Tindale had been a founding

member of the Anthropological Society of South

Australia during 1926 and he continued to

participate in its activities and administration. His

advice was directly sought by government upon

such issues as the creation of a reserve for

Pitjantjatjara people in South Australia’s north-

west during the 1930s, and the policy issues

arising out of his survey of what had become

known as the ‘half-caste problem’, documented

during the 1938-39 expedition. Tindale also found

time to apply his anthropological expertise and

experience of other museums to new exhibits. He

undertook a full-scale renovation of the Museum’s

Pacific Gallery shortly after his return from war

service during 1946. He superintended the

construction of an Indonesian Hall during 1952—

54 (since dismantled) dealing with the cultures

and history of the Asian-Pacific region., and

periodically made important changes to the

Museum’s long-standing Aboriginal display in the

Stirling Gallery. His role as an ‘expositor of

science’ (Day 1966) was best exemplified perhaps

by his co-authorship of three popular children’s

books on the subject of indigenous peoples. ‘The

First Walkabout’ (1954, Longmans), written with

H. A. Lindsay, was awarded a prize for the best

Australian children’s book of the year in 1955.

Another successful historical novel, ‘Rangatira. A

Polynesian Saga’, also co-written with Lindsay,

168 P. G. JONES

was published in 1959 (Rigby), followed by a

factual illustrated children’s book, “The Australian

Aborigines’ (1971, Lloyd O’Neill), co-written

with his daughter Beryl. Tindale was a committed

bibliophile, but as a user, rather than an owner.

His own extensive reference library was often

annotated and full of bookmarks. His early

exposure to the Adelaide Public Library collection

enabled him to save many important reference

works marked for the discard pile; these are now

incorporated within the South Australian Museum

library. Tindale was a great advocate for binding

library volumes. He served as honorary libarian to

the Royal Society of South Australia from 1952 to

1966, as its Secretary during 1935 and as

President during 1949.

There is no doubt that Tindale had developed

formidable skills as an administrator and policy-

maker during his long career. It was unfortunate

that Herbert Hale’s postponement of his own

retirement made it impossible for Tindale to be

considered as an applicant for the job of Director

at the South Australian Museum. Tindale acted in

this position from 1959 until 1960, having spent

the previous year as Visiting Professor of

Anthroplogy at the University of California in Los

Angeles. The lack of opportunity in Adelaide

made the decision to take further American

academic postings easier. Tindale spent terms as

Visiting Professor of Anthropology at the

University of Colorado from 1966-67 and

from 1970-71, and a further term at UCLA

from 1967-68.

This academic recognition in America, where

Tindale gained a widespread reputation among

students and fellow academics for his lecturing

skills, culminated in the University of Colorado’s

award to him of an honorary doctorate during

1967, the result of John Greenway’s initiative.

Australian recognition was less forthcoming.

Through Professor Derek Freeman Tindale was

awarded a Research Fellowship to the Australian

National University during 1973, in company with

his long-term colleague and friend Joseph

Birdsell. This posting enabled him to complete

the final details for his major work, ‘Aboriginal

Tribes of Australia: Their Terrain, Environmental

Controls, Distribution, Limits, and Proper

Names’, a compendium of data dealing with every

known Aboriginal group in Australia. Often

criticised, both at the level of detail and for its

controversial definition of ‘tribe’, the book and

accompanying maps remain incontrovertibly as a

classic work of Australian anthropology.

Tindale was awarded the Verco Medal of the

Royal Society of South Australia during 1956, the

Australian Natural History Society Medallion

during 1968 and the John Lewis Medal of the

Royal Geographical Society of Australasia during

1980. During that year also, Tindale was awarded

his second honorary doctorate, by the Australian

National University. By this time though, Tindale

had made his home in America. His wife of forty-

five years, Dorothy May, had died of leukemia

during 1969. She had accompanied him, together

with the Birdsells, across Australia on the 1938-

39 Harvard-Adelaide expedition, and had

encouraged an appreciation of anthropology in

their two children, Anthony and Beryl. In 1970

Tindale married an old family friend, Muriel

Nevin, whom he had first met in Honolulu during

his 1936 Carnegie Fellowship visit to America.

Apart from occasional research trips to Australia

and butterfly collecting trips elsewhere in North

America, they continued to live at Palo Alto near

Stanford University in Muriel’s small timber

house, bursting at the seams with his research

materials, library, and butterfly specimens. An

adjacent shed provided more storage space and a

workbench for constructing his neat wooden

butterfly boxes.

While Tindale relished the relative seclusion of

his retirement in the United States, he was never

aloof from family or friends. It became almost

standard for Australian anthropologists and

linguists visiting America to adjust their

itineraries to take in a side-trip to discuss points

and issues with ‘Tinny’. A boyish sense of

humour, a readiness to engage with researchers on

their own terms, and an enthusiasm for new

information sustained him through accidents and

episodes of ill health from his mid-80s. He

impressed all visitors during his later years with

the same qualities recorded by earlier

colleagues—an indefatigable commitment to

making an enduring record of Aboriginal life

before the transformations wrought by European

contact. His career’s output of several books and

nearly 200 scientific papers on anthropology,

geology and entomology were used as working

texts for future papers; he did not preserve

bookshelf copies of any of his publications. By

1989 he knew that he would not complete his

Milerum book, nor several other projects.

Unfazed, he scaled his work programme back and

supplied data for Aboriginal place names to the

South Australian Department of Lands. He was

never happier nor more animated than when

confirming a new detail and putting it on the

record for others to use.

OBITUARY — NORMAN BARNETT TINDALE 169

Tindale remained an Honorary Associate of the

South Australian Museum until his death, an

association which spanned more than seven

decades. During this time all of his former

colleagues had left the scene and he observed the

gap between museum and academic anthropology

develop, widen, but then, encouragingly, begin to

close. His letters to the Museum were like those

from someone who has stayed away in the field

too long: they were always completed with the

touching epigram, ‘Please give greetings to all

those who remember me’. During 1993 Tindale

received unofficial confirmation of the award of

Officer in the Order of Australia; this was

presented posthumously, to his widow Muriel.

But the South Australian Museum Board’s 1993

decision to name a public gallery in his honour

may have meant most to him—a ‘museum man’

to the last.

The last word, and most cogent summary of

Tindale’s achievements, rests with an old friend

and scientific colleague, Professor W. V.

MacFarlane, writing in support of Tindale’s

honorary doctorate from.the University of

Colorado:

It is not common in our time to find men with the

skills and insights which Mr Tindale has shown

through his active and productive life. He is basically

a scientist, while skills with language and human

relations fit him for anthropology. His special

interests in entomology, geology and botany

broadened the scope of his ethnographic studies. As

the use of film, tape, carbondating and blood

grouping came into anthropological work he readily

made use of these for the data that they could bring

towards his final synthesis of the cultural history of

the Aboriginals in time and space. In addition, Mr

Tindale has shown throughout his work a tolerance,

humility, honesty and adaptability which made it

easy for him to find collaborators amongst both black

and white men. (MacFarlane 1966)

REFERENCES

BIRDSELL, J. B. 1966. Letter in support of award to N.

B. Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

BROWN, F. M. 1966. Letter in support of award to N.

B. Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

CALABY, J. H. 1966. Letter in support of award to N.

B. Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

CLENCH, H. K. 1966. Letter in support of award to N.

B. Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

COMMON, I. F. B. 1966. Letter in support of award to

N. B. Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

DAILY, B. 1966. Letter in support of award to N. B.

Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

DAY, M. F. 1966. Letter in support of award to N. B.

Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

GILL, E. D. 1966. Letter in support of award to N. B.

Tindale of D.Sc. by University of Colorado. J.

Greenway file, N. B. Tindale Papers, South Australian

Museum.

GREENWAY, J. 1973. ‘Down Among the Wild Men’.

Hutchinson: London.

JONES, P. G. 1985-89. Interviews with Norman B.

Tindale. Tapes and notes in author’s possession.

MACFARLANE, W. V. 1966. Letter in support of

award to N. B. Tindale of D.Sc. by University of

Colorado. J. Greenway file, N. B. Tindale Papers,

South Australian Museum.

TINDALE, N. B. n.d. Biographical data. 2 vols. Copy in

Anthropology Archives, South Australian Museum.

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

their terrain, environmental controls, distribution,

limits, and proper names’. With four sheet map.

University of California Press, Berkeley and Canberra.

TINDALE, N. B. 1986. Milerum. ‘Australian Dictionary

of Biography’ vol. 10: 498-99.

WALTER, K. 1988. ‘The Proper Breadth of Interest’.

Norman B. Tindale: The Development of a

Fieldworker in Aboriginal Australia 1900-1936. M.A.

Thesis, A.N.U., Canberra.

BIBLIOGRAPHY OF NorMAN B. TINDALE

This bibliography is arranged chronologically

under four headings: Ornithology and Botany (10

entries), Geology and Palaeontology (11 entries),

Entomology (42 entries), and Anthropology (135

entries).

Ornithology and Botany

1922

Notes on the birds of Groote Eylandt, Gulf of

Carpentaria. South Australian Naturalist. 9: 10-21.

Birds observed at Roper River, Northern Territory. South

Australian Naturalist. 9: 21-24.

170

1924

Visits to the islands of the Sir Joseph Banks Group.

South Australian Naturalist. 5: 130-132.

1925

Notes on the birds of Groote Eylandt, Gulf of

Carpentaria. Birds observed at Roper River, Northern

Territory. South Australian Ornithologist. 10: 10-24.

1929

Bird notes. Noisy Miner on Nepean River, N.S.W. South

Australian Ornithologist. 10: 73.

1930

Notes at a camp on the River Murray. South Australian

Ornithologist. 10 : 209-212.

1931

Spur-winged Plover (Lobibyx novae-hollandiae) at

‘Poltalloch’ near Wellington. South Australian

Ornithologist. 11: 109.

1934

Bird notes. Paradise Whydah from Fulham Gardens.

South Australian Ornithologist. 12: 216.

1938

Silver gulls feeding on native currants. South Australian

Ornithologist. 14: 171.

1941

A list of plants collected in the Musgrave and Mann

Ranges, South Australia, 1933. South Australian

Naturalist. 21: 8-12.

Geology and Palaeontology

1931

Geological notes on the Iliaura country, North West of

the Macdonnell Range, Central Australia.

Transactions of the Royal Society of South Australia.

55: 32-38.

1933

Tantanoola Caves, South East of South Australia,

geological and physiographical notes. Transactions of

the Royal Society of South Australia. 57: 130-142.

1933

Geographical notes on the Cockatoo Creek and Mount

Leibig country, Central Australia. Transactions of the

Royal Society of South Australia. 57: 206-217.

1935

Hall, F. J. , Fenner, F. J. & Tindale, N. B. Mammal bone

beds of probable Pleistocene age, Rocky River,

Kangaroo Island. Transactions of the Royal Society

of South Australia. 59: 103-106.

P. G. JONES

1952

Gill, E.D., Fairbridge, R.W. & Tindale, N. B. Research

on Eustatic sea levels — South Australia. Report of the

Australian & New Zealand Association for the

Advancement of Science. Sydney, 336.

1954

Eustatic Terrace Committee. Report for South Australia.

Australian Journal of Science. 32(6): 228-229.

1955

Report of the Eustatic sea levels committee. Australian

and New Zealand Association for the Advancement

of Science. Melbourne, July 1955.

1956

Australian and New Zealand Research in Eustacy —

South Australia. Australian Journal of Science. 19:

56.

1957

Committee for Investigation of Quaternary Strandline

changes. Report from South Australia to A.A.A.S.

Dunedin, January, 1957. Australian Journal Science

20(1): 7-8.

1959

Pleistocene strandlines of the upper South East of South

Australia. Transactions of the Royal Society of South

Australia. 82: 119-120.

1983

The Woakwine Terrace in the South East of South

Australia and indications of the very early presence of

man Pp.583—600 in. P.M. Masters & N.C. Flemming

(eds.) ‘Quaternary Coastlines and Marine

Archaeology’. Academic Press: London.

Entomology

1922

On a new genus and species of Australian Lycaeninae.

Transactions of the Royal Society of South Australia.

46: 537-538.

1923

On Australian Rhopalocera. Transactions of the Royal

Society of South Australia. 47: 342-354.

Flora and fauna of Nuyts Archipelago and the

Investigator Group. No. 13 — Orthoptera.

Transactions of the Royal Society of South Australia.

47: 362-364.

Review of Australian Mantidae. Records of the South

Australian Museum. 2 (3): 425-457.

1924

Review of Australian Mantidae, Pt. II. Records of the

South Australian Museum. 2 (4): 547-552.

OBITUARY — NORMAN BARNETT TINDALE 171

Notes on the life history of the moth (Cacoecia). South

Australian Naturalist. 6: 7-9.

1927

New butterfly of the genus Papilio from Arnhem Land.

Records of the South Australian Museum. 3(3): 339-

341.

1928

Australian mole-crickets of the family Gryllotalpidae

(Orthoptera). Records of the South Australian

Museum. 4(1): 1-42.

Preliminary note on the life history of Synemon,

(Lepidoptera, Family Castniidae). Records of the

South Australian Museum. 4(1): 143-144.

Species of Chlenias attacking pines (Lepidoptera, Family

Boarmiidae). Records of the South Australian

Museum. 4(1): 43-48.

1930

Mantidae in the Australian Museum. Records of the

Australian Museum. 17(8): 343-354.

1932

Revision of the Australian Ghost Moths (Lepidoptera

Homoneura, Family Hepialidae). Records of the

South Australian Museum. 4(4): 497-536.

1933

Revision of the Australian Ghost Moths (Lepidoptera

Homoneura, Family Hepialidae) Pt. II. Records of the

South Australian Museum. 5(1): 13-43.

1935

Revision of the Australian Ghost Moths (Lepidoptera

Homoneura, Family Hepialidae), Pt. III. Records of

the South Australian Museum. 5(3): 275-536.

1937

Tindale, N. B. & Womersley, H. Lepidoptera. BANZARE

Reports. Ser.B, 4( 3): 83-86.

1939

Tindale, N. B. Ghost moths of the family Hepialidae.

South Australian Naturalist. 19(1): 1-6.

1941

Tindale, N. B. Life history of a Convolvulus feeding

moth Aedia acronyctoides (Guenee 1854);

Lepidoptera Heteroneura, Family Noctuidae. Records

of the South Australian Museum. 7(1): 47-50.

1941

Revision of the Ghost Moths (Lepidoptera Homoneura,

Family Hepialidae), Pt. IV. Records of the South

Australian Museum. 7(1): 15-45.

1942

Revision of the Ghost Moths (Lepidoptera Homoneura,

Family Hepialidae), Pt. V. Records of the South

Australian Museum. 7(2): 151-168.

1945

Triassic insects of Queensland. Proceedings of the Royal

Society of Queensland. 56(5): 37-46.

1947

New race of Tisiphone abeona Donovan (Lepidoptera

Rhopalocera) from South Australia. Records of the

South Australian Museum. 8(4): 613-617.

1949

New Satyridae of the genus Oreixenica from South

Australia and New South Wales, together with notes

on the recent climate of Southern Australia. Records

of the South Australian Museum. 9(2): 143-155.

1952

A new butterfly of the Ogyris. South Australian

Naturalist. 27(2): 31-33.

On a new form of Heteronympha penelope Waterhouse

(Lepidoptera Rhopalocera, Family Satyridae).

Transactions of the Royal Society of South Australia.

75: 25-29.

1953

New Rhopalocera and a list of species from the

Grampian Mountains, Western Victoria. Records of

the South Australian Museum. 11(1): 43-68.

On some Australian Cossidae including the moth of the

Witjuti (Witchety) grub. Transactions of the Royal

Society of South Australia. 76: 56-65.

On a new species of Oenetus (Lepidoptera, Family

Hepialidae) damaging Eucalyptus saplings in

Tasmania. Transactions of the Royal Society of South

Australia. 76: 77-79.

1954

On a new genus of Hepialid moth from Rarotonga in the

Pacific Islands. Annals and Magazine of Natural

History. Ser. 12. 8: 13.

1955

Revision of the Ghost Moths, Part 6. Records of the

South Australian Museum. 11(4): 307-344.

Notes on the eucosmid (olethreutid) moth,

Cryptophlebia ombrodelta (Lower). Transactions of

the Royal Society of South Australia. 78: 97-98.

1958

Revision of the Ghost Moths (Lepidoptera Homoneura,

Family Hepialidae), Pt. 7. Records of the South

Australian Museum. 13(2): 157-197.

Witchety Grub. ‘Australian Encyclopedia’. Sydney, 9:

339-340.

1961

A new species of Chlenias (Lepidoptera Boarmiidae) on

Acacia aneura, with some Central Australian native

beliefs about it. Records of the South Australian

Museum. 14(1): 193-196.

1962

The chlorocresol method for field collecting. Journal of

the Lepidopterists Society. 15(3): 195-197.

1963

Origin of the Rhopalocera stem of the Lepidoptera. Proc.

XVI International Congress of Zoology. Washington.

1: 304.

1964

Revision of the Ghost Moths (Lepidoptera Homoneura,

Family Helpalidae). Part VIII. Records of the South

Australian Museum. 14(4): 663-668.

1965

A new species of Holochila (Rhopalocera, Family

Lycaenidae) from Victoria and Southern New South

Wales. Records of the South Australian Museum.

15(1): 10.

1968

On a new Oxycanus (Lepidoptera Homoneura, Family

Hepialidae) from New Guinea. Zoologische

Mededlingen. 42(27): 303-305.

1970

Tindale, N. B. & Le Souef, J. C. A new subspecies of

Virachola_ smilis (Hewitson) (Lepidoptera

Lycaenidae) from Northern Australia. Journal

Australian Entomology Society. 9: 219-222.

1980

Origin of the Lepidoptera with description of a new Mid-

Triassic species and notes on the origin of the

butterfly stem. Journal of the Lepidopterists Society.

34: 263-285.

1981

The original of the Lepidoptera relative to Australia. Pp.

957-976 in ‘Ecological Biogeography of Australia’.

Ed. A. Keast. Junk, The Hague.

1982

The Clench method of relaxing and spreading

butterflies for the collection. Victorian Entomologist.

12(1): 6-8.

Anthropology and Archaeology

1925

Natives of Groote Eylandt and of the West Coast of the

Gulf of Carpentaria, Pt. I. Records of the South

Australian Museum. 3(1): 61-102.

Hale, H. M. & Tindale, N. B. Observations on

P. G. JONES

Aborigines of the Flinders Ranges and records of rock

carvings and paintings. Records of the South

Australian Museum. 3(1): 45-60.

1926

Natives of Groote Eylandt and of the West Coast of the

Gulf of Carpentaria, Pt. II. Records of the South

Australian Museum. 3(2): 103-134.

Tindale, N. B. & Mountford, C. P. Native markings on

rocks at Morowie, South Australia. Transactions of

the Royal Society of South Australia. 50: 156-159.

Tindale, N. B. Native burial at Pedler’s Creek, South

Australia. South Australian Naturalist. 8(1): 10.

1927

Tindale, N. B. & Sheard, H. L. Aboriginal rock

paintings, South Para River, South Australia.

Transactions of the Royal Society of South Australia.

51: 14-17.

1928

Native rock shelters at Oenpelli, Van Diemen Gulf,

North Australia. South Australian Naturalist. 9(2):

35-36.

Natives of Groote Eylandt and the West Coast of the

Gulf of Carpentaria, Pt. III. Transactions of the Royal

Society of South Australia. 52: 5-27.

Ethnological notes from Arnhem Land and from

Tasmania. Transactions of the Royal Society of South

Australia. 52: 223-224.

1929

Hale, H. M. & Tindale, N. B. Further notes on

Aboriginal rock carvings in South Australia. South

Australian Naturalist. 10(2): 30-34.

1930

Hale, H. M. & Tindale, N. B. Notes on some human

remains in the Lower Murray Valley, South Australia.

Records of the South Australian Museum. 4(2): 145—

218.

1931

Tindale, N. B. & Maegraith, B. G. Traces of an extinct

Aboriginal population on Kangaroo Island. Records

of the South Australian Museum. 4(3): 275-289.

1932

Primitive art of the Australian Aborigines. Manuscripts.

No. 3, November, 38-42.

Notes on the supposed primitive stone implements from

the Tableland regions of Central Australia. Records of

the South Australian Museum. 4(4): 483-488.

1933

Hale, H. M. & Tindale, N. B. Aborigines of Princess

Charlotte Bay, North Queensland, Pt. I. Records of

the South Australian Museum. 5(1): 64-116.

OBITUARY — NORMAN BARNETT TINDALE 173

Tindale, N. B. & Hackett, C. J. Preliminary report on

field work among the Aborigines of the north-west of

South Australia. Oceania. 4(1): 101-105.

1934

Vanishing tribes of Arunta Desert. The Advertiser. 17

September.

Initiation ceremonies. The Advertiser. 29 December.

Hale, H. M. & Tindale, N. B. Aborigines of Princess

Charlotte Bay, North Queensland, Pt. 2. Records of

the South Australian Museum. 5(2): 117-172.

1935

Legend of Waijungari, Jaralde tribe, Lake Alexandrina,

South Australia, and the phonetic system employed in

its transcription. Records of the South Australian

Museum. 5(3): 261-274.

Initiation among the Pitjandjara natives of the Mann and

Tomkinson Ranges in South Australia. Oceania. 6(2):

199-224.

Cultural status of the Australian Aborigine. Mankind.

1(11): 264.

Rock markings in South Australia. Antiquity. 9: 93-95.

General report on the Anthropological expedition to the

Warburton Range, Western Australia, July—

September, 1935. Oceania. 6(4): 481-485.

1936

Tindale, N. B. & Mountford, C. P. Results of the

excavation of Kongarati Cave near Second Valley,

South Australia. Records of the South Australian

Museum. 5(4): 487-502.

Notes on the natives of the Southern portion of Yorke

Peninsula, South Australia. Transactions of the Royal

Society of South Australia. 60: 55-69.

Tindale, N. B. Legend of the Wati Kutjara, Warburton

Range, Western Australia. Oceania. 7(2): 169-185.

Tindale, N. B. General report on the anthropological

expedition to the Warburton Range, Western

Australia. Oceania. 6(4): 481-485.

Tindale, N. B. Australian Aboriginal Songs. Inst.

International D’Coop. Intell. Musique et Chansons

Populaires. League of Nations: Paris. 2: 1-3.

Tindale, N. B. & Ward, L. K., Campbell, T. D. & Hale,

H.M. Fossil Man in the State of South Australia.

Report of XVI International Geological Congress.

Washington, 1933: 1271-1273.

Cleland, J. B. & Tindale, N. B. Natives of South

Australia. Pp.16-29 in ‘The Centenary History of

South Australia’. Supplement to Proceedings of the

Royal Geographical Society of Australasia, South

Australian Branch 36.

1937

‘Vocabulary of Pitjandjara. The language of the natives

of the Great Western Desert’. Pp.1—138. (Limited

edition of 6 copies, copyright registered in Public

Library of South Australia, 25 August 1939).

Two legends of the Ngadjuri tribe from the middle north

of South Australia. Transactions of the Royal Society

of South Australia. 61: 149-153.

Relationship of the extinct Kangaroo Island culture of

Australia, Tasmania and Malaya. Records of the

South Australian Museum. 6(1): 39-60.

Native songs of the South East of South Australia.

Transactions of the Royal Society of South Australia.

61: 107-120.

Natives of the Western Desert of Australia. Man. 34: 33.

Tindale, N. B. & Bartlett, H. K. Notes on some clay pots

from Panaeati Island, South East of New Guinea.

Transactions of the Royal Society of South Australia.

61: 159-162.

Tasmanian Aborigines on Kangaroo Island, South

Australia. Records of the South Australian Museum.

6(1): 29-37.

1938

A game from the Great Western Desert of Australia.

Man. 38: 128-129.

1939

Prupe and Koromarange. A legend of the Tanganekald,

Coorong, South Australia. Transactions of the Royal

Society of South Australia. 62: 18-23.

Australie du Sud. Contribution de M. Norman B.

Tindale. Folklore Musical, Dept, D’Art,

D’ Archaeologie et D’Ethnologie Inst. International de

Cooperation Intellectuelle, Paris, pp. 1-3.

Eagle and crow myths of the Maraura tribe, Lower

Darling River, N.S.W. Records of the South

Australian Museum. 6(3): 243-261.

Summary of lectures by Norman B. Tindale on July 3rd,

1939. Mankind. 2(7): 233.

Notes on the Ngaiawung tribe, Murray River, South

Australia. South Australian Naturalist. 20(1): 10-11.

1940

Adelaide-Harvard Universities 1939 Expeditions.

Summary of lecture read before South Australian

Society. Mankind. 2(8): 281.

A curious arrangement of sticks near Lyndoch, South

Australia. South Australian Naturalist. 20(2): 24-25.

A grooved stone mace-head? South Australian

Naturalist. 20: 26.

Distribution of Australian Aboriginal tribes: A field

survey. Transactions of the Royal Society of South

Australia. 64(1): 140-231.

Some Japanese Prints. Bulletin of the National Gallery

of South Australia. 2(2).

Stone Figure of Shou Lao. Bulletin of the National

Gallery of South Australia. 2(3): 1.

174

1941

Antiquity of man in Australia. Australian Journal of

Science. 3: 144-147.

Exhibit of stone implements. Mankind. 3(2): 69-71.

Native songs of South East of South Australia, Pt. II.

Transactions of the Royal Society of South Australia.

65(2): 233-243.

Polychrome incised pottery ware from Mt. Turu, New

Guinea. Records of the South Australian Museum.

6(4): 357-362.

The hand axe used in the Western Desert of Australia.

Mankind. 3(2): 37-41.

Tasmanian stone implement made from bottle glass.

Papers and Proceedings of the Royal Society of

Tasmania. 1941, pp.1-2.

Survey of the half-caste problem in South Australia.

Proceedings of Royal Geographical Society of

Australasia, S.A. Branch 42: 66-161.

Tindale, N. B. & Birdsell, J. B. Tasmanoid tribes in

North Queensland. Records of the South Australian

Museum. 7(1): 1-9. (Results of Harvard-Adelaide

Universities Anthropological Expedition, 1938-1939).

Tindale, N. B. & Noone. H. V. Analysis of an Australian

Aboriginal’s horde of knapped flint. Transactions of

the Royal Society of South Australia. 65(1): 116—

122%

1945

Microlithic mounted stone engraver from Western

Queensland. Queensland Naturalist. 12(5): 83-84.

1946

Australian Aborigines. Encyclopedia of Literature. New

York, vol. 1, pp. 74-78.

1947

Subdivision of Pleistocene time in South Australia.

Records of the South Australian Museum. 8(4):

619-652.

1949

Large biface implements from Mornington Island,

Queensland, and from South Western Australia.

Records of the South Australian Museum. 9(2):

157-166.

1950

Positive approach to an absent subject. South Australian

Numismatic Journal. 1(5): 39.

Palacolithic Kodj axe of the Aborigines and its

distribution in Australia. Records of the South

Australian Museum. 9(3): 257-274.

1951

Palaeolithic Kodj axe of the Aborigines. Further notes.

Records of the South Australian Museum. 9(4):

371-374.

P. G. JONES

Only a strong Australian can survive. Trade Digest.

February, pp.10-11.

Aboriginal net making. Mankind. 4(6): 257-258.

Comments on supposed representations of giant bird

tracks at Pimba. Records of the South Australian

Museum. 9(4): 381-382.

1952

Rock paintings of Cairns area, North Queensland. North

Queensland Naturalist. 20(102): 25-28.

1953

Cassowary-horn native ornament from Arnhem Land.

South Australian Ornithologist. 21: 11.

Growth of a people. Records of the Queen Victoria

Museum. 2: 1-64.

Tribal and intertribal marriage among the Australian

Aborigines. Human Biology. 25(3): 169-190.

1954

Tindale, N. B. & Cleland, J. B. Ecological surroundings

of the Ngalia natives in Central Australia and native

names and uses of plants. Transactions of the Royal

Society of South Australia. 77: 81-86.

Tindale, N. B. & Lindsay, H. A. ‘The First Walkabout’.

Longmans Green & Co: London.

1955

Archaeological site at Lake Menindee, New South

Wales. Records of the South Australian Museum.

11(3): 269-298.

1956

The tasty witchetty grub. A.B.C. Weekly. 18(27): 5.

Peopling of South Eastern Australia. Australian Museum

Magazine. 12(4): 115-120.

Place names. Pp.6—7 in ‘Hiking in the Flinders Ranges,

South Australia’. Boy Scouts Association of South

Australia: Adelaide.

First Australian: the Aborigine, past, present and

prospect. Pacific Discovery. 9(5): 6-13.

1957

Men roamed Australia 12 000 years ago. A.B.C. Weekly.

July 10: 10.

Culture succession in South Eastern Australia from Late

Pleistocene to the Present. Records of the South

Australian Museum. 13(1): 149.

A dated Tartangan implement site from Cape Martin,

South East of South Australia. Transactions of the

Royal Society of South Australia. 80: 109-123.

Future progress of archaeology in Australia. A.G.M.A.

News Bulletin, 10 October, 4-5. (I. Wood, Brisbane).

1958

Australian Aborigines. ‘The New International Illustrated

Encyclopaedia’. Melbourne, vol. 1: 268-276.

OBITUARY — NORMAN BARNETT TINDALE We)

The White Contact. ‘Australian Encyclopaedia’. Sydney,

vol. 1: 87-95

Tindale, N. B. & Lindsay, H. A. The legacy of the

Aborigines. Beaumaris Tree Preservation Society.

Native plants and Seaside Gardens. Melbourne

Rn Coon

1959

A Trobriand Medusa? Man. 59: 49-50.

Totemic beliefs in the Western Desert. Part. I. Records

of the South Australian Museum. 13(3): 305-332.

Ecology of Primitive Aboriginal Man in Australia.

Bodenheimer, F. S., Monographiae Biologicae. 8:

36-S1.

1959

Tindale, N. B. & Cleland, J. B. The native names and

uses of plants at Haast Bluff, Central Australia.

Transactions of the Royal Society of South Australia.

82: 124-140.

Tindale, N. B. & Lindsay, H. A. ‘Rangatira (The High-

born) A Polynesian Saga’. Harrap: London.

1960

Man of the hunting age. Colorado Quarterly. 8(3): 229-

245.

1961

Tribal distribution and populations. Social Science

Research Council of Australia. Conference on

Aboriginal Studies. Data Paper 7: 1-14.

Archaeological excavation of Noola rock shelter: a

preliminary report. Records of the South Australian

Museum. 14(1): 193-196.

Some population changes among the Kaiadilt of

Bentinck Island, Queensland. Tenth Pacific Science

Congress. Abstracts of Symposium papers: 87-88.

1962

Geographical knowledge of the Kaiadilt people of

Bentinck Island, Queensland. Records of the South

Australian Museum. 14(2): 259-296.

Some Population Changes Among the Kaiadilt People of

Bentinck Island, Queensland. Records of the South

Australian Museum. 14(2): 297-336.

Tindale, N. B., Simmons, R. T & Birdsell, J. B. Blood

group genetical survey in Australian Aborigines of

Bentinck, Mornington and Forsyth Islands, Gulf of

Carpentaria. American Journal of Physical

Anthropology. 20(3): 303-320.

1963

Totemic Beliefs in the Western Desert of Australia. Part

IJ. Musical rocks and associated objects of the

Pitjandjara people. Records of the South Australian

Museum. 14(3): 499-514.

A Tjurunga-like Stone Pendant from New South Wales.

Records of the South Australian Museum. 14(3):

555-559

1964

Comments on flint implements found near Nipa, Central

Papua Highlands. Records of the South Australian

Museum. 14(4): 670-673.

Notes. Pp.14-15 in Blainey, J.M. Clues to the Murray’s

biggest floods. Riverlander. August, 1964.

Radiocarbon dates of interest to Australian

archaeologists. Australian Journal of Science.

27(1): 24.

Simmons, R. T., Graydon, J. J. & Tindale, N. B. Further

blood group genetical studies on Australian

Aborigines of Bentinck, Mornington and Forsyth

Islands and the mainland, Gulf of Carpentaria,

together with frequencies for natives of the Western

Desert, Western Australia. Oceania. 35(1): 66-80.

1965

Stone implement making among the Ngadadjara and

Pitjandjara of the Great Western Desert. Records of

the South Australian Museum. 15(1): 131-164.

Progress in Australian Archaeology. Archaeology and

the Living People in Australia. A.N.Z.A.A.S. Hobart.

Summary of papers in Section F, August 1965, 3

pages.

1966

Tindale, N.B. Insects as food for the Australian

Aborigines. Australian Natural History. 15(6):

179-183.

Review. Berndt, R. M. & Berndt, C. H. ‘World of the

first Australians’, American Anthropology. 68(4):

1031-1033.

1967

Peopling of the lands southeast of Asia. Colorado

Quarterly. Spring, 1967, pp. 339-353.

1968

Review. Commonwealth Film Unit. Desert people

(Australia), American Anthropology. 70(2): 437-438.

Nomenclature of archaeological cultures and associated

implements in Australia. Records of the South

Australian Museum. 15(4): 615-640.

1970

H. M. Cooper — A tribute. Pp. vii — viii in Cooper, H.

M., Kenny, M. & Scrymgour, J. ‘Hallett Cove. A Field

Guide’. South Australian Museum: Adelaide.

Obituary. Harold M. Cooper — 1886-1970. Mankind. 7:

314-315.

1971

Tindale, N. B. & George, B. ‘The Australian

Aborigines’. Lloyd O’ Neill: Windsor.

176 P. G. JONES

1972

The Pitjandjara. Pp. 217-268 in ‘Hunters and Gatherers

Today’. Ed. M. G. Bicchieri. Holt, Rienhart &

Winston: New York.

1972

Curtain, C. C., Tindale, N. B., et al. Distribution of the

immunoglobulin markers at the IgGI, IgG2, IgG3,

IgA2, and K-chain loci in Australian Aborigines:

comparison with New Guinea populations. American

Journal of Human Genetics. 24: 145-155.

1974

Tindale, N. B. ‘Aboriginal tribes of Australia, their

terrain, environmental controls, distribution, limits,

and proper names’. With four sheet map. University

of California Press: Berkeley and Canberra.

Aboriginal tribes of Australia...simplified version of map,

the 601 tribes. Sydney Morning Herald. Special

Supplement. Australia Unlimited. July 15, 1974, p.50.

Notes for an introduction to a Festschrift in honour of

Joseph B. Birdsell. Submitted May 1974 to Dr. Mai,

ULC IL SN,

1975

‘Aboriginal tribes of Australia. Geographic II spelling

version of 4 sheet map’. Australian Institute of

Aboriginal Studies: Canberra.

Introduction. ‘Time before Morning. Art and Myth of

the Australian Aborigines’ by Louis A. Allen. Thomas

Y. Crowell Co.: New York.

1976

Adaptive significance of the Panara or grass seed culture

of Australia. Pp. 345-349 in ‘Stone Tools as Cultural

Markers’. Ed. R.V.S. Wright. Australian Institute of

Aboriginal Studies: Canberra. .

Notes of a few Australian Aboriginal concepts. Pp. 156—

163 in ‘Australian Aboriginal Concepts’. Ed. L.R.

Hiatt. Australian Institute of Aboriginal Studies:

Canberra.

1977

Further report on the Kaiadilt people of Bentinck Island,

Queensland. Pp. 247-273. in ‘Sunda and Sahul’. Ed.

J. Allen et al. Academic Press: London.

1978

Aboriginal landscape of the Lower Murray Valley, South

Australia. With map. South Australian Department of

Lands, [and] South Australian Museum: Adelaide.

Role of Aboriginal people from the Lower Murray in

helping to ensure that records of their traditional life

would be preserved in the South Australian Museum.

Pp. 8-15 in ‘Unesco Regional Seminar. Excursion

Guide’. South Australian Museum: Adelaide.

1981

The Aborigines: an introduction. Pp. 1745-1748 in

‘Ecological Biogeography of Australia’. Ed. A. Keast,

Junk: The Hague.

Prehistory of the Aborigines: some interesting

considerations. Pp. 1763-1795 in ‘Ecological

Biogeography of Australia’. Ed. A. Keast. Junk: The

Hague.

1982

A South Australian looks at some beginnings of

archaeological research in Australia. Aboriginal

History 6(3): 93-110.

1983

The Woakwine Terrace in the South East of South

Australia and indications of the very early presence of

man. Pp. 583-600 im ‘Quaternary Coastlines and

Marine Archaeology’. Ed. P. M. Masters & N. C.

Flemming. Academic Press: London.

1986

Anthropology. Pp. 235-249 in ‘Ideas and Endeavours —

The Natural Sciences in South Australia’. Eds. C. R.

Twidale, M. J. Tyler & M. Davies. Royal Society of

South Australia: Adelaide.

Milerum. ‘Australian Dictionary of Biography’ vol. 10:

498-99.

1987

The wanderings of Tjilbruke: a tale of the Kaurna people

of Adelaide. Records of the South Australian Museum

20: 5-13.

Kariera views on some rock engravings at Port Hedland,

Western Australia. Records of the South Australian

Museum 21: 43-63.

P.G. Jones, South Australian Museum, North Terrace, Adelaide, South Australia 5000. Records of the South

Australian Museum 28(2): 159-176.

Joseph B. Birdsell was born in South Bend,

Indiana on 30 March, 1908 and died in Santa

Barbara, California on 5 March, 1994. Known as

‘Jo’ to his colleagues and friends, Birdsell spent

his academic years mostly at the University of

California, Los Angeles, where he taught from

1947 to his retirement as Professor of

Anthropology in 1974. His scholarly life as a

physical/biological anthropologist started with a

Ph.D. from Harvard in 1941 where he was trained

by Hooton, Boyd and later influenced by Clyde

Kluckhohn. His work and publications spanned

five and a half decades of fieldwork in Australia,

teaching at UCLA and writing on a broad range

of subjects. Two valuable retrospective

commentaries are available on his work. A

volume of essays in his honor appeared in 1981,

and Shanklin’s and Mai’s (1981) article provides

an overview of his life and an appraisal of his

conceptual work. Birdsell (1987) himself, also

offered discussion of major intellectual influences

on his scholarly growth in a general review which

appeared about eight years ago.

For those of us who had Jo as a teacher, the

experience conveyed the power of his ability to

relate empirical facts to model building to

understanding how evolution operates as a broad

orchestration of Darwinian principles in micro-

OBITUARY

JOSEPH B. BIRDSELL

30 March 1908 — 5 March 1994

evolutionary changes, and to the analyses of issues

as diverse as race, racism and population ecology.

This obituary discusses Birdsell’s contribution to

the study of the Australian Aboriginal, his overall

impact on biological anthropology and the kinds

of influences which his writings have had on the

field, and finally my own personal reflections on

the man and his ideas.

Birdsell’s fieldwork and publications on the

Australian Aboriginal were always linked with his

colleague Norman B. Tindale. It was Tindale

through Hooton who brought up the idea for

Birdsell to work in Australia in 1938, the start of

the first Harvard-Adelaide joint venture. It was

during this two year period that Birdsell and

Tindale collected a vast range of data from blood

samples to phenotypical features on a broad

spectrum of Aboriginal populations from

Queensland to Tasmania and Western Australia.

Later, in the early 1950s, both scholars carried out

two more years of fieldwork which focused on the

Kimberley and the northwest part of Australia.

The Birdsell-Tindale collaboration, which lasted

for nearly fifty years, permitted each scholar to

develop a diversity of theoretical and empirical

concerns about which they could exchange ideas

and mutually play off one another. Birdsell’s

commitment to empirically grounded theory

178

represents a lifelong development which is

initially apparent in his 1941 dissertation from

Harvard and continues through to his recent

massive 455 page volume which summarises

more of his empirical findings an Australia

(Birdsell 1993).

Birdsell’s contributions to the biological

analysis of Aboriginal Australians covers three

areas: the trihybrid theory of origin, the issue of

dating Homo sapiens’ first entrance to continental

Australia, and a focus on the continental level of

evolutionary developments. Prior to Birdsell’s

analysis, the overwhelming view was that the

Australian Aboriginal was a single, homogeneous

population which spread throughout the continent,

and the dating indicated recent human arrival in

Australia possibly as recently as 5 000 to 6 000

years ago. Furthermore, most of the physical

anthropology of Australia stressed only local and

regional developments in the fossil record.

Birdsell saw the problem along broader

continental perspectives which related the

particular to the whole of Australia and also

compared the Australian situation to population

processes in southeast Asia, East Asia and Africa.

In identifying the problem of origins as one of

micro-evolutionary differences related to

differential ecological and population forces,

Birdsell argued that the category ‘Australoid’ was

simply wrong and should be abandoned. In his

writings in the 1940s and also in his 1993

volume, Birdsell claimed that Australia was

inhabited by three waves of population expansion:

the Barrinean wave with Negritic features similar

to some southeast Asian populations, the

Murrayian wave with affinities to Ainu-like

groups (the original inhabitants of Japan), and the

Carpentarians with Veddoid features. The

trihybrid hypothesis addressed the concern that the

previous Australoid categorisation embraced

significant internal variation thus blurring micro-

evolutionary processes as well as other hypotheses

of internal migration within Australia.

While proposing the trihybrid theory as it

related to broader processes within the Asian

context, Birdsell also speculated that early man

arrived in Australia at least 35 000 years ago if

not earlier. In the 1940s when Birdsell suggested

this early inhabitation of Australia, his claim was

anathema to most scholars in Australia. There

were no C14 dates, very few fossil remains and

the commonly accepted scholarly idea was that

humans entered Australia only 6 000 years ago.

Both of these opinions made Birdsell and his

views totally unacceptable to biological

A. YENGOYAN

anthropologists and comparative anatomists in

Australia. When I arrived in Sydney in 1966 to

start fieldwork in the Centre, I was asked by many

academicians at the University of Sydney and

elsewhere what had stimulated my interest in

Australia. Although I had a Ph.D. from the

University of Chicago based on fieldwork among

the Mandaya of southeast Mindanao, Philippines,

it was Jo Birdsell who had whetted my intellectual

appetite for working in Australia. When I

mentioned Birdsell’s name, I was simply laughed

at, no one took Birdsell seriously. This attitude

held for most Australian Aboriginal scholars

(except for W. E. H. Stanner). Late in the summer

of 1966/1967, I met A. A. Abbie at Amata in the

Musgrave Ranges. When Abbie heard me talk

about Birdsell as pivotal to my working in

Australia, he quickly and quietly walked away.

However, by 1974 when C14 dating indicated that

30 000 B.P. was plausible for the earliest date of

human entrance into Australia, Birdsell and

Tindale were honored with a conference held in

Canberra, and Birdsell was asked to spend a year

at the Australian National University. Yet, by the

late 1970s, even Birdsell’s estimates were viewed

with caution, since C14 dates had pushed human

entrance back to 45 000 to 50 000 B.P.

Birdsell’s early speculations and visions were

based on a well developed idea of population

genetics combined with migration theory which

stressed the rapidity by which early hominoid

forms could move from South and East Africa to

the Asian mainland. These interests indicated that

Birdsell saw local and regional changes and

developments from a broader continental and even

global perspective. Birdsell’s (1951) lengthy

article on the peopling of the Americas applied

the same perspective by attempting to extrapolate

what could be said about the Americas from Asia.

To this day, most of his generalisations have been

validated, with subsequent archaeological research

supporting his early speculations, just as they did

within the Aboriginal Australian context. C14

dates from the Bering Straits, as well as those |

from the extreme tip of South America, add |

credence to his early ‘educated speculations’.

Although Birdsell’s lifelong interest in

Australia was well anchored in the history of

physical and biological anthropology in the United

States, his intellectual aim even as early as the

systematics from biology and genetics to physical |

anthropology, where for decades the dominant |

concern was primarily taxonomic and |

phenotypical analysis which had no bearing on |

OBITUARY - JOSEPH B. BIRDSELL

ideas of evolutionary biology. This new paradigm

meant that physical and anthropologists had to

turn to the works of Mayr, Dobzhansky, Haldane,

Simpson and Fisher as they established a true and

vital evolutionary basis for the study of human

populations. But of all the evolutionary biologists

who were critical to the new physical

anthropology, it was Sewall Wright’s writings

which had the greatest impact on how Birdsell

understood micro-evolutionary processes and the

extent to which population genetics related to

broader evolutionary forces. Birdsell’s fifty-five

years of analysis and writing all addressed the

multi-facets of evolutionary issues and

Mendelianism as Wright envisioned the problem,

and Birdsell’s comprehensive 1993 volume is

dedicated to Wright.

As a biological anthropologist representing the

new systematics in biology, Birdsell’s writings

represent a broad range of interests covering

population genetics, primate studies, racial

analysis, environmental regulation among hunting

and gathering populations, analysis of the

paleontological records along spatial axioms, and

the rigorous application of modeling to

understanding population processes and

diachronic changes. The application of population

genetics to human societies was most problematic

when Birdsell first entered academia. Fruit flies

and controlled populations were ideal to ascertain

how evolutionary forces worked in a population,

but the human scene, even hunting and collecting

societies, was too complex and variable as a unit

of analysis. Jo saw the problem as one which

required both a strict conceptual definition of

population and a realisation that certain

_ evolutionary processes simply could not be limited

to the population per se. Thus, his use of genetic

space and gene flow models was the beginning of

cline analysis in small populations in which

spatial models on a regional or continental scale

were necessary for understanding microlevel

changes. Furthermore, Birdsell’s investigation of

international drift and mutational processes

among western desert Aboriginal groups was

among the first direct application of Sewall

Wright’s (1931, 1939, 1978) theories to human

populations. Until Birdsell’s analysis, genetic drift

) has been employed as a factor only if no other

evolutionary process could explain the observed

variation. Also following Mayr (1942), Birdsell

pioneered in understanding the dynamics of the

founder effect among small human societies.

Systematics in biology reached a new plateau

when Birdsell took their insights and re-thought

the genetic basis of human societies.

In 1953, Bartholemew and Birdsell published a

pivotal and now classic article on protohominids.

Although this analysis was not empirical, it

established the agenda in regard to how

comparative primate studies should be

approached. By establishing a baseline from data

on non-human mammals and comparing this with

hunting and gathering societies, the authors

created a framework of the biological and minimal

cultural attributes which protohominids must have

expressed. Over the past four decades, primate

studies have filled in the empirical foundations

through field studies, and, Birdsell’s plea for

integrating population genetics and social ecology

resonates in each particular case study. Even the

interest in vocal behaviour is found in this central

piece, and the past three decades again

demonstrate the clarion call of this highly creative

and venturesome kind of thought.

Birdsell’s form of ecological analysis, especially

in analysing the Australian data, indicated that

certain demographic and environmental

regulations were present. Spacing mechanisms,

both spatial and demographic, were essential in

determining how hunters and gatherers survived.

A number of his papers, the most general one

being in the ‘Man the Hunter’ (1968) volume, set

forth the essential dynamics of spacing, one of

which was preferential female infanticide.

Infanticide is common in hunting and gathering

societies, however, the stress on preferential

female infanticide must be revised and questioned

as a universal feature among such societies (see

Yengoyan, 1981). Nevertheless, it was Birdsell’s

direction and thinking along such lines that has

stimulated reconsideration of such issues.

Birdsell’s stress on population characteristics

(fertility, fecundity, mortality rates, morbidity,

population control, the population pyramid, rates

of in-migration and out-migration) was based on

the assumption that demographic variables are the

connecting link between environmental factors (eg

rainfall) and social organisation/structure. This

type of ecological analysis was markedly different

from Julian Stewart’s idea of cultural ecology, and

in most respects avoided the pitfalls of Stewart’s

conception of the culture core.

His interest in population dynamics was not

limited to the Australian case. His 1953 paper

drew the connection between mean annual rainfall

and Aboriginal population densities in Australia,

and his analysis has been duplicated in other areas

of the world such as the Great Basin in the

western United States. In my opinion, Birdsell’s

180

most original and creative thinking about

paleontological issues was published in 1957. By

working through various models of population

dispersion and the effects of how and when a

population buds off from a parental population

into unoccupied lands, Birdsell first asked the

question: ‘how fast could early man occupy

continental Australia?’ Using a relatively

conservative assumption that populations would

double in each generation when entering

unexploited territories, Birdsell concluded that

early man could have spread throughout Australia

(and Tasmania) within a period of 7 000 years,

which in evolutionary terms is virtually nothing.

From this framework, Birdsell asked how fast

would the Intrinsic Rate of Increase (IRI) be from

South/East Africa through the Middle East, India,

Southeast Asia and to Australia? He concluded

that the time differential would have been from 22

000 to 25 000 years, which again is an

evolutionary split-second. Thus, the old idea that

isolation and slow population growth were

barriers to human expansion had to be dismissed

simply from what is known of population and

demographic processes. This particular paper has

led to a rethinking of the whole fossil-man story,

and the picture is still far from complete, but the

essential parameters were set forth by Jo in his

usual insightful way, a mix of good empirical

observations combined with a creative and fertile

approach to problems. Without this essential

creativity, palaeontologists simply could not

analyse their data productively, even if the fossil

record were nearly complete.

Birdsell’s collaboration with Carleton Coon and

Stanley Garn (1950) represents the first major

break in the study of race and racial formation.

Under Hooton and other physical anthropologists

throughout the first five decades of this century,

the notion of race was based on a set of

phenotypical features which could be measured

and compared, but these features had little or no

adaptive utility. Thus, Hooton conceived a race as

a cluster of non-adaptive features based primarily

on forces of inheritance, although virtually nothing

was known about inheritance and genetic

transmission. While the distribution of blood type

alleles was known to manifest different

proportions according to race, as a taxonomy,

racial classification was dealt with in a virtually

non-biological way.

The Coon, Garn and Birdsell approach moved

away from gross phenotypical contrasts and

stressed race as a continuing historical process of

adaptation to particular broad environmental

A. YENGOY AN

forces. Furthermore, race was treated as a micro

breeding unit through which one could show how

various markers are transmitted as specific genetic

features which ideally can be related to gene

frequencies. It was on the level of local Mendelian

populations or breeding units that race manifested

and expressed evolutionary processes. In the

1940s, these ideas were highly important, since

for the first time they moved the study of racial

formation to specific environmental and biological

parameters which acted selectively with the

genotypoe as well as certain phenotypical features.

My initial contact with Jo Birdsell was in 1956

when I enrolled in the Department of

Anthropology as the first year graduate student at

ULCA. Coming in with only three undergraduate

anthropology courses meant that virtually

everything I encountered was new, exciting and

challenging, and during my two years at ULCA I

took Jo’s undergraduate courses as well as his

seminars. Jo’s style of lecturing was inspiring to

all his students in that he conveyed a deep

knowledge of the subject matter combined with

an openness of ideas which provoked students to

challenge him on the spot. Above all, he insisted

that we work through ideas and clusters of data in

a method in which “educated speculation”

emerged as a means of seeing new problems

derived from older and possibly less interesting

issues. It was in these seminars that we had a

chance to work through various ideas which he

was developing and which soon appeared in the

journals.

Furthermore, a few of us worked for him

tabulating his data from his first field trip to

Australia as well as his last one in the early

1950s. Clyde Wilson, Robert Littlewood and I

spent hours in the late afternoons, evening and

sometimes on the weekends tabulating all the

findings on a single variable. It was this

experience which brought forth Jo’s insistence that

incorporating maximum biological, social and

interactional data on a genealogical grid was the

most essential strategy in dealing with a vast

range of data. His genealogies were impressive,

and, being a neophyte, I was always wondering

what it meant and where was it going. Thus he

was a gifted teacher, one who was articulate

beyond expectation, one who knew his subject

matter, and one who knew how to think about his

data in a venturesome and creative way. At the

same time we all sensed that his broad and bold

thinking would culminate in a work which was

readily approachable. In 1972 Birdsell published

a textbook titled ‘Human Evolution’, a book

OBITUARY - JOSEPH B. BIRDSELL

which has since gone through various editions.

Reading Jo’s textbook is equivalent to hearing

vintage Birdsell in person. Ideas abound on the

pages, the case studies and data are fascinating,

and his ability to convey a broad spectrum of

issues, concerns and doubts in an engaging way

express an intellectual love of his subject matter

that never waned. :

Two different styles of Jo’s pedagogy should be

noted. First was his insistence that empiricism

was fundamental to what biological

anthropologists must set forth as theories or as

hypotheses. Models and the craft of modeling

were based on empirical knowledge, but Birdsell

always felt that empiricism could only take one so

far. There are some limits to explanation and

interpretation intrinsically due to a lack of

information, but the wisdom and experience that

students received from him were that one need not

be curtailed and/or limited by empiricism.

Educated speculations were as valid as pure

empirical information, and in most cases the

speculations were more interesting and led to a

discussion of new problems and future kinds of

research.

Second was Jo’s insistence on examining

problems, issues and debates in a different and

unique way. Here Jo would ask what an event,

thing, theory or strategy could not be. By focusing

on conditions which either empirically and/or

logically could not be the case, we were forced to

imagine the range of all possibilities, even if some

of these simply could not occur. In each case, the

feasibility of the occurrence of an event or thing

compelled seminar participants to show the

empirical basis and/or logic relating to the

presence and absence of factors. In contemporary

philosophy this form of analysis is called

counterfactual thinking, and it has had an

important effect on the way that philosophical and

logical arguments are crafted. Birdsell understood

this form of thought well before it came into

vogue. Nearly four decades after that two year

experience with Jo, the impact of his thought and

the way he saw problems is still central to my

own work.

After finishing an MA at UCLA in 1958, I

moved on to the University of Chicago to pursue

my doctorate in social anthropology. For the

following thirty to forty years, Jo and I were in

communication on various Australian matters,

especially every time I returned from Australia.

It goes without saying that he will be sorely

missed by his friends and former students. The

experience of his seminars, of seeing him craft an

argument, either on paper or verbally, and the kind

181

of deep probing which he asked of himself and his

students will all be missed. Those of us who

experienced the man hold him in deep

appreciation and are much in his debt, for there

will never be another ‘Jo’ in our midst.

SELECTED BIBLIOGRAPHY

BIRDSELL, J. B. 1940. Field impressions of an

ethnologist [abstract]. Mankind, 2 (8):277.

BIRDSELL, J. B. 1941. The trihybrid origin of the

Australian Aborigines. Doctoral dissertation filed in

the Peabody Museum, Harvard University,

Cambridge, Massachusetts.

BIRDSELL, J. B. 1941. A preliminary report on the

trihybrid origin of the Australian Aborigines.

American Journal of Physical Anthropology

[abstract], 28: 6.

BIRDSELL, J. B. 1947. New data on racial analysis.

American Journal of Physical Anthropology

[abstract] (ns), 5(2): 232.

BIRDSELL, J. B. 1947. Genes, P-genes and racial

analysis. American Journal of Physical Anthropology

[abstract] (ns), 5(2):237.

BIRDSELL, J. B. 1949. The racial origins of the extinct

Tasmanians. Records of the Queen Victoria Museum,

2(3): 105-122.

BIRDSELL, J. B. 1950. Some implications of the genetic

concept of race in terms of spatial analysis. Pp. 259-

314 in ‘Origin and Evolution of Man’. Cold Spring

Harbor Symposia on Quantitiative Biology, Vol. 15,

Cold Spring Harbor: Long Island, NY.

BIRDSELL, J. B. 1950. A collaborative genetical survey

of the human populations of the Pacific areas.

Science, 112(2897): 25-26.

BIRDSELL, J. B. 1951. The problem of the early

peopling of the Americas as viewed from Asia. Pp. 1—

68a in ‘Papers on the Physical Anthropology of the

American Indian’. Ed. W. S. Laughlin, Edwards

Brothers Inc.: Ann Arbor.

BIRDSELL, J. B. 1951. Review of Genetics and the

Races of Man. (W. C. Boyd). American Journal of

Physical Anthropology (ns), 9: 219-223.

BIRDSELL, J. B. 1952. On various levels of objectivity

in genetical anthropology. American Journal of

Physical Anthropology (ns), 10(3): 355-362.

BIRDSELL, J. B. 1953. Some environmental and

cultural factors influencing the structuring of

Australian Aboriginal populations. American

Naturalist, 87(334): 171-207.

BIRDSELL, J. B. 1957. Some population problems

involving Pleistocene man. Population Studies,

Animal Ecology and Demography. Cold Spring

Harbor Symposia on Quantitative Biology, 22: 47-

69.

BIRDSELL, J. B. 1957. On methods of evolutionary

182

biology and anthropology. Part Il, Anthropology.

American Scientist, 45(5): 393-400

BIRDSELL, J. B. 1958. On population structure in

generalized hunting and collecting populations.

Evolution 12(2): 189-205.

BIRDSELL, J. B. 1959. ‘Australian Bibliography’ (area

22). Council of Old World Archaeology (COWA),

Cambridge.

BIRDSELL, J. B. 1963. Review of: ‘The Origin of

Races’, by C. S. Coon, The Quarterly Review of

Biology, 38(2): 178-185.

BIRDSELL, J. B. 1966. Comment on: Population

distances: Biological, linguistic, geographical and

environmental by W.W. Howells. Current

Anthropology, 7(5): 531-540.

BIRDSELL, J. B. 1967. Preliminary data on the

trihybrid origin of the Australian Aborigines.

Archaeology and Physical Anthropology in Oceania

2(2), 100-155.

BIRDSELL, J. B. 1968. Some predictions for the

Pleistocene based on equilibrium systems among

recent hunters. Pp. 229-40 in ‘Man the Hunter’. Eds.

R. Lee and I. DeVore, Aldine Press: Chicago.

BIRDSELL, J. B. 1970. Local group composition among

the Australian Aborigines; a critique of the evidence

from field work conducted since 1930. Current

Anthropology 11(2): 115-142.

BIRDSELL, J. B. 1971. Australia: ecology, spacing

mechanisms and adaptive behaviour in Aboriginal

land tenure. Pp. 334-361 in ‘Land Tenure in the

Pacific’ . Ed. R. C. Crocombe. Oxford University

Press: Melbourne.

BIRDSELL, J. B. 1972. ‘Human Evolution’. Rand

McNally: Chicago.

BIRDSELL, J. B. 1972. The problem of the evolution of

human races: classification or clines? Social Biology

19: 136-162.

BIRDSELL, J. B. 1973. A basic demographic unit.

Current Anthropology 14 (4): 337-356.

BIRDSELL, J. B. 1975. A preliminary research on man-

land relations in aboriginal Australia. American

Antiquity, Memoirs, 40 (2), pt. 2 mem. 30: 34-37.

BIRDSELL, J. B. 1976. Realitites and transformations:

The Tribes of the Western Desert of Australia. Pp.

95-120 in ‘Tribes and Boundaries in Australia’. Ed..

Nicolas Peterson, Australian Institute of Aboriginal

Studies. Social Anthropology Series, Humanities

Press: New Jersey.

BIRDSELL, J. B. 1977. The recalibration of a pradigim

for the first peopling of Greater Australia. Pp. 113-67

in ‘Sunda and Sahul’. Eds. J. Golson and R. Jones,

Academic Press: New York.

BIRDSELL, J. B. 1978. Spacing mechanisms and

adaptive behaviour of Australian Aborigines. Pp. 213—

44 in ‘Population Control by Social Behaviour’. Eds.

F. J. Ebling and D. M. Stocklart, Institute of Biology,

London.

A. YENGOY AN

BIRDSELL, J. B. 1979. Ecological influences on

Australian Aboriginal social organisations. Pp 117—

151 in ‘Primate Ecology and Human Origins’. Eds. I.

S. Bernstein and E. O. Smith, Garland STPM Press:

New York.

BIRDSELL, J. B. 1979. Physical anthropology in

Australia today. Pp. 417-30 in ‘Annual Review of

Anthropology’, Vol. 8. Eds. B. J. Siegel, A. B. Beals,

and S. A. Tylor, Annual Reviews Inc.: Palo Alto.

BIRDSELL, J. B. 1987. Some Reflections on Fifty Years

in Biological Anthropology. Annual Review of

Anthropology. 16: 1-12.

BIRDSELL, J. B. 1993. ‘Microevolutionary Patterns in

Aboriginal Australia’. Oxford University Press: New

York.

BIRDSELL, J. B., & BARTHOLEMEW, G. A. Jr. 1953.

Ecology and the Protohominids. American

Anthropologist, 55: 481-498.

BIRDSELL, J. B. & BOYD, W. 1940. Blood groups in

the Australian Aborigines. American Journal of

Physical Anthropology 27(1): 69-90.

BIRDSELL, J. B. with COON, C. S., GARN, S. 1950. A

Study of the Problem of Race Formation in Man.

American Lecture Series, Publication No. 77. Charles

C. Thomas: Springfield, Illinois.

BIRDSELL, J. B. & SIMMONS R. T., & GRAYDON,

J.J. 1953. High R* frequency in the blood of

Australian Aborigines. Nature, 172: 500.

BIRDSELL, J. B., SIMMONS, R. T. & GRAYDON, J.

J. 1976. Micro-differentiation in blood group gene

frequencies among twenty-eight adjacent aboriginal

tribal isolates in Western Australia. Australian

Institute for Aboriginal Studies. Occasional Papers

in Human Biology, 2: 1-38.

BIRDSELL, J. B., SIMMONS, R. T. & TINDALE, N. B.

1962. A blood group genetical survey in Australian

aborigines of Bentinck, Mornington and Forsythe

Islands, Gulf of Carpentaria. American Journal of

Physical Anthropology, 20(3): 303-320.

BIRDSELL, J. B. & TINDALE, N. B. 1941. Results of

the Harvard-Adelaide Universities Anthropological

Expedition, 1938-39: Tasmanoid tribes in Northern

Queensland. Records of the South Australian

Museum, 7(1): 1-9.

IN PRESS

BIRDSELL, J. B. & MAI, L. L. (ms). Heat stress in

Australian Aboriginal man, together with some

evolutionary complications. [Submitted February

1995 to the American Journal of Physical

Anthropology].

REFERENCES

BIRDSELL, J. B. 1951. The problem of the early

OBITUARY — JOSEPH B. BIRDSELL

peopling of the Americas as viewed from Asia. Pp. 1-

68a in ‘Papers on the Physical Anthropology of the

American Indian’. Ed. W. S. Laughlin, Edwards

Brothers Inc.: Ann Arbor.

BIRDSELL, J. B. 1968. Some predictions for the

Pleistocene based on equilibrium systems among

recent hunters. Pp. 229-40 in ‘Man the Hunter’. Eds.

R. Lee and I. DeVore, Aldine Press: Chicago.

BIRDSELL, J. B. 1987. Some Reflections on Fifty Years

in Biological Anthropology. Annual Review of

Anthropology. 16: 1-12.

BIRDSELL, J. B. 1972. ‘Human Evolution’.

McNally: Chicago.

BIRDSELL, J. B. 1993. “Microevolutionary Patterns in

Aboriginal Australia’. Oxford University Press: New

York.

BIRDSELL, J. B., & BARTHOLEMEW, G. A. Jr. 1953.

Ecology and the Protohominids. American

Anthropologist, 55: 481-498.

BIRDSELL, J. B. with COON, C. S., GARN, S. 1950. A

Study of the Problem of Race Formation in Man.

American Lecture Series, Publication No. 77. Charles

C. Thomas: Springfield, Illinois.

MAYR, E. 1942. ‘Systematics and the Origin of

Species’. Columbia University Press: New York.

SHANKLIN, E. & MAI, L. L. 1981. Joseph B. Birdsell:

A Conceptual Biography. Pp. 21-53 in ‘The

Perception of Evolution: Essays Honoring Joseph B.

Birdsell’. Eds. L. L. Mai, E. Shanklin, & R. W.

Sussman. University of California: Los Angeles.

WRIGHT, S. 1931. Evolution in Mendelian Populations.

Genetics 16: 97-159.

Rand

183

WRIGHT, S. 1939. Statistical genetics in relation to

evolution. Pp. 5-64 in ‘Actualities Scientific et

Industrielle’. No. 802. Hermann et Cie: Paris.

WRIGHT, S. 1978. Variability Within and Among

Natural Populations ‘Evolution and Genetics of

Populations’. Volume 4. University of Chicago Press:

Chicago.

YENGOYAN, A. A. 1981. Infanticide and Birth Order.

An Empirical Analysis of Preferential Female

Infanticide Among Australian Aboriginal Populations.

Pp. 255-273 in ‘The Perception of Evolution: Essays

Honouring Joseph B. Birdsell’. Eds. L. L. Mai, E.

Shanklin, & R. W. Sussman. University of California:

Los Angeles.

UNFINISHED MANUSCRIPTS

BIRDSELL, J. B. (ms). The Ituri Pygmies: No to the

Mull Hypothesis.

BIRDSELL, J. B. (ms). Englishmen and Australian

Aborigines, 1788-1989.

BIRDSELL, Joseph B. (ms) The biology of inheritance

in Australian-European hybrids. [This manuscript has

not been located, although it is mentioned in his will

as a book-length companion monograph to

Microevolutionary Patterns..., which was published by

Oxford in 1993. Birdsell (1987) also commented that

“the all-important hybrid series containing almost 400

first-generation crosses between European and

Aborigines, is just now undergoing analysis...”

(pg.9)].

A. YENGOYAN, Department of Anthropology, University of California, Davis, California 95616, USA. Records of

the South Australian Museum 28(2): 177-183.

THE WATER SKINKS (LACERTILIA: EULAMPRUS) OF VICTORIA

AND SOUTH AUSTRALIA

MARK N. HUTCHINSON & PETER A. RAWLINSON

HUTCHINSON, M. N. & RAWLINSON, P. A. 1995. The water skinks (Lacertilia: Eulamprus)

of Victoria and South Australia. Records of the South Australian Museum 28(2): 185-207.

The taxonomy, biology and distribution of the Victorian and South Australian water skinks

of the Eulamprus quoyii species-complex are reviewed. A lectotype is designated for Scincus

vittatus Quoy & Gaimard, 1824. Five taxa are recognised: E. quoyii Duméril & Bibron from

the lower Murray River valley and Mt Lofty Ranges, E. heatwolei Wells & Wellington from

warm temperate eastern Victoria and the Fleurieu Peninsula, E. kosciuskoi Kinghorn from

alpine northeastern Victoria and E. tympanum Lénnberg & Andersson, the latter with two

subspecies, E. t. tympanum (including E. herseyi Wells & Wellington, 1985), widespread

through cool temperate habitats and E. t. marnieae ssp. nov., restricted to the stony rises east

and north of Lake Corangamite where its survival is threatened by habitat degradation.

M. N. Hutchinson, South Australian Museum, North Terrace, Adelaide, South Australia 5000,

and P. A. Rawlinson', Department of Zoology, La Trobe University, Bundoora, Victoria 3083.

Manuscript received 31 October, 1994.

The water skinks form a group of closely related

medium sized to moderately large (SVL to 118

mm) Australian lygosomine scincid lizards that

inhabit the margins of watercourses throughout

the coastal drainage systems of eastern and

southeastern Australia (excluding Tasmania).

Water skinks are conspicuous, active diurnal

heliotherms (Spellerberg 1972b) which readily use

water as a refuge, swimming on and under the

surface (Daniels & Heatwole 1990).

Traditionally these lizards have been placed in

Sphenomorphus, either as a subgenus of the

catchall Lygosoma (Smith 1937) or, following

Loveridge (1934) and Mittleman (1952), as a full

genus. Recent Australian usage (Cogger 1992,

Greer 1989, 1992) has recognised the paraphyletic

nature of Sphenomorphus (see Greer 1979a, Greer

& Parker 1967) by employing the long-disused

name Eulamprus Fitzinger (type species quoyii)

for the larger Australian viviparous

‘Sphenomorphus’, including the E. quoyii

complex. As yet there has not been a rigorous

phylogenetic assessment of this assemblage, but

Shea & Peterson (1985) and Greer (1989, 1992)

discuss some potentially useful characters which

indicate that Eulamprus is at least more likely to

be a natural unit than the much larger and

obviously heterogeneous assemblage represented

' Peter Rawlinson died in April 1991. His unpublished studies

of type material and his taxonomic insights form the basis of

this paper.

by Sphenomorphus, and it is provisionally

accepted in this paper.

The first described member of this group was

Quoy & Gaimard’s (1824) Scincus vittatus, the

“Scinque a flancs noirs’, from the Sydney region.

Duméril & Bibron (1839) redescribed the species

using the replacement name Gongylus

(Lygosoma) quoyii, and as Lygosoma or

Sphenomorphus quoyii, the water skink has

become a familiar and relatively well-studied

eastern Australian lizard (e.g. King 1964, Veron

1969, Spellerberg 1972a—d, Daniels 1987).

Early workers in Victoria and South Australia

(Lucas & Frost 1894, Waite 1929) were familiar

with a lizard they referred to as Lygosoma quoyii,

although specimens from southern Australia do

not conform in all respects to the east coastal

populations. Two new water skink taxa were

described during the early part of this century,

Lygosoma tympanum from near Melbourne

(Lonnberg & Andersson 1913) and Hinulia quoyii

kosciuskoi from Mt Kosciusko (Kinghorn 1932).

Loveridge (1934), in spite of some uncertainty,

synonymised kosciuskoi with tympanum, treating

the latter as a southern and highland subspecies of

quoyii. Worrell (1963) recognised tympanum as a

full species, distinct from quoyii, but did not query

the synonymy of tympanum and kosciuskoi.

Rawlinson (1969) reported that tympanum and

kosciuskoi were distinct species, and that both

were distinct from quoyii, the three being

collectively referred to as the Sphenomorphus

186

quoyii species complex. He also reported a fourth,

undescribed member of the complex, referred to

as the ‘Warm Temperate’ form of S. tympanum.

The data on which Rawlinson’s conclusions were

based were not presented at the time but these

conclusions have become widely used (Cogger et

al. 1983).

Five species of water skinks are recognised by

Cogger (1992). In addition to Eulamprus quoyit,

E. tympanum and E. kosciuskoi, E. heatwolei

Wells & Wellington (1984) is applied to New

South Wales populations of Rawlinson’s ‘Warm

Temperate’ form of E. tympanum (Shea &

Peterson 1985, Cogger 1992) while E. leuraensis

Wells & Wellington (1984) is applied to the Blue

Mountains population formerly referred to E.

kosciuskoi.

Shea & Peterson (1985) have summarised data

pertinent to New South Wales water skink

populations but variation in the southerly parts of

their distributions is not well documented and the

distributions themselves are not mapped with

sufficient resolution in Cogger’s books. Much of

what is known, including the taxonomic

distinctiveness of tympanum from quoyii and in

tum of heatwolei from tympanum stems from

unpublished work of the second author. In

addition, a hitherto unreported, morphologically

distinctive population of water skinks has been

found in central southwestern Victoria. The

purpose of this paper is to stabilise the

nomenclature, document the distinguishing

features and geographic distribution of the four

described water skinks in Victoria and South

Australia, and to describe the newly discovered

Victorian form.

MatERIALS AND METHODS

The specimens on which this study is based are

primarily those of the Museum of Victoria (NMV)

and the South Australian Museum (SAMA).

Except where indicated, specimen descriptions are

based only on Victorian and South Australian

specimens. Other Museum abbreviations used

here (following Leviton et al. 1985) are: AMS,

Australian Museum, Sydney; BMNH, Natural

History Museum, London; MNHN, Muséum

Nationale d’Histoire Naturelle, Paris; NHRM,

Naturhistoriska Riksmuseet, Stockholm; NMW,

Naturhistorisches Museum, Vienna.

Head shield terminology used in this paper is

illustrated in Figure 1. Supraciliaries were counted

to the last scale in contact with both the upper

palpebrals and the supraoculars. Presuboculars are

M. N. HUTCHINSON & P. A. RAWLINSON

the scales between the posterior loreal and the

subocular supralabial. The postsubocular scale

row is formed by the series of scales beginning on

the orbital margin of the postsubocular supralabial

and running dorsally to the posterior supraciliary.

The last infralabial was difficult to distinguish

from adjacent scales when the mouth was closed;

it was identified here as the last scale contacting

the ventral margin of the last supralabial. Scale

counts were made using standard criteria (e.g.

Greer 1982). Shea & Peterson (1985) employed a

different method for counting paravertebral scales

than the one used here: they stopped the count at

the level of the anterior edge of the hind limb,

whereas we followed Greer (1982) in taking the

counts posteriorly to the first scale posterior to a

line level with the posterior edge of the hind limb

held at right angles to the body. This count

estimates the number of scales overlying the trunk

vertebrae (including the sacrals). Measurements

of snout—vent length (SVL), tail length and hind

limb length (HLL) were made to the nearest

millimetre using a ruler. Head width (HW,

measured across temporal jaw muscle bulge) and

head length (HL, measured from snout to anterior

edge of ear opening) were measured using dial

calipers to the nearest 0.1 millimetre. Size at

sexual maturity was estimated from smallest

female with enlarged ova or male with enlarged

testes. Osteological data were obtained primarily

from the water skink skeletal collection assembled

by S. J. Tilley, now in the collection of the

Museum of Victoria.

SYSTEMATICS

SCINCIDAE Gray, 1825

LYGOSOMINAE Mittleman, 1952

Eulamprus Fitzinger, 1843

A group of lygosomines belonging to the

Sphenomorphus Group (Greer, 1979b), sharing

the derived features of moderately expanded

palatal rami of the pterygoids and viviparous

reproduction, but lacking the derived character

states of other members of this Group. Within this

genus, the E. quoyii species group shares four

derived character states: third pair of chin shields

separated by five smaller scales; inguinal fat

bodies absent; distal supradigital scales in a single

row; subdigital lamellae grooved, divided basally

(after Greer 1989).

Other features shared by all species in the

complex are as follows. Nasals separated

medially. Supranasal and postnasal scales absent.

WATER SKINKS 187

preocular

supraciliaries

rostral

1cm

infralabials

presuboculars

suboculars

postoculars,

primary temporal

secondary temporals

postsuboculars

rostral

nasal ve frontonasal

_— mental

i \ prefrontal

PD iN supraoculars

Gh Na

frontoparietals [RAY —~ Ve} vp) “! postoculars

pa pe: parieta

interparietal

gly,

WX ee

See

nuchal

post parietal scales

tertiary temporal

ve Ly

NC SI ee!

Cesare.

FIGURE 1. Head of a water skink (Eulamprus t. tympanum; NMV D50716) showing head shield nomenclature used

in this paper.

Two subequal loreals. A single preocular. Lower

eyelid scaly. Three presuboculars, the third

penetrating downwards in front of the subocular

supralabial. Two or three subocular scales. Four

supraoculars, the first three (rarely first two)

contacting the frontal. Two or three postoculars lie

between the posterior supraciliary and the parietal;

the dorsalmost and often the largest of these could

be regarded as a small fifth supraocular, and its

designation as a postocular here is arbitrary,

reflecting traditional treatment of these species

(e.g. Cogger 1992) as having only four

supraoculars. Frontoparietals paired; interparietal

distinct. Primary temporal small, variable

188

intraspecifically, sometimes scarcely distinct from

postsubocular scales. Upper secondary temporal

longer than deep, contacting lateral margin of

parietal scale and bordered below by single lower

secondary temporal which is deeper than long;

upper temporal may be divided by a vertical suture

and lower by a horizontal suture in a minority of

specimens. First pair of chin shields in broad

median contact; second pair separated by a single

scale. Ear opening large, three-quarters the size of

the eye, its margin smooth-scaled and without

projecting auricular lobules. Median pair of

preanals much larger than lateral preanals.

Water skinks show ontogenetic variation in two

significant scale characters. In _ the

Sphenomorphus Group, each parietal is usually

bordered along its posterior margin by a

transversely enlarged nuchal (medially) and along

its lateral margin by the upper secondary temporal

and an additional large scale intercalated between

the nuchal and the upper secondary temporal.

Most adult water skinks, the alpine species

excepted, lack this scale morphology, having

instead four or five variably enlarged and often

obliquely oriented and asymmetrically arranged

scales filling the gap between the upper secondary

temporal on each side. Neonates (Fig. 2),

however, have the more common lygosomine

arrangement of two transversely aligned nuchals.

Positive allometric growth of the upper secondary

temporal relative to the parietal appears to crowd

secondary temporal

tertiary temporal

nuchal

keeled dorsals

FIGURE 2. Head and forebody of a neonate water skink

(Eulamprus t. tympanum; NMV D13654). showing the

arrangement of post-parietal scales (compare to Fig. 1)

and the keeled dorsal body scales present in juveniles but

lost early in ontogeny.

M. N. HUTCHINSON & P. A. RAWLINSON

the post-parietal scales in adults. Water skinks as

subadults and adults have smooth body scalation

but neonates have all dorsal and lateral body, tail

and limb scales keeled, with up to four low keels

or pustules on each dorsal scale producing a wavy

trailing edge. The scales become smooth and

cycloid as the carination is lost at a snout vent

length of about 40-45 mm.

Presacral vertebrae 26. Phalangeal formula of

manus and pes 2.3.4.5.3 and 2.3.4.5.4

respectively. Postorbital bone present,

intraspecifically variable in dorsal exposure from

elongate and extending to the supratemporal

fenestra to greatly reduced and restricted to the

region of thejugal articulation. Supratemporal

fenestra large. Ectopterygoid without or with

variably developed palatal process which may

extend to the palatine, excluding the pterygoid

from the infraorbital vacuity. Hemipenis elongate

with deep distal bifurcation. Iris of eye black,

indistinguishable from pupil in life.

The water skinks show slight sexual

dimorphism in proportions, females reaching a

slightly larger SVL and males having relatively

longer limbs and larger heads. Within species,

dimorphism becomes more obvious with

increasing body size. Both sexes attain sexual

maturity at similar sizes.

Eulamprus quoyii (Duméril & Bibron, 1839)

(Figs 3-4)

Scincus vittatus Quoy & Gaimard, 1824: 178.

Lectotype (designated herein): MNHN 7112,

Neutral Bay, Port Jackson, New South Wales, F.

Péron. (Junior homonym of Scincus vittatus [=

Mabuya vittata] Olivier, 1804).

Gongylus (Lygosoma) quoyii Duméril & Bibron,

1839: 728. Lectotype (Wells & Wellington 1985):

MNHN 7113, Neutral Bay, Port Jackson, New

South Wales.

Eulamprus quoyii Fitzinger, 1843: 22.

Hinulia quoyii Gray, 1845: 70.

Hinulia gastrosticta Giinther, 1875: 11. Lectotype

(Wells & Wellington 1985): BMNH 1946.8.15.34

‘Queensland’, purchased from G. Krefft.

Lygosoma (Hinulia) quoyii Boulenger, 1887: 230.

Sphenomorphus quoyi Barbour, 1914: 204.

Sphenomorphus quoyii quoyii Loveridge, 1934:

349,

Lygosoma (Sphenomorphus) quoyi Smith, 1937:

220.

WATER SKINKS 189

Eulamprus gastrostictus Wells & Wellington,

1984: 93.

Types

As the oldest available name relating to this

group of lizards, Duméril & Bibron’s (1839)

Gongylus (Lygosoma) quoyii must be confidently

allocated before other names can be applied. Five

syntypes used by Duméril & Bibron are still

identifiable in the MNHN collection, four of

which are members of the E. quoyii species

complex, the fifth (7114) being a specimen of the

Asian species Scincella reevesi. Of the remaining

four, two (7112-13) were collected by Péron and

are identified by the MNHN as syntypes of

Scincus vittatus Quoy & Gaimard. Both have a

colour pattern which includes well-developed

narrow pale dorsolateral lines, matching the figure

provided by Quoy & Gaimard (1824) and the

description of Duméril & Bibron (1839). The

remaining syntypes, 2976 (Port Macquarie, J.

Verreaux) and 2977 (Nouvelle Hollande, Lesson

et Garnot) are conspecific with 7112 and 7113.

Examination of the types thus confirms that the

current taxonomy is correct in applying the name

quoyii to the east-coastal Australian member of

the complex. One of Quoy & Gaimard’s syntypes,

7113, SVL 105 mm, was designated by Wells &

Wellington (1985) as the lectotype of Gongylus

(Lygosoma) quoyii Duméril & Bibron, 1839. This

specimen is not in good condition, the mouth and

neck being badly mutilated; however there is no

doubt of its specific identity . The other syntype,

7112, is in excellent condition and would have

been a better choice. It is hereby designated as the

lectotype of Scincus vittatus Quoy & Gaimard,

1824.

Four syntypes of Giinther’s Hinulia gastrosticta

are in the collection of the Natural History

Museum, London (BMNH 1946.8.4.99 and

1946.8.15.34-36) and two more identified as

‘Typus’ are in the NMW (16656:1-—2). All six are

conspecific with the lectotype of Gongylus

(Lygosoma) quoyii, confirming the correctness of

Boulenger’s (1887) synonymisation of

gastrosticta with quoyii. One of the BMNH

syntypes, 1946.8.15.34 (at 101 mm SVL, the

largest of the four), was designated as lectotype of

Hinulia gastrosticta Giinther, 1875, by Wells &

Wellington (1985) who gave no reason for

resurrecting the species from synonymy, nor did

they provide any distinguishing features. We here

return gastrosticta to the synonymy of quoyii.

Diagnosis

A large water skink (adults reaching over 110

a, iy,

LIE

SABES

FIGURE 3. Head shields of Eulamprus quoyii (SAM

R33009).

mm snout—vent) with sharply-defined narrow pale

yellow dorsolateral stripes but without a black

vertebral stripe.

Description

36-42 (x 39.3, n = 36) longitudinal scale rows

at midbody. Paravertebral scales 74-88 (x 79.7, n

= 36), no larger or only slightly broader than

adjacent dorsals. Subdigital laellae on fourth toe

24-32 (K 27.4, n = 35), most with a median

groove and those at the base of the toe divided.

Prefrontals usually broadly contacting (narrowly

separated in 4 out of 36). Interparietal elongate,

approximately one and a half times as long as

wide, but never separating parietals. Each parietal

bordered posteriorly by one to three nuchal scales

and laterally by the upper secondary temporal.

Supraciliaries 9-12 (x 9.7, mode 9); first to third

or fourth forming decreasing series, next three to

five smallest, last two larger, usually penetrating

dorsally each side of the fourth supraocular.

Supralabials 7-8 (mode 7), fifth or sixth

subocular. Infralabials 7-10 (modes 8 and 9), first

and second always and third sometimes in contact

with (single) postmental.

Premaxillary teeth usually 9 (n = 9); single

specimens each with 7 and 8.

Dimensions (of adults, n = 32). SVL 82-112

mm (X 94.9 mm). HW 11.2-17.2 mm. HL 17.3-

24.9 mm. HL/HW 1.36-1.68, (« 1.53). HW/SVL

0.118-0.177 (x 0.146). HLL 31-43 mm. HLL/

190 M. N. HUTCHINSON & P. A. RAWLINSON

3 7 PS

/§ “i

if,

/ D

Cs,

< Pe

iS Set

4 eS iN

3 7 a \

oO E ee

7) 5 Le

g : |

no. "

(L ap.

E ey

Se)

2 ow Og

; =

o zs

5 =

cae

mein ane

FIGURE 4. Distribution of Eulamprus quoyii and E. kosciuskoi in Victoria and South Australia.

WATER SKINKS 19]

SVL 0.333-0.464 (x 0.401). Tail length/SVL (n =

8) 1.71-1.89 (« 1.80).

Colour (in preservative) light to medium brown

on the dorsal surface of the head, body, tail and

limbs. Dorsum of head and body with a few

irregularly scattered flecks. Limbs brown with

irregular black bars. A narrow, well-defined pale

yellow dorsolateral stripe runs from the

supraciliaries posteriorly to half way along the

back, whence it breaks up and fades before

reaching the hips. This stripe often margined

medially by a narrow black line. Upper lateral

zone black, the colour extending anteriorly to the

ear opening. Two to three uneven series of pale

dots (each covering one to two scales) overlie the

black. Temporals and posterior supralabials

usually obscurely spotted with darker pigment.

Lower lateral zone greyish-yellow with black

flecks tending to align along scale rows to form

weak barring. In adults of both sexes infralabials,

chin shields and throat pale to dark grey, spotted

with cream, the spotting smaller and arranged in

longitudinal lines on the throat. Juveniles and

subadults lack this pattern, having instead uniform

greyish-yellow throat colouring. Underside

yellowish white with fine black dots.

In life the colour is similar to that in

preservative.

Distribution

The Murray River from its junction with the

Darling downstream to about Tailem Bend, South

Australia (Fig. 4). A disjunct population occurs in

the Mt Lofty Ranges. Extralimitally the species

occurs along the Darling River and along the east

coast of Australia from Cairns, Qld, south to about

Jervis Bay, New South Wales.

Ecology

Eulamprus quoyii is only found adjacent to

water, living beside permanent lakes, swamps and

billabongs or on the banks of perennial streams,

typically being observed on logs or rocks

emerging from the water’s edge. It appears to have

colonised this part of its range by expansion over

the Great Divide into the headwaters of the

Darling system, and along its course to the

Murray. This species thus extends into a

climatically unsuitable (arid) environment by

restricting its activity to the mesic riparian

corridors (Littlejohn & Rawlinson 1971).

Aspects of the ecology of E. quoyii have been

examined by Veron (1969), Spellerberg (1972b—

d), Daniels (1987) and Daniels & Heatwole

(1984, 1990), all studies relating to humid, warm

temperate, east coastal populations.

Females ovulate late October-November, with

litter size (oviducal eggs) ranging 2-5 (x 4.0, n=

5) in Murray Valley and Adelaide Hills

specimens. Testes are enlarged between April and

August, and regress over spring to a minimum in

November—January. The species is viviparous, the

young being born during January and February.

Mating has been recorded by Veron (1969) as

occurring in spring.

Discussion

This species is the type of Fitzinger’s genus

Eulamprus, and Gray’s Hinulia (Mittleman,

1952), and featured in several early studies of

lizard anatomy (Siebenrock, 1892, 1895, Busch

1898). More recently it has been subjected to a

variety of anatomical and physiological studies

(King 1964, Daniels 1985, Daniels, Heatwole &

Oakes 1987, Daniels, Oakes & Heatwole 1987).

Eulamprus quoyii is much the largest of the

species in the E. quoyii complex, with a mean

adult SVL of 95 mm and a maximum of 118 mm

(extralimitally), both values being roughly 15 mm

greater than the corresponding values for the next

largest species (E. heatwolei and E. tympanum). It

shares with E. heatwolei the most gracile

proportions seen in the group, with the longest

extremities and slenderest head.

Distribution in the area considered is limited to

the Murray River valley and three west-flowing

river systems (Torrens, Sturt and Onkaparinga) of

the adjacent Mt Lofty watershed. The grey and

cream mottled throat pattern is more weakly

developed in Mt Lofty populations compared with

the Murray Valley populations, suggesting slight

differentiation of the two. In this feature, the Mt

Lofty populations are more similar to the east

coast New South Wales populations in which

throat patterning is also weakly developed or

absent.

Eulamprus tympanum tympanum (Lonnberg &

Andersson, 1913)

(Figs 1, 5)

Lygosoma tympanum Lonnberg & Andersson,

1913: 9. Holotype: NHRM 3094 ‘neighbourhood

of Melbourne’, Victoria.

Sphenomorphus quoyit

Loveridge, 1934: 350.

Sphenomorphus tympanus Mittleman, 1952: 31.

tympanum (part)

Sphenomorphus tympanum (part) Worrell, 1963:

333},

192

Sphenomorphus tympanum Cool Temperate Form,

Rawlinson, 1969: 119.

Eulamprus tympanum Wells & Wellington, 1984:

94.

Eulamprus herseyi Wells & Wellington, 1985:

29. Holotype: AMS R111949 (formerly AM Field

Series 16791), Dora Dora National Park Proposal

Area near Albury, New South Wales.

Type Specimens

The holotype of Lygosoma tympanum, NHRM

3094, is in good condition and clearly identifiable

as belonging to the ‘Cool Temperate Form’ of

Rawlinson (1969). The anterior margin of the ear

opening is pale, the throat and chin are smudged

with grey and there is no trace of a pale post-

supraciliary streak. Midbody scales are in 37

rows. The specimen is immature, with a SVL of

46 mm. The appearance of the specimen is

consistent with the collection data—‘said to have

been collected in the neighbourhood of

Melbourne, July 1911’.

Wells & Wellington (1985) failed to

differentiate Eulamprus herseyi from its

congeners. Their purported diagnosis merely listed

a number of scalation and meristic parameters

none of which, either individually or collectively,

differentiates E. herseyi from E. heatwolei, E.

tympanum or even E. quoyii. The only exception

is the supposed five supraoculars. The holotype

does have five supraoculars on the right side due

to an abnormally divided first supraocular but the

normal count of four is present on the left side and

the specimen is otherwise a typical E. tympanum.

Wells & Wellington define their genus Eulamprus

(restricted to the quoyii complex) as having four

supraoculars, although two of the three species

they described, E. heatwolei and E. herseyi, were

said to have five. Possibly the small scale at the

posterior end of the supraoculars (see above, p.

187) is responsible for this inconsistency.

Diagnosis

A water skink lacking any trace of longitudinal

dorsal striped pattern, with a pale anterior margin

to the ear opening, usually 42 or fewer midbody

scale rows and without transversely oriented dark

dorsal markings.

Description

3644 (x 39.2, n = 116) longitudinal scale rows

at midbody. Paravertebral scales 68-89 (x 75.3, n

= 116), scarcely broader than adjacent dorsals.

Subdigital lamellae on fourth toe 18-29 ( 22.4, n

= 109), most with a median groove and those at

M. N. HUTCHINSON & P. A. RAWLINSON

the base of the toe divided.

Prefrontals separated (freq. 0.45) or in point to

moderately broad contact. Interparietal elongate,

approximately twice as long as wide, but usually

not separating parietals (frequency of separation

0.11). Each parietal bordered posteriorly by one to

three nuchal scales and laterally by the upper

secondary temporal. Supraciliaries 8-10 (> 3 >

rest. Supralabials 6-9 (x 7.1, mode 7, n = 40),

fourth, fifth or sixth subocular. Infralabials 6-9 (x

7.5, mode 7, n = 40), first and second in contact

with (single) postmental.

Premaxillary teeth usually 8 (n = 9), less often 9

(n = 4) or 7 (n= 1).

Dimensions (adults, n = 90). SVL 66-93 mm

(* 81.7). HW 10.0-14.6 mm. HL 14.7-19.9 mm.

HL/HW 1.27-1.59 (x 1.43). HW/SVL 0.137-

0.168 (x 0.151). HLL 26-36 mm. HLL/SVL

0.311-0.449 (x 0.379). Tail length/SVL (n = 26)

1.40-1.83, x 1.58.

Colour (in preservative) light to very dark

brown on the dorsal surface of the head, body, tail

and limbs, immaculate or with few to numerous

irregular black flecks. No suggestion of a pale

dorsolateral stripe. Tail with dark flecks better

developed laterally than dorsally; limbs overlain

by heavy black network. Upper lateral zone black,

the colour usually fading to brown on the temples.

Several uneven series of widely spaced pale dots

(each covering only a single scale) overlie the

black. A pale horizontal streak runs posteriorly

from the dorsal rim of the ear opening and is

continuous with the pale anterior edge of ear

opening. Lower lateral zone pale grey, lightly

dotted with pale yellow and dark grey, less often

with weak black barring. Chin and throat light to

dark grey in most populations, sometimes with

darker smudges. Otway Ranges specimens

sometimes with black throats. Remainder of

underside yellowish white, sometimes immaculate

but usually with black markings which may occur

as small dark flecks or as black pigment

concentrated along the edges of scale rows,

forming thin lines.

In life the general colour is similar in most

populations. Ventral colour of Otway Ranges

specimens usually bright yellow.

Distribution

The Great Dividing Range, continuous from the

New South Wales border to about Ballarat,

extending south from the Divide into southern

Gippsland. Disjunct populations occur around the

Pyrenees, Grampians and Otway Ranges and far

southwestern Victoria and southeastern South

WATER SKINKS 193

E. t. tympanum @

E. t. marniae

Intergrades

Eulamprus tympanum

FIGURE 5. Distribution of Eulamprus tympanum in Victoria and South Australia. The stars denote localities where

E. heatwolei and E. t. tympanum have been collected in syntopy.

194 M. N. HUTCHINSON & P. A. RAWLINSON

Australia (Fig. 5). Extralimitally the species

extends northward along the Great Dividing

Range as far as the Blue Mountains, west of

Sydney, New South Wales.

Ecology

Aspects of the ecology of E. t. tympanum have

been covered by Rawlinson (1969, 1971, 1974),

Spellerberg (1972b-d), Pengilley (1972), Tilley

(1986), Brown (1991) and Schwarzkopf (1992). It

is a diurnal and heliothermic skink, restricted to

the margins of water courses only in the lower

rainfall portions of its range (e.g. the northern and

western margins of Melbourne). Over much of

cool temperate southeastern Australia, E. t.

tympanum is a widespread forest-dwelling

species. In montane forests in eastern Victoria, the

Australian Capital Territory and southern NSW

this is one of the most commonly encountered

reptile species. Activity is generally centred

around rotting fallen logs and stumps which are

used as perches for thermoregulation and the

cavities of which are used for shelter (Mather

1978, Tilley 1986). Tilley’s study demonstrated

that the species is probably non-territorial, has low

juvenile survivorship but potentially long-lived

adults, females living for up to 13 years and males

to 11. Brown (1991) found that this species is a

generalised invertebrate carnivore, taking only a

small proportion of plant matter in its diet.

Females ovulate late October-November, with

litter size (oviducal eggs) ranging 2-6 (X 4.5,n=

16). Testes are enlarged between April and

August, and regress over spring to a minimum in

November—January. The species is viviparous, the

young being born during January and February.

The time of mating has not been recorded.

Rawlinson (1974) stated that E. tympanum mated

in autumn, with overwintering of sperm by

females. This latter conclusion was drawn directly

from the above mentioned observations of the

testicular cycle coupled with the belief that

testicular enlargement ought to be correlated with

male sexual activity. However, as Greer (1989)

noted, both E. quoyii (Veron 1969) and E.

heatwolei (Pengilley 1972) are known to mate in

spring even though, like E. tympanum, they have

a testicular maximum in late autumn—winter.

Observational data are needed to establish the

time of mating for E. tympanum.

Discussion

Eulamprus tympanum is most obviously

different from the other water skinks in having a

broader head relative to body size than the other

species. In body and limb proportions it is

intermediate between the gracile E. quoyii and E.

heatwolei and the dumpy E. kosciuskoi.

The nominate subspecies shows little

geographic variation even though several western

populations appear to be isolated from one

another. Local trends include larger scales and a

greater development of linear black ventral

markings in Otway Ranges specimens and longer

tails and smaller body scales in Grampians

specimens. A general trend is for rock-dwelling,

Streamside populations to have a greater

development of black dorsal flecking than log-

dwelling, forest populations.

Eulamprus tympanum marnieae subsp. nov.

(Figs 5-7)

Eulamprus tympanum ssp. nov. Cogger et al.

ICL MOT,

Types

HOLOTYPE: NMV D52921, adult male, 5.5

km E. of Dreeite, Victoria, 38°11'S, 143°34'E, P.

A. Rawlinson; P. Robertson and M. Hutchinson,

1 November, 1979.

PARATYPES: 30 specimens, all from Dreeite

area. D49377, D49385—92, D52912-20, D52922—

52926, D52955-56, D53977-80, D62035 (see

appendix for details of localities).

Diagnosis

A water skink distinguished from all other

members of the E. quoyii species complex by the

very small midbody scales (usually in 43 or more

rows, versus usually 42 or fewer), the black dorsal

markings arranged as short irregular transverse

bars, and bold ventral pattern of black longitudinal

bars on a bright yellow (in life) background.

Description

40-48 (x 44.8, n = 36) longitudinal scale rows

at midbody. Paravertebral scales 76-95 (* 84.4, n

= 36), no larger or only slightly broader than

adjacent dorsals. Subdigital lamellae on fourth toe

20-26 (X 22.9, n = 36), most with a median

groove and those at the base of the toe divided.

Prefrontals separated (freq. 0.47) or in point to

moderately broad contact. Interparietal elongate,

approximately twice as long as wide, and almost

or actually (0.19) separating parietals. Each

parietal bordered posteriorly by one to three nuchal

scales and laterally by the upper secondary

temporal. Supraciliaries 7—9 (X 8.0); first to third

forming decreasing series, next three smalles, last

two larger, usually penetrating dorsally each side

WATER SKINKS 195

of the fourth supraocular. Supralabials 6-8, fifth

or sixth subocular. Last supralabial sometimes

horizontally divided (n = 2) to give the count of

six. Infralabials 7—9, first and second in contact

with (single) postmental.

Premaxillary teeth 8 (n = 3).

Dimensions (adults, n = 27). SVL 72-97 mm.

HW 10.4-15.7 mm. HL 15.3—20.6 mm. HL/HW

1.32-1.50 (x 141). HW/SVL 0.132-0.171 (x

0.141). HLL 27-36 mm. HLL/SVL 0.323-0.420

(x 0.368). Tail length/SVL (n = 6) 1.52-1.72

163),

FIGURE 6. Head shields of the holotype of Eulamprus tympanum marnieae n. subsp. (NMV D52921).

Colour (in preservative) light to very dark

brown on the dorsal surface of the head, body, tail

and limbs, overlain by black markings as follows:

head shields with irregularly scattered flecks; back

with numerous irregular patches, generally

laterally expanded to form transverse bars, some

contacting the black upper lateral zone; tail with

closely-spaced transverse wavy bars better

developed laterally than dorsally; limbs overlain

by heavy black network. Upper lateral zone black,

the colour extending anteriorly to the eyes. One or

two uneven series of widely spaced pale dots

196 M. N. HUTCHINSON & P. A. RAWLINSON

FIGURE 7. Live paratype female of Eulamprus tympanum marnieae (NMV D53980) (photo: M. Hutchinson).

(each covering two or three scales) overlie the

black, including a whitish temporal spot and a

horizontal streak running posteriorly from the

dorsal rim of the ear opening and continuous with

its pale anterior edge. Dorsal margin of upper

lateral black colouring with jagged projections,

with a pale spot in the notches between the

projections. Lower lateral zone greyish yellow

with irregular black barring. Anteriorly to the

axilla, the lower lateral zone resolves itself into a

midlateral pale stripe. Chin shields white spotted

with black. Throat grey-white with large blackish

patches laterally which form the ventral margin of

the midlateral pale stripe. Underside yellowish

white with black pigment forming interrupted

longitudinal black patches.

In life the general colour is similar but suffused

with yellowish. The dorsal ground colour is

frequently bright brassy and the underside of the

belly, limbs and tail is bright yellow.

Features of Holotype

Midbody scale rows 47. Paravertebral scales 78.

Subdigital lamellae under fourth toe 23.

Supralabials 7/8. Infralabials 7/8. Supraciliaries

7/7. Anomalies of scalation include a super-

numerary anterior chin shield on the left side and

three loreals also on the left side. SVL 80 mm.

Tail length 133 mm (1.66 SVL). HLL 33 mm

(0.413 SVL). HW 13.0 mm (0.163 SVL). HL 18.0

mm. HL/HW 1.38. Testes regressing.

Etymology

Named for Marnie Lincoln Rawlinson.

Distribution

Rocky outcrops and drystone walls on the

northern margins of the basaltic ‘stony rises’ east

and north of Lake Corangamite, southwestern

Victoria (Fig. 5). An outlying population occurs

beside Lake Bolac to the northwest.

Ecology

A diurnal, evidently heliothermic skink,

normally observed perched on a rockpile or fence.

Unlike other water skinks this is an extremely shy

lizard, fleeing to cover even when a human

observer is tens of metres distant and seldom

coming into open view while being watched. Like

other water skinks, E. t. marnieae will dive into

water and swim submerged to escape pursuit, but

most individuals are observed away from standing

water and take refuge in deep gaps in rockpiles.

Favoured localities combine remnant arboreal

vegetation (notably Hymenanthera, Violaceae),

deeply fissured basaltic rock piles and permanent

or ephemeral swamps. Its habitat in summer can

appear very arid compared with the areas

WATER SKINKS 197

inhabited by its relatives, but in winter the terrain

is extensively flooded, with almost all depressions

and low-lying areas holding water, and it seems

likely that the deep rock piles favoured by this

subspecies provide cool and humid refuges even

during the dry summer months.

Specimens have been collected between the end

of winter and mid-summer and in late autumn.

Females ovulate late October—early November,

with recorded litter size (oviducal eggs) ranging

2-6 (x 4.3, n = 21). Males with enlarged testes

have been collected in April and September, with

testes regressing in October-November and

completely regressed in January. Viviparity is the

mode of reproduction in all other water skinks for

which mode is known (not confirmed for E.

leuraensis) so it is highly probable that E. t.

marnieae is also viviparous. Birth would be

expected to occur in January, reinforced by the

collection date of the smallest individual in the

series, a juvenile (D53977, SVL 39 mm) collected

on 24—25 January.

Discussion

The subspecies is known from _ three

populations, each slightly differentiated (see Table

1). Aside from the type population described

above, series have been collected from 8 km S of

Lismore, Vic., (NMV D36075-084) and from

Lake Bolac (D52600-01, D52901-09). The

Lismore series is notable for its higher

paravertebral counts (up to 98), high frequency of

separation of the parietals by the interparietal (8

out of 10) and for a relatively high incidence of

division of the last supralabial (4 out of 10). The

Lake Bolac specimens are darker, with less yellow

pigmentation in life, have the highest midbody

scale counts (up to 53), high frequency of

separated parietals (7 out of 11) and a common

scalation abnormality, fusion of the last two

supralabials (5 out of 11).

This taxon is readily distinguished from other

water skinks by its small scales and bold dorsal

pattern, and the initial conclusion on discovering

this form was that it represented a new species.

However, subsequent collections suggest that it

intergrades with typical E. tympanum. North and

west, relatively arid conditions provide a barrier

between the two; no water skinks have been found

in the arc running from between Lake Bolac and

the Pyrenees east to about Rokewood and

southeast to about Winchelsea. However, to the

south and southwest, typical E. tympanum is more

continuously distributed and specimens from

Cororooke (south of Dreeite) and from north of

Camperdown (Kariah and Lake Colongulac) are

intermediate in colour pattern and scalation

between typical tympanum and marnieae (Table

Such populations have become taxonomically

problematic with the recent acceptance of the

inadequacy of the old subspecies concept (Collins

1992, Frost et al. 1992). The overuse of

‘subspecies’ to arbitrarily name geographically

isolated but undifferentiated populations or to

artificially partition continuous or clinal variation

has devalued the term, but there remain cases such

as the present in which a relatively consistent

phenotype confined to a specified geographic area

appears to be genetically continuous with adjacent

TABLE |. Comparison of Victorian populations of Eulamprus tympanum. Colour pattern characters scored are: 1,

black transverse dorsal bars present; 2, black longitudinal ventral bars present; 3, lateral margins of throat black; 4,

pale lateral flecks cover more than a single scale.

Population MBSR PVS Colour pattern characters

(n) (x) (x) 1 DD 3 4

E. t. marnieae

Dreeite area (36) 44.8 84.4 + + + +

8 km S of Lismore (10) 48.4 92.9 + + + +

Lake Bolac (11) Sill 85.0 + + + +

Intermediate

Camperdown area (5) 41.0 79.6 - + + -

Cororooke (3) 40.7 VIB +/— + + -

E. t. tympanum

Pirron Yallock (2) 39.0 80.0 /+

Otway Ranges (30) SHE) WS. /+ /+

Grampians (30) 40.7 77.0 /+

Pyrenees—Mt Macedon (26) 395) 76.8

Eastern Victoria (30) 39:3 74.6

198 M. N. HUTCHINSON & P. A. RAWLINSON

but phenotypically and geographically discrete

populations (Frost & Hillis 1990). In the case of

the water skinks reticulate evolution probably

occurs at least on the southern margins of the

distribution of marnieae. The possibility exists of

introgression of genes beyond the limits of any

putative hybrid zone, suggested by an increase in

midbody scale counts south to north moving away

from the contact with tympanum. To combine the

two simply as a binomial E. tympanum would

bury this most distinctive population, while to

recognise marnieae as a full species would be to

imply an evolutionary independence which is

denied by the circumstantial evidence available.

Further specimens and biochemical genetic data

would illuminate the degree of gene flow currently

occurring and would reveal the degree to which

the intermediate populations are acting as a bridge

or barrier to gene flow; should the latter be

established, elevation of marnieae to full species

status would logically follow. The fact is,

however, that such data will be difficult to obtain

due to the general difficulty of locating any water

skinks in the highly modified intergrade areas.

Eulamprus t. marnieae inhabits a geo-

graphically peculiar Victorian landform,

technically part of the Newer Volcanics, a region

of extensive late Tertiary sheet basalt lava flows,

and colloquially known as the ‘stony rises’. The

‘rises’ are basalt ridges and boulder heaps left by

the collapse of lava tunnels. The area supports

other distinctive herpetofauna, including a small-

scaled, speckled form of Pseudemoia

entrecasteauxii (Hutchinson & Donnellan 1992)

and a green morph of the normally brown treefrog,

Litoria ewingii (Hero et al. 1991).

Within its limited geographic range this

subspecies occurs only patchily, with only one

large colony known. Most of the area in which the

species probably once occurred has been

extensively modified, with total clearing of

vegetation for grazing and continuing demolition

of drystone walls and removal of boulders for

‘mossy rock’ landscape gardening. These

processes show no signs of ceasing and the

subspecies must be regarded as threatened. The

Lismore population was probably wiped out

during the rock clearing activities that produced

the specimens.

Eulamprus kosciuskoi (Kinghorn, 1932)

(Figs 4, 8)

Lygosoma (Hinulia) quoyii kosciuskoi Kinghorn,

1932: 359. Holotype: AMS R4654, Mt Kosciusko,

New South Wales.

Sphenomorphus quoyii

Loveridge, 1934: 350.

Sphenomorphus kosciuskoi Mittleman, 1952: 26.

Sphenomorphus tympanum (part) Worrell, 1963:

53,

Eulamprus kosciuskoi Wells & Wellington, 1984:

3},

tympanum (part)

Types

The specimens forming the type series, a

holotype, AMS R4654, and four paratypes (AMS

R558-9, R4832 and R5061), are clearly

identifiable as belonging to the southern

population. They have a light colour pattern, with

weakly marked head and clear light ground colour

between the black vertebral and dorsolateral

stripes. The dark upper lateral zone does not

extend to the ventrolateral area. Midbody scales

are in 31-34 rows?. The holotype has 34 midbody

scale rows, 19 subdigital lamellae and a SVL of

76 mm.

Diagnosis

A small water skink (largest adult 86 mm

snout-vent) distinguished from all but E.

leuraensis by the presence of a black vertebral

stripe. Said to differ from E. leuraensis by its

paler dorsal colouring (including weakly marked

head), broader paravertebral stripes and pale-

spotted black lateral pattern stopping at the

midlateral level rather than extending to the

ventrolateral angle of the trunk (Shea & Peterson,

1985).

Description

30-34 (x 32.3, n = 37) longitudinal scale rows

at midbody. Paravertebral scales 58-68 (x 63.6, n

= 32), slightly broader than adjacent dorsals.

Subdigital lamellae on fourth toe 18-23 (x 20.5,

n = 32), most with a median groove and those at

the base of the toe divided.

Prefrontals moderately separated to broadly

contacting (frequency of separated prefrontals

0.38); an azygous ‘interprefrontal’ sometimes

present (frequency 0.16). Interparietal elongate,

approximately twice as long as wide, usually

(frequency 0.71) separating parietals. Each

parietal bordered posteriorly by two moderately

2 Shea & Peterson (1985) gave the range for this series as 32—

35, but obtained the same average across all five, 32.2, as we

did.

WATER SKINKS 199

1icm

FIGURE 8. Head shields of Eulamprus kosciuskoi

(NMV D42202).

expanded nuchal scales and laterally by a large

upper secondary temporal. Supraciliaries 7—9; size

1 >2= last > 3 > rest. Supralabials 6-7 (mode 7),

fourth or fifth subocular. Infralabials 7-8 (mode

7), first and second in contact with (single)

postmental.

Premaxillary teeth 8 (n = 2) or 9 (n= 2).

Dimensions (of adults, all females, n = 13).

SVL 66-74 mm (X 70.6). HW 9.2-10.9 mm. HL

13.8-15.4 mm. HL/HW 1.40-159 (x 1.49). HW/

SVL 0.129-0.152 (x 0.140). HLL/SVL 0.292-

0.361 (x 0.326). Tail length/SVL (for individuals

>50 mm SVL, n = 8) 1.07-1.25 (« 1.17).

Colour (in preservative) light brown on the

dorsal surface of the head, body, tail and limbs.

Back with a black vertebral stripe running from

the nape to the base of the tail. A narrow yellow

dorsolateral stripe runs from the supraciliary

region to the base of the tail, becoming less

brightly coloured. This stripe edged medially by a

wider black stripe. Black upper lateral zone,

commencing at the ear and breaking up on the

tail, with three to four series of yellowish dashes

or dots, tending to align longitudinally on adjacent

scale pairs. A horizontal streak runs posteriorly

from the top of the ear opening and is continuous

with the pale anterior margin of the ear. Tail with

regularly spaced blackish lateral blotches. Limbs

brown mottled with black. Lower lateral zone

greyish with black spots, the pattern continuing

across the belly. Infralabials, chin, throat and

remainder of underside greyish white with

scattered black spots.

In life the general colour is similar but overlain

by a yellow-green opalescent gloss.

Distribution

Davies Plains-Mt Cobberras area and the

Bogong High Plains, Victoria (Fig. 4). The

southern, typical populations of this species are

confined to the Snowy Mountains of New South

Wales and the adjacent alpine areas of Victoria. A

second morphologically distinct group of

populations occurs in the Barrington Tops region

and on the New England Plateau (Shea &

Peterson, 1985).

Ecology

The Snowy Mountains—Victorian High Plains

populations of this species have a restricted

habitat, being found only in subalpine to alpine

bogs and sluggish creek margins, the habitats

being characterised by being perennially wet, cool

and densely vegetated at ground level. Coventry &

Robertson (1980) and Mansergh (1982) briefly

describe the habitat of this species at Davies

Plains and Mt Cope, respectively.

We have no explanation for the absence of

males in this sample, and their under-

representation in the samples of E. heatwolei and

E. tympanum. Certainly those females that are

pregnant are more catchable than non-pregnant

females or males, but excluding pregnant females

still leaves a deficiency of males. Further study of

wild populations could determine whether there is

a sex ratio bias against males, or whether

behavioural attributes of the sexes make males

harder to collect.

Most specimens have been collected during late

January, at which time about half of the females

(5) have full term young, while the rest (6) contain

no young or enlarged ova. A single adult female

collected in December contains advanced

embryos, while a single adult female collected in

February is non-breeding. At present it is not

possible to say whether the females without young

in January had just given birth or had not bred

that year, but this would be worth exploring in

view of the fact that females of the Tasmanian

alpine skinks (Niveoscincus) only breed every

second year (Greer, 1982, Hutchinson et al. 1989).

Litter size is 2-4 (x 3.2).

200

Discussion

Shea & Peterson (1985) noted that the topotypic

(Mt Kosciusko area) population of E. kosciuskoi

is more similar to E. leuraensis than is the New

England population, suggesting some

intermediacy, but maintained the specific

distinctness of the latter taxon. In fact, where one

draws the line between populations depends upon

the character chosen; if colour pattern, then most

Snowy Mountain—Victorian and New England E.

kosciuskoi are more similar to each other than

either is to E. leuraensis, while if limb proportions

and head and body scalation are considered, then

the new England E. kosciuskoi stand apart.

Moreover, the more melanised individuals in the

Victorian sample approach the description of E.

leuraensis in that the head is heavily black-

flecked, the laterodorsal black stripes are broader

and leave little dorsal ground colour showing and

the midlateral pattern extends ventrally to the

lower lateral area. It appears therefore that no

absolute distinctions separate any populations

formerly referred to E. kosciuskoi. It is evident

from comparison with other water skink

populations that degree of dorsal melanisation can

be labile in water skinks and that isolated

populations can undergo shifts in scale count

frequencies. The situation of E. tympanum

discussed above also shows that much greater

qualitative differences in colour pattern and scale

size are not necessarily indicative of independently

evolving entities (species).

Allopatric populations are a problematic group,

even if one discards the increasingly unpopular

biological species concept (BSC, e.g. Frost &

Hillis 1990). Recent debate on the North

American fauna (Collins 1991, 1992, Montanucci

1992, Van Devender et al. 1992, Frost et al.

1992) highlights the differing views that can exist

concerning populations which until recently most

authors would have identified as subspecies, more

for convenience than as a positive expression of

the degree of historical independence. Further

consideration of the status of E. leuraensis is

beyond the scope of this study, but the appropriate

taxonomic treatment for the alpine water skink

populations could stand further analysis.

Eulamprus heatwolei Wells & Wellington, 1984

(Figs 9, 10)

Sphenomorphus tympanum Warm Temperate

Form, Rawlinson, 1969: 119.

M. N. HUTCHINSON & P. A. RAWLINSON

Eulamprus heatwolei Wells & Wellington, 1984:

93. Holotype: AMS R116967 (formerly AM Field

Series 27987), Macquarie Rivulet, just east of

Robertson, New South Wales, R. W. Wells, 20

October, 1982.

Sphenomorphus heatwolei Shea & Peterson,

1985: p. 144.

Type Specimen

The holotype of Eulamprus heatwolei, AMS

R116967, is in moderate condition, with the tail

almost broken and is clearly identifiable as

belonging the ‘Warm Temperate Form’ of

Rawlinson (1969). Wells & Wellington (1984)

purported to diagnose the species by listing the

attributes of the holotype, but only aspects of the

colour pattern description are unique to heatwolei.

Our observations on the holotype disagree with

those of Wells and Wellington (in square

brackets) in several significant respects. Midbody

scale rows 40 [38]. Tail regenerated, so subcaudal

count [73] irrelevant. Supraoculars 4 [5]; first

supraocular on left fragmented, with two

abnormal small scales contiguous with the

supraciliary row. Postnasal scales absent [said to

be present]. One [2] preocular, if the lower of two

antorbital scales is regarded, as here, as the first

presubocular. Supraciliaries 10/9 [6]. Adpressed

limbs strongly overlap, the fourth toe of the hind

foot reaching about the level of the elbow

[adpressed limbs just fail to meet]. Other

important points not mentioned in the type

description are throat colouring, the chin shields

being heavily edged with black and the throat grey

with four irregular blackish longitudinal bars, and

size, the snout—vent length being 71 mm.

Diagnosis

A water skink lacking longitudinal dorsal

stripes, most similar to E. tympanum, but differing

in its longer appendages, pale post-supraciliary

streak, black anterior edge of the tympanic

opening and immaculate venter (bright yellow in

life).

Description

36-44 (« 39.9, n = 73) longitudinal scale rows

at midbody. Paravertebral scales 69-89 (x 77.5, n

= 73), scarcely or not broader than adjacent

dorsals. Subdigital lamellae on fourth toe 23-29

(X 25.1, n = 73), most with a median groove and

those at the base of the toe divided.

Prefrontals in point to broad contact (freq. =

0.32). Interparietal elongate, approximately twice

as long as wide, but seldom separating parietals

WATER SKINKS 201

‘

FIGURE 9. Head shields of Eulamprus heatwolei (SAM

R38610).

(3 out of 73 scored). Each parietal bordered

posteriorly by one to three nuchal scales and

laterally by the upper secondary temporal.

Supraciliaries 7-11 (x 9.1, mode 9); size 1 > 2 =

last > 3 > rest. Supralabials 7-9 (mode 7), fifth or

sixth subocular. Infralabials 6-9 (mode 8), first

and second in contact with (single) postmental.

Premaxillary teeth usually 9 (n = 8), less often 8

@=3),

Dimensions (of adults, n = 47). SVL 74-101 («x

84.3). HW 10.1-14.0 mm. HL 16.3-20.7 mm.

HL/HW 1.42-168 (x 1.51). HW/SVL 0.126—

0.162 (* 0.143). HLL 30-36 mm. HLL/SVL

0.327-0.439 (x 0.379). Tail length/SVL (n = 13)

1.58-1.92 (x 1.76).

Colour (in preservative) light to very dark

brown on the dorsal surface of the head, body, tail

and limbs. Head shields with irregular black

flecks. A paler brown streak, suggesting the start

of a pale dorsolateral stripe, runs posteriorly from

the supraciliary region to the neck. Back usually

with numerous irregular black flecks; immaculate

in very few specimens (those from the Murray

River, both Victorian and South Australian). Tail

with black, closely-spaced wavy bars laterally;

limbs overlain by wavy black bars. Upper lateral

zone black, the colour extending over the

temporals and eye to the loreals. Several uneven

series of pale dots (each covering only a single

scale) overly the black. A pale horizontal streak

runs posteriorly from the dorsal rim of the ear

opening; anterior edge of ear opening black.

Lower lateral zone greyish with black scales

forming irregular vertical or backward-sloping

bars or a black reticulum. Chin and throat white,

most populations with black edges on chin shields

and elongate black blotches on the throat.

Remainder of underside yellowish white,

immaculate.

In life the general colour is similar but suffused

with yellowish. The dorsal ground colour is

frequently brassy and the underside of the belly,

limbs and tail is bright yellow...

Distribution

Eastern Victoria, west to about the Goulburn

River (Fig. 10). Absent from higher elevations

along the Great Dividing Range. An isolated

record from Great Western (Victoria) and a

disjunct group of populations on the lower

Fleurieu Peninsula, South Australia, from Deep

Creek to the northern shore of Lake Alexandrina.

Extralimital in eastern New South Wales north to

the New England plateau.

Ecology

This species is very similar in habits to E.

quoyii, being mostly restricted to creek margins.

Most populations of this species favour rocky

substrates, but the species does occur along

muddy river banks, notably the Murray in north—

central Victoria and at its mouth in South

Australia. Its reproductive characteristics appear

similar to those of E. quoyii, with similar ovarian

and testicular cycles. Litter size among the

specimens examined ranged 2-4 (x 3.3, n = 9).

The distributions of E. heatwolei and E. t.

tympanum are complementary to a remarkable

degree; although the distributions abut and

interdigitate throughout eastern Victoria there are

few places (a total of ten) where specimens have

been collected in syntopy (indicated by stars in

Figs. 5 and 10). Syntopy generally occurs along

stream valleys where E. heatwolei can extend

along the warm valley floor to contact E.

tympanum populations inhabiting the cooler,

elevated valley slopes. In East Gippsland, the

aspect of a section of stream channel can

determine the species present, with north facing

slopes occupied by E. heatwolei while south-

facing slopes are occupied by E. t. tympanum.

Meandering watercourses may show an

alternation of species (A. J. Coventry, pers.

comm.) depending on the degree of shading. No

202 M. N. HUTCHINSON & P. A. RAWLINSON

Eulamprus heatwolei

FIGURE 10. Distribution of Eulamprus heatwolei in Victoria and South Australia. The stars denote localities where

E. heatwolei and E. t. tympanum have been collected in syntopy.

WATER SKINKS

studies have yet been directed to determining the

degree to which competition or the documented

physiological differences (Spellerberg, 1972b,c)

determine range limits in these two species.

Discussion

The type locality of E. heatwolei, Robertson,

New South Wales, is adjacent to this region so

that as yet the species is not certainly recorded

from other parts of Australia. Cogger (1992)

mapped the species as far south as the New South

Wales-Victoria border. For many years workers

have used Rawlinson’s informal taxonomy to

distinguish two species level taxa, the ‘Cool

Temperate’ and ‘Warm Temperate’ forms of

Eulamprus (as Sphenomorphus) tympanum (e.g.

Spellerberg 1972b-—d, Rawlinson 1974, Jenkins &

Bartell 1980). The forms have been distinguished

mainly on ventral colouring, as well as dorsal

head colour pattern and proportions. Examination

of the holotype of E. heatwolei and other New

South Wales specimens shows that they do not

differ from the Victorian ‘Warm Temperate

Form’. Accordingly, the concept of E. heatwolei

is expanded to include these populations.

Shea & Peterson (1985) listed variation within

this species based on Blue Mountains area

specimens only. Their data are similar to ours,

with slightly higher midbody scale counts (x

40.8) and maximum snout—vent length a little less

(92 mm).

South Australian populations of this species are

geographically remote from the main eastern

Australian populations, and are restricted to only

a few known sites. The terminus of the Murray

River on the northern margin of Lake Alexandrina

supports a colony living immediately adjacent to

the water on eroding banks next to cleared grazing

land. On the lower Fleurieu Peninsula E.

heatwolei is confined to a few perennial rocky

streams which retain some streamside vegetation.

In spite of their isolation, these populations differ

little from the continuous populations of eastern

Victoria and New South Wales. Body scales are

small, with midbody scale counts restricted to the

upper half of the total range of variation seen in

the species as a whole (midbody scale rows 40—

44, x 41.5 in South Australia), and higher

paravertebral counts (74-89, X 80.2, versus 69—

82, X 76.4 in Victoria). The South Australian

colonies appear vulnerable to habitat changes. At

Lake Alexandrina changes in Murray River flow,

either decreases leading to drying out or increases

causing erosion of the banks, may destroy local

populations. In the rocky streams of the Fleurieu

203

Peninsula, modification of the banks, with either

clearing of the vegetation or silt build-up reducing

exposed rocks, is likely to be deleterious.

A Key To Tue Eutamprus OF VictorIA AND SOUTH

AUSTRALIA

Pale yellow dorsolateral line runs from

behind the eye to at least the posterior one-

thindto fathes trun kerseeerees eee cater

— No pale yellow dorsolateral line

A black vertebral stripe......... E. kosciuskoi

— No black vertebral stripe.............. E. quoyii

Dark dorsal markings arranged as irregular

transverse bars; usually more than 42

midbody scale rows ............ E. t. marnieae

— Dark dorsal markings (if present) in the

form of small black flecks; usually fewer

than 43 midbody scale rows ...............06 4

Anterior margin of ear opening black; belly

in life immaculate bright yellow, most

intense under chest and groin; throat white

with black patches ................. E. heatwolei

— Anterior margin of ear opening cream;

belly in life pale yellow to greenish yellow

(bright yellow striped with black in some

Otway Ranges specimens) with or without

black flecks; throat light to dark grey, with

or without darker grey smudges.................

Bess ete neato setees E. t. tympanum

ACKNOWLEDGMENTS

The Curators of the European collections

housing type material showed Peter Rawlinson

great hospitality during his tour of collections in

1973, for which he was very grateful. In Australia,

R. Sadlier (AMS) and A. J. Coventry (NMV)

loaned specimens in their care. MNH particularly

thanks Marnie Rawlinson for freely allowing the

loan of detailed notes Peter compiled on type

material and his unpublished notes on the

taxonomy of this lizard group. A. J. Coventry

provided encouragement for the project and many

useful discussions concerning water skinks and

Peter Rawlinson’s thoughts on the group. I thank

Adrienne Edwards for her help in compiling some

of the scale count and morphological data and

Jennifer Thurmer (line drawings) and Trevor

Peters (photography) for preparation of the

illustrations.

204 M. N. HUTCHINSON & P. A. RAWLINSON

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APPENDIX

Specimens examined.

Eulamprus heatwolei.

New South Wales: AMS R116967, Macquarie Rivulet,

just E of Robertson (Holotype).

Victoria: MV D552, Great Western, 37°09'S 142°52'E;

D2286, Wangaratta, 36°21'S 146°19'E; D34392, 5.6 km

N.E. of Wodonga, 36°10'S 146°51'E; D34488, Nug Nug,

Ovens River, 1.6 km. W. of Myrtleford, 36°39'S

146°43'E; D 34541-48, Poddys Creek, 24.1 km. W. of

Cann River, 37°36'S 148°54'E; D34549, 14.5 km N. of

Nariel, 36°18'S 147°50'E; D 34550, Still Creek, 30.6 km

S.E. of Eildon, 37°25'S 146°09'E; D34551, 12.9 km S. of

Sardine Creek, 37°31'S 148°40'E; D34552, 16 km E. of

Orbost, 37°41'S 148°35'E; D34553, 24.1 km N. of

Orbost, 37°34'S 148°34'E; D34554, Snowy River,

Tulloch Ard Gorge, 37°16'S 148°19'E; D34560-62,

Goulburn River, 16 km S.E. of Seymour, 37°08'S

145°16'E; D36839, Lake Tali Karng, 37°33'S 146°47'E;

D36841, 2nd last crossing of Wellington River, Lake Tali

Karng Track; D36842, 9.7 km N. of Culloden on Dargo

River, 37°41'S 147°08'E; D36843, Wangareta

[=Wangaratta], 36°22'S 146°19'E; D36844—45, 14.5 km

N. of Stockdale, 37°40'S 147° 11'E; D36846—48, 16.1 km

N.N.W. of Stockdale, 37°39'S 147°06'E; D36849-53, 5.6

km E. of Wodonga, 36°09'S 146°52'E; D36854-SS, 19.3

km S.W. of Walwa, 36°05'S 147°34'E; D36856, 0.4 km.

E. of Genoa, 37°28'S 149°36'E; D42564—86, Sheepwash

Lagoon, 37°10'S 145°32'E; D56624, Seven Creeks,

Goorum Falls, 36°54'S 145°35'E; D57154, Two Mile

Creek, 13.4 km N.N.W. of Picolo, 35°59'S 144°S6'E;

D57164, Ovens River, Naughtons Bend, 14.3 km N.W.

of Peechelba, 36°06'S 146°08'E.

South Australia: MV D34467-68, 3.2 km. E. of

Myponga; D34487, D36880, Hindmarsh Falls, 35°27'S

138°35'E; D34555-56, D34559, Angas River,

Strathalbyn, 35°16'S 138°54'E; D34558, Currency

Creek, 35°27'S 138° 46'E. SAMA R2878, Tapanappa

Rocks, 35°38'S 138° 15'E; R3018, Deep Creek, 35°36'S

138°15'E; R13436A-B, Withers Creek, tributary of Deep

Creek, 35°38'S 138°15'; R17074, 1 km S of Mount

Compass, 35°22'S 138°38'E; R18572-73, Tower of

Babel, Inman Valley, 35°28'S 138°33'E; R24077, Lake

Alexandrina [north shore], 35°24'S 139°19"E; R38557,

Hindmarsh Falls, 35°27'S 138°35'E; R38610-12, North

shore of Lake Alexandrina at exit of Murray River,

Nalpa Stn, 35° 24'S 139° 20'E.

Eulamprus kosciuskoi.

New South Wales: AMS R558-59, R4654, R5061, Mt

Kosciusko (Types of Lygosoma (Hinulia) quoyii

kosciuskoi)

Victoria: MV D42060, Charlies Creek, Davies Plains

M. N. HUTCHINSON & P. A. RAWLINSON

Track, 36°17'S 147°59'E; D42075, Charlies Creek,

Davies Plains Track, 36°17'S 147°59'E; D42110,

Charlies Creek, Daveys Plain Track, 36°17'S 147°59'E;

D42201-02, Charlies Creek, Davies Plains Track,

36°17'S 147°59'E; D42203, King Plain, Davies Plains

Track, 36°39'S 148°04'E; D 47513-15, Davies Plains,

36°17'S 147°59'E; D47539-49, D47589-90, D47654—

59, Davies Plains, 36°17'S 147°59'E; D48557, The

Playgrounds, Mt. Cobberras, 36°52'S 148°09'E; D5511 il.

Mt. Cope area, 36°56'S 147°17'E,; D55630, Rocky

Plains, 36°56'S 148°10'E; D56465-66, 1 km E. of Mt.

Cope, 36°56'S 147°17'E; D59849, The Playgrounds, Mt.

Cobberras, 36°52'S 148°09'E.

Other: AMS R4832, no data (‘Tonga’, in error).

Eulamprus quoyii.

New South Wales: MNHN 2976, Port Macquarie;

7112-3, Neutral Bay, Port Jackson. (Types of Gongylus

(Lygosoma) quoyii).

Victoria: MV D1352, D1372, No other data; D13871,

Lindsay River, Berribee Station, 34°0'S T4ieOrks

D34326-31, Murray River, 6.4 km W. of Lock No.9;

D39078-79, Potterwalkalgee Creek, 34°08'S 141°23'E;

D39080-81, Murray River & Potwatagee

[=Potterwalkalgee] Creek junction, 34°08'S 141°23'E;

D39085, Potterwalkalgee Creek, 34°08'S 141°23'E;

D56882-83, Potterwalkalgee Creek, 5 km E. of Neds

Station corner, 34°08'S 141°23'E; D61906, Dedman

Creek, 13 km W. of Horseshoe Lagoon, Wallpolla Island,

34°08'S 141°42'E.

South Australia: MV D5297, Purnong, 34°51'S

139°38'E; D34282-83, Kingston [on Murray]; D34284—

89, Torrens River Gorge, 34°52'S 138°46'E; D 39070—

71, Cattambal, Torrens River Gorge. SAMA R2393A-B,

Sturt River, Eden, 35°01'S 138°36'E; R2874, Mylor,

35°03'S 138°46'E; R2889, Bridgewater, 35°00'S

138°46'E; R11177, Torrens Gorge, 34°53'S 138°44'E;

R13481, just N of Sinclair Flat, River Murray, 34°15'S

139°38'E; R13482, Waterfall Gully, 34°57'S 138°41'E;

R18519, Glenforslan Ranch, 34°15'S 139°39'E;

R22306-07, Sturt Creek, FlagstaffHill area, 35°02'S

138°32'E; R33009, Monoman Creek, Chowilla,

33°56'35"S 140°52'45"E; R33350, Clarendon, 35°07'S

138°38'E; R33782, Punkah Island, Chowilla Stn,

33°57'45"S 140°57'45"E; R37931, Scott’s Lagoon, 8.5

km S of Morgan, 34°07'S 139°40'E; R38017, Lake

Garnett, 20 km NE of Mannum, 34°53'S 139°31'E.

Other: MNHP 2977 ‘Nouvelle Hollande’ (Syntype of

Lygosoma quoyii). BMNH 1946.8.4.99, ‘Kangaroo

Island, S.A.’ (in error); 1946.8.15.34-35, ‘Queensland’,

1946.8.15.36, ‘Australia’; (Types of Hinulia

gastrosticta). NMW 16656:1—2, ‘Kangaroo Island’ (in

error) (Types of Hinulia gastrosticta).

Eulamprus tympanum tympanum.

New South Wales: AMS R111949, Dora Dora National

Park proposal area, 35°55'S 147°35'E (Holotype of

Eulamprus herseyi).

WATER SKINKS

Victoria: NHRM 3094, Neighbourhood of Melbourne

(Holotype of Lygosoma tympanum). MV D11881-

11883, Gellibrand River, 38°31'S 143°32'E; D12207—

12216, Gellibrand, 38°31'S 143°32'E; D 13642-13645,

Mt. Sabine 38°47'S 147°19'E; D13653-13654,

Gellibrand River 38°31'S 143°32'E; D14056-14065,

Dellys Dell 37°12'S 142°32'E; D15594, Near Halls Gap,

37°08'S 142°31'E; D17571, Gellibrand River 38°32'S

143°32'E; D33424, Junction of Mairs Track & Syphon

Road, 37°11'S 142°20'E; D33491, 3 km S. of Ben Nevis,

37°15'S 143°12'E; D33492-33493, 4.8 km N. of Mt.

Cole, 37°17'S 143°16'E; D33500, Grampian Ranges,

37°07'S 142°26'E; D33874, 14.5 km S. of Elmhurst,

37°18'S 143°15'E; D35846, Syphon Road, 37°10'S

142°20'E; D35847, Moora Hut, 34°14'S 142°26'E;

D35848-35849, 3.2 km W. of Pirron Yallock, 38°21'S

143°23'E; D3579, Back River Bridge, 8 km W. of

Bentleys Plains, 37°14'S 147°49'E; D35794, 12.9 km E.

of Moutys Hut; D35795, Ridge over Nuniong Plains,

37°08'S 147°57'E; D35798-800, 3.2 km W. of Cape

Horn, 38°44'S 143°34'E; D35802, Great Ocean Road,

3.2 km W. of Apollo Bay, 38°46'S 143°37'E; D 35806,

D35808-09, Dargo High Plains, 37°06'S 147°09'E;

D35830-33, D35835-39, Lake Mountain road, 6.8 km.

from Cumberland road, 37°31'S 145°52'E; D35852-—

35854, Matlock road, 54.7 km N. of Noojee, 37°24'S

146°0'E; D35855—35856, Matlock road, 37 km N. of

Noojee, 37°34'S 146°0'E; D35857, Big River Camp,

37°32'S 145°57'E; D35858-35859, 51.5 km N. of

Noojee, 37° 26'S 146° 0'E; D35860-35861, 24 km N. of

Noojee, on Matlock Road, 37°41'S 146°0'E; D35862, 66

km N. of Noojee on Frenchmans Spur road, = 29 km E.

of Big R. camp,37°32'S 145°57'E; D35863, Wartook,

37°02'S 142°21'E; D35864, 13.7 km N.W. of

Peterborough, 38°30'S 142°45'E; D35865, Lake

Wartook, 37°05'S 142°27'E; D35866, 4.8 km N. of

Tyers, 38°06'S 146°28'E; D35867, 4 km N. of Streiglitz,

37°S0'S 144°11'E; D35868, Badger Ck., Healesville,

37°34'S 145°35'E; D35869, Memorial Gardens, Mt.

Macedon, 37°22'S 144°35'E; D35870-35871, 8.8 km E.

of Marysville, 37°31'S 145° 50'E; D35872, 21 km S.E. of

Cumberland Junction 37°42'S 146°04'E; D35881, Fyans

Creek, 61 km. N. of Dunkeld, 37°05'S 142°34'E;

D35891, Wartook Reservoir, 37°05'S 142°27'E; D35985,

Chimney Pot Gap, 37°24'S 142°18'E; D 35985,

D35987-90, Chimney Pot Gap, 37°24'S 142°18'E;

D35993, Lake Wartook, 37°05'S 142°27'E; D36001,

Blanket Bay, 38°49'S 143°35'E; D36002, 8 km N. of

Cape Horn, 38°39'S 143°37'E; D36018-36028, 8 km N.

of Cape Horn, 38°39'S 143°37'E; D36063-36067, 3.2

km W. of Cape Horn, 38°44'S 143°34'E; D36092-

36093, Enfield, 12.8 km S. of Ballarat, 37°45'S

143°47'E; D36173, Lake Wartook, 37°05'S 142°27'E;

D36307-36309, Mt. Sabine, 38°38'S 143°44'E; D39399,

Mt. William, 37°13'S 144°48'E; D39401—39406, 4 km S.

of Ben Nevis fire tower, 37°16'S 143°12'E; D39407,

Lake Wartook, Grampian Road 37°05'S 142°27'E;

D47759, 4 km W.S.W. of Enfield, 37°45'S 143°45'E;

D47760, 3 km W. of Enfield, 37°45'S 143°45'E;

D47804-47807, Glenisla Shelter, Victoria Range,

37°09'S 142°15'E; D47808, Mt. William, 37°13'S

207

144°48'E; D47915, 0.5 km N. of Mt. Langi Ghiran,

37°17'S 143°08'E; D48690, Nowhere Creek, 37°07'S

143°17'E; D48691, Mt. Avoca, 37°06'S 143°21'E;

D48709-11, 5 km N.E. of Glenlofty, 37°05'S 143°15'E;

D50183-50185, 1 km S. of Mt. Lonarch, 37°16'S

143°21'E; D50257-58, 4 km W. of Mt. Buangor, 37°18'S

143°11'E; DS0716-19, Mt. Avoca, 37°06'S 143°21'E;

D50951-52, 1 km S.W. of Mt. Sabine, 38°38'S

143°43'E; D50966, Mt. Sabine 38°37'S 143°44'E;

D56657, Hopkins Falls, 38°20'S 142°37'E.

South Australia: SAMA R11263, Sect. 123, Hund. of

Young, County Grey, 37°43'S 140°45'E; R12400,

Southernmost point, Sect. 123, Hund. of Young, County

Grey, 37°43'S 140°46'E; R12982, NW of Port

Macdonnell, 37°59'S 140°33'E; R13076, SW of Mount

Gambier, 37°54'S 140°41'E; R14123, N of Mount

Gambier, 37°44'S 140°50'E; R14868, SE of Mount

Gambier, 37°55'S 140°57'E; R15163A-—B, Sect. 391,

Hund. of Caroline, 37°58'S 140°51'E; R16826A-C,

Rivoli Bay, 37°32'S 140°06'E; R17889, 5 km N of

Wandilo siding, 37°41'S 140°44'E; R19082-83, N of

Mount Gambier, 37°42'S 140°46'E; R23890, R23952,

Woolwash Creek near Port Macdonnell, 38°03'S

140°45'E; R23926, 10 km NW of Port Macdonnell,

38°00'23"S 140°36'11"E.

E. t. tympanum x marnieae intergrade specimens.

Victoria: D39412, 16 km N. of Camperdown, 38°01'S,

143°09'E; D52910-11, 1.5 km N. of Kariah, 38°10'S,

143°13'E; D56721, 11.5 km NNE of Camperdown,

38°08'S, 143°12'E; D56802, 4.5 km E (100°) of Bookar,

Lake Colongulac, 38°10'S, 143°10'E; D56803, 4.6 km

ESE (105’) of Cororooke, 38°10'S, 143°32'E; D56804,

D56878, 3.4 km SE (130°) E of Cororooke, 38°17'S,

143°33'E.

Eulamprus tympanum marnieae

Victoria: HOLOTYPE: NMV D52921, 5.5 km E. of

Dreeite, Victoria, 38°11'S, 143°34'E, P. A. Rawlinson &

M. Hutchinson, | November, 1979.

PARATYPES: 5.5 km E. of Dreeite: D49377, D49391—

92, P. A. Rawlinson & M. Hutchinson, 1 September

1977; D52912-20, D52922-529266, P.A. Rawlinson &

M. Hutchinson, 30 October and 1 November, 1979;

D62035, M. Hutchinson & S. Donnellan, September,

1986. 5 km E of Dreeite, 38°11'S, 143°34'E, D52955—S6,

G. Brown, 9 November 1979; D53977-80, G. Brown,

25 January, 1980. Dreeite, Taits Road, 38°11'S,

143°3 1'E: D49385-87, M. Hutchinson & G. Ingram, 26

August 1977; D49388-90, P. A. Rawlinson & M.

Hutchinson, | September 1977.

REFERRED SPECIMENS: D36075-84, 8 km S. of

Lismore, 38°01'S, 143°20'E, S. Hosgood, 16 April, 1963;

D52600-01, Lake Bolac, near caravan park, 37°44'S,

142°52'E, M. Hutchinson & G. Brown, 23 February

1978; D52901-09, Lake Bolac, near caravan park,

37°44'S, 142°52'E, P. A. Rawlinson & M. Hutchinson, 31

October 1979.