NEW SPECIES AND NEW RECORDS OF CLOACINA VON LINSTOW, 1898

(NEMATODA: STRONGYLOIDEA) PARASITIC INMACROPODID

MARSUPIALS FROM PAPUA NEW GUINEA

I. BEVERIDGE

BEVERIDGE, I. 2002. New species and new records of Cloacina von Linstow, 1898

(Nematoda: Strongyloidea) parasitic in macropodid marsupials from Papua New Guinea.

Records of the South Australian Museum 35(1): 1-32.

New species described from macropodid marsupials in Papua New Guinea are : Cloacina

cretheis sp. nov. from tree kangaroos, Dendrolagus inustus (type host), D. dorianus, D.

goodfellowi, D. matschiei and D. scottae; C. cunctabunda sp. nov. from D. mbaiso; C.

eurynome sp. nov. from D. dorianus and D. scottae; C. hecale sp. nov. from D. dorianus; C.

theope sp. nov. from D. matschiei and D. dorianus; C. erigone sp. nov., C. hyperaea sp. nov.,

C. nephele sp. nov., C. polymena sp. nov., C. praxithea sp. nov. and C. procris sp. nov. from

the scrub wallaby, Dorcopsis hageni, and C. oweni n. sp. and C. papuensis n. sp. from the

agile wallaby, Macropus agilis. New host records are: C. caballeroi Mawson, 1977, C. sterope

Beveridge & Speare, 1999 and C. syphax Beveridge & Speare, 1999 in Do. hageni. Additional

geographical records are given for C. cornuta in M. agilis. A key to the known species of

Cloacina in Papua New Guinea is provided.

I. Beveridge, Department of Veterinary Science, University of Melbourne, Parkville, Victoria

3052, Australia. Manuscript received 7 May 2001.

Species of the nematode genus Cloacina von

Linstow, 1898 occur exclusively in the stomachs

of macropodid marsupials. Currently, 103 species

are recognised as valid (Beveridge 1998, 1999;

Beveridge and Speare 1999) although substantial

numbers of species remain to be described

(Beveridge 1998). Most of the species described

to date are from Australian kangaroos and

wallabies, with relatively few records from Papua

New Guinea. This situation reflects the paucity of

knowledge of the parasite fauna of macropodids

from Papua New Guinea, which is based currently

on a small number of incidental collections.

Nevertheless, material currently available from

various kangaroos and wallabies consists of a

number of novel species of Cloacina. Thirteen

new species are described in the current paper as

well as new host and distribution records. The

opportunity is taken to provide a summary of

species of Cloacina known from Papua New

Guinea together with a key to facilitate their

identification. Finally, a preliminary comparison

is made between the Australian and Papua New

Guinean species and their host distributions.

Beveridge (1998) noted the occurrence of a

number of species of Cloacina in Papua New

Guinea, specifically C. australis (Yorke &

Maplestone, 1926) in the agile wallaby, Macropus

agilis (Gould, 1842); C. caballeroi Mawson, 1977

in the grey scrub wallaby, Dorcopsis luctuosa

(D’ Albertis, 1874) and the brown scrub wallaby,

D. muelleri (Schlegel, 1866); C cloelia Beveridge,

1998 in the pademelons Thylogale calabyi

Flannery, 1992 and T. stigmatica (Gould, 1860);

C. cornuta (Davey & Wood, 1938) in M. agilis;

C. cybele Beveridge, 1998 in T. stigmatica; and

C. dahli von Linstow, 1898 in Thylogale browni

Ramsay, 1877, T. calabyi and T. stigmatica.

Subsequently, based on an examination of the

parasites of four small scrub wallabies,

Dorcopsulus vanheurni (Thomas, 1922),

Beveridge & Speare (1999) described seven new

species, C. sancus, C. sciron, C. sappho, C. solon,

C. solymus, C. sterope and C. syphax. Beveridge

(1998) noted the presence of undescribed species

from the white-striped scrub wallaby, Dorcopsis

hageni Heller, 1897 in the collections of the South

Australian Museum, Adelaide, and from Do.

luctuosa in the collections of The Natural History

Museum, London. Flannery et al. (1996) also

reported species of the genus Cloacina in a variety

of tree kangaroos: Dendrolagus inustus Mueller,

1840; D. goodfellowi Thomas, 1908; D. dorianus

Ramsay, 1883; D. scottae Flannery & Seri, 1990;

and D. mbaiso Flannery, Boeadi & Szalay, 1995,

a group of kangaroos not previously known to

harbour Cloacina. These various undescribed

collections form the basis of the current report.

2 I. BEVERIDGE

MATERIALS AND METHODS

Nematodes were examined from collections

held in the Australian Helminthological Collection

(AHC), South Australian Museum (SAM),

Adelaide, the Natural History Museum (BMNH),

London and the United States National Parasite

Collection (USNPC), Beltsville, Maryland.

Nematodes were washed in water and cleared in

lactophenol. Drawings were made using a drawing

tube attached to an Olympus BH2 microscope.

Drawings of apical views of the heads of

nematodes are oriented with the dorsal aspect

uppermost; drawings of the bursa have the ventral

surface uppermost. All drawings are of paratype

specimens. Measurements were made with an

ocular micrometer. All measurements are in

millimetres and are presented as the range

followed by the mean in parentheses.

Morphological terminology follows Beveridge

(1998). Because of the relative uniformity of the

genus Cloacina, the descriptions presented are

abbreviated and follow the format used by

Beveridge (1998), in which a full description of

the genus was followed by individual species

descriptions concentrating on features of

diagnostic significance. A detailed description of

the genus is available in Beveridge (1998).

Features which are relatively invariable, such as

the disposition of ventral and lateral bursal rays,

are not included in descriptions but are,

nonetheless, illustrated.

Types of the new species have been deposited

in SAM, BMNH and USNPC. Host nomenclature

follows Flannery (1995) and Flannery et al.

(1995).

Following previous practice (Beveridge 1998),

the names of new species are based on those of

classical deities since Cloacina was the Roman

goddess of the toilet. An invocational quatrain on

this subject, usually attributed to Byron (see

Lewin 1999, p. 31), occurs in a number of

variations, but the rendering by Lewin (1999) is

presented here:

O Cloacina, goddess of this place,

Look on thy suppliants with smiling face,

Soft yet cohesive let their offerings flow,

Not rashly swift nor insolently slow.

SYSTEMATICS

Cloacina caballeroi Mawson, 1977

Material examined: 73, 132, from stomach of

Dorcopsis hageni, Usino, Madang, Papua New

Guinea, coll. T. Reardon, May 1987, SAM AHC

31425.

Remarks

Cloacina caballeroi was described by Mawson

(1977) and Beveridge (1998) from Dorcopsis

muelleri and Do. luctuosa from Irian Jaya and

Papua New Guinea. This represents the first

record from Do. hageni.

Cloacina cornuta (Davey & Wood, 1938)

Material examined: 23, 12, from stomach of

Macropus agilis, Bula Plain, Bensbach, Papua

New Guinea, coll. I. Owen, May, 1998, BMNH

1998.9.28.24-26.

Remarks

Cloacina cornuta was reported from Macropus

agilis from Port Moresby and Cape Rodney by

Beveridge (1998). The present record adds an

additional locality. Data provided by Dr I. Owen

indicate that in four wallabies examined, numbers

of C. cornuta ranged from 1650-9600 (mean

4700). Although based on a very small number of

wallabies, these data contrast strikingly with those

of Speare et al. (1983) for M. agilis from northern

Australia in which C. cornuta was found in only

41% of hosts examined, and with Beveridge et al.

(1998) who found the nematode in 35% of

wallabies in Queensland. In Australia, C. cornuta

invariably occurs in much lower numbers than the

synhospitalic species C. australis (unpublished

observations).

Cloacina cretheis sp. nov.

(Figs 1-13)

Types: From stomach of Dendrolagus inustus,

Mt Somoro, Sandaun Province, Papua New

Guinea, coll. T. Flannery, 10.iii.1990. Holotype

3, SAM AHC 31426; allotype 2, SAM AHC

31427. Paratypes: 12d, 122, SAM AHC 31428;

1d, 12, BMNH 2001.4.10.1-2; 1¢, 12, USNPC

91133. Slide preparations of male, apical views of

mouth and bursa, SAM AHC 28378.

Material examined: From Dendrolagus inustus:

types. From Dendrolagus dorianus: 13, 49,

Gunung Ki, Tembagapura, Irian Jaya, coll. T.

Flannery, 19.v.1994, SAM AHC 31430; 2d, 10,

Lake Trist, Papua New Guinea, coll. I. Redmond,

BMNH, 1979, 4486-4505. From Dendrolagus

CLOACINA FROM PAPUA NEW GUINEA 3

FIGURES 1-13. Cloacina cretheis sp. nov., specimens from Dendrolagus inustus. 1. Anterior end, lateral view. 2.

Cephalic extremity, lateral view, dorsal aspect on right-hand side. 3. Cephalic extremity, dorsal view. 4. Submedian

cephalic papilla. 5. Cephalic extremity, apical view. 6. Cephalic extremity, transverse optical section through

buccal capsule. 7. Genital cone, ventral view. 8. Genital cone, dorsal view. 9. Gubernaculum, ventral view. 10.

Bursa, apical view. 11. Distal tip of spicule, lateral view. 12. Female tail, lateral view. 13. Female genital system,

lateral view. Scale bars: 1, 10, 12, 13, 0.1 mm; 2-9, 11, 0.01 mm.

4 I. BEVERIDGE

goodfellowi: 5d, 58, Mt Machold, Sandaun

Province, Papua New Guinea, coll. T. Flannery

and K. Vula, 13.xii.1990, SAM AHC 31431.

From Dendrolagus matschiei Forster &

Rothschild, 1907: 26¢, 282, Huon Peninsula,

Papua New Guinea, coll. J. Mayer, July 1999,

USNPC 91146. From Dendrolagus scottae: 93,

52, Sweipini, Sandaun Province, Papua New

Guinea, coll. T. Flannery, 15.vi.1991, SAM AHC

31429.

Description

Small nematodes; cervical cuticle very

slightly inflated to beyond level of excretory

pore; transverse cuticular annulations widely

spaced, 0.030 apart. Submedian cephalic

papillae prominent, 0.015 long, projecting

anteriorly from peri-oral cuticle; distal segment

sub-spherical, 0.005 long, shorter than

cylindrical proximal segment, 0.010 long.

Buccal capsule shallow, symmetrical in dorsal

and lateral views, circular in transverse section,

wider than deep, wall without striations. Leaf

crown elements 8 in number, slightly recurved

at tips; peri-oral cuticle not inflated into lobes

attached to each element. Dorsal sector of

oesophagus bearing opening of dorsal

oesophageal gland, not projecting into lumen of

buccal capsule. Oesophagus slender, claviform,

slightly wider at posterior end; lining without

sclerotised bosses; denticles absent. Nerve ring

in mid-oesophageal region; deirids posterior to

Nerve ring; excretory pore at level of

oesophago-intestinal junction.

Male. Measurements of 5 specimens, types.

Total length 6.7—9.0 (7.8); maximum width 0.35—

0.45 (0.40); buccal capsule 0.010 (0.010) x 0.023—

0.025 (0.023); oesophagus 0.46-0.57 (0.52);

nerve ring to anterior end 0.20—0.24 (0.22);

excretory pore to anterior end 0.39—-0.48 (0.45);

deirid to anterior end 0.28—0.36 (0.32); spicules

2.34-2.62 (2.54); gubernaculum 0.03 (0.03) long.

Measurements of 5 specimens from D.

goodfellowi: total length 7.9-9.3 (8.8); maximum

width 0.48-0.56 (0.51); buccal capsule 0.010—

0.012 (0.011) x 0.023-0.025 (0.023); oesophagus

0.53-0.64 (0.59); nerve ring to anterior end 0.23-

0.24 (0.24); excretory pore to anterior end 0.44—

0.53 (0.47); deirid to anterior end 0.20—-0.34

(0.29); spicules 2.53-2.94 (2.74); gubernaculum

0.030—0.040 (0.034) long.

Measurements of 3 specimens from D. scottae:

total length 6.9-9.7 (8.4); maximum width 0.43—

0.47 (0.45); buccal capsule 0.010 (0.010) x 0.023—

0.025 (0.023); oesophagus 0.48-0.62 (0.56);

nerve ring to anterior end 0.20; excretory pore to

anterior end 0,40-0.46 (0.42); deirid to anterior

end 0.25-0.31 (0.28); spicules 2.63—2.66 (2.65);

gubernaculum 0.030—0.035 (0.032) long.

Measurements of single specimens from D.

dorianus: total length 9.4 ; maximum width 0.43;

buccal capsule 0.010 x 0.025 ; oesophagus 0.50 ;

excretory pore to anterior end 0.50; deirid to

anterior end 0.26; spicules 2.97; gubernaculum

0.035 long.

Dorsal ray elongate; external branchlets arise

at 1/3 length, before major bifurcation; angle of

bifurcation acute; external branchlets as long as

internals, directed postero-laterally, not

reaching margin of bursa; internal branchlets

originate 2/3 along dorsal ray, directed postero-

laterally, almost reaching margin of bursa.

Externo-dorsal ray not reaching margin of

bursa. Gubernaculum prominent, wider than

long. Spicule tip blunt; ala terminates anterior

to spicule tip; anterior lip of genital cone

conical; posterior lip with paired projections

and cuticular inflation of internal surface of

bursa on either side.

Female. Measurements of 5 specimens, types.

Total length 7.7—9.0 (8.2); maximum width 0.52-

0.54 (0.52); buccal capsule 0.008—0.010 (0.010) x

0.023-0.025 (0.023); oesophagus 0.48-0.55

(0.53); nerve ring to anterior end 0.20-0.30

(0.23); excretory pore to anterior end 0.39-0.50

(0.44); deirid to anterior end 0.26-0.33 (0.30); tail

0.23-0.31 (0.27); vulva to posterior end 0.35—0.40

(0.38); vagina 0.50-0.73 (0.62); egg 0.075—0.080

(0.080) x 0.040—0.050 (0.045).

Measurements of 5 specimens from D. scottae:

total length 8.8-10.7 (10.0); maximum width

0.49-0.64 (0.59); buccal capsule 0.008—0.010

(0.009) x 0.020—-0.025 (0.023); oesophagus 0.53—

0.58 (0.55); nerve ring to anterior end 0.20;

excretory pore to anterior end 0.48-0.50 (0.49);

deirid to anterior end 0.29-0,32 (0.30); tail 0.23-

0.32 (0.28); vulva to posterior end 0.38-0.40

(0.39); vagina 0.53—0.58 (0.56).

Measurements of 3 specimens from D.

dorianus: total length 8.8—9.2 (9.0); maximum

width 0.49-0.72 (0.60); buccal capsule 0.010-

0.013 (0.012) x 0.023-0.025 (0.024); oesophagus

0.53-0.60 (0.57); nerve ring to anterior end 0.23

(0.23); excretory pore to anterior end 0.48-0.51

(0.49); deirid to anterior end 0.20—0.29 (0.24); tail

0.21-0.25 (0.23); vulva to posterior end 0.49-0.50

(0.49); vagina 0.64-0.67 (0.66); egg 0.070—0.075

(0.078) x 0.040—-0.050 (0.045).

Female tail short, conical; vagina elongate,

convoluted; egg ellipsoidal.

CLOACINA FROM PAPUA NEW GUINEA 5

Remarks

C. cretheis sp. nov. is characterised by

submedian papillae with a small sub-spherical

distal segment, a simple buccal capsule, eight

elements to the leaf crown, a slender,

unornamented oesophagus, deirids posterior to

the nerve ring, a dorsal ray in which the

external branchlets arise before the major

bifurcation and an elongate, convoluted vagina.

The shape of the dorsal ray alone separates it

from all congeners except C. caballeroi, C.

enyo Beveridge, 1998, C. ips Beveridge, 1998,

and C. syphax. It differs from all of these

except C. caballeroi in having the deirid

posterior to the nerve ring, although in some

highly contracted specimens of C. cretheis, in

which the anterior oesophagus becomes

sinuous, the deirid may appear more anteriorly

on the contracted cuticle. C. cretheis is further

distinguished from C. syphax which has a

sinuous anterior margin to its buccal capsule

and submedian cephalic papillae with enlarged

distal segments. C. cretheis is distinguished

from C. caballeroi by spicule lengths, which

are 2.34—2.97 mm in C. cretheis compared with

1.23-1.45 mm in C. caballeroi. The spicules of

C. enyo are 0.68—0.88 mm long while those of

C. ips are 1.36-1.39 mm in length, providing a

further means of distinguishing the latter

species.

C. cretheis occurs in a wide range of tree

kangaroo species in Papua New Guinea and was

by far the most frequently encountered species in

tree kangaroos in this study. It does not occur in

either Dendrolagus bennettianus DeVis, 1887 or

D. lumholtzi Collett, 1884 in Australia (Spratt et

al. 1991). Although incomplete, the measurements

of specimens from different hosts suggest that

there are no major differences induced by the host

species. Some of the minor differences observed

are due to the state of contraction or of

preservation of specimens, and the incomplete sets

of measurements reflect the poor state of

preservation of many of the specimens, precluding

the measurement of internal organs.

Cloacina cunctabunda sp. nov.

(Figs 14-23)

Types: From stomach of Dendrolagus mbaiso,

Camp Ridge, Tembagapura, Irian Jaya, coll. T.

Flannery, 24.v.1994. Holotype ¢, SAM AHC

31432; allotype 2, SAM AHC 31433. Paratype

3, 3 slides, SAM AHC 28379.

Description

Small nematodes; cervical cuticle slightly

inflated to beyond level of excretory pore;

transverse cuticular annulations widely spaced,

0.015 apart. Submedian cephalic papillae

prominent, 0.010 long, projecting anteriorly from

peri-oral cuticle; distal segment ovoid, 0.006 long,

longer than cylindrical proximal segment, 0.004

long. Buccal capsule shallow, symmetrical in dorsal

and lateral views, oval in transverse section,

slightly deeper dorso-ventrally, wider than deep,

wall without striations. Leaf crown elements 6 in

number, recurved at tips; peri-oral cuticle, inflated;

inflations not attached to each element of leaf

crown. Dorsal sector of oesophagus bearing

opening of dorsal oesophageal gland, not projecting

into lumen of buccal capsule. Oesophagus slender,

claviform, slightly wider at posterior end; lining

without sclerotised bosses; denticles absent. Nerve

ring in mid-oesophageal region; deirids anterior to

nerve ring, mid-way between nerve ring and buccal

capsule; excretory pore anterior to level of

oesophago-intestinal junction.

Male. Measurements of holotype and paratype.

Total length 3.30, 3.32; maximum width 0.16,

0.17; buccal capsule 0.008, 0.010 x 0.025, 0.025;

oesophagus 0.34, 0.35; nerve ring to anterior end

0.16, 0.18; excretory pore to anterior end 0.26,

0.30; deirid to anterior end 0.11, 0.14; spicules

1.39, 1.57; gubernaculum 0.015, 0.015 long.

Dorsal lobe of bursa and dorsal ray elongate,

ray broad at origin; external branchlets arise at 1/2

length, immediately before major bifurcation;

angle of bifurcation acute; external branchlets

shorter than internals, directed postero-laterally,

not reaching margin of bursa; internal branchlets

originate soon after externals arise from dorsal

ray, directed postero-laterally, almost reaching

margin of bursa. Externo-dorsal ray not reaching

margin of bursa. Gubernaculum prominent, wider

than long. Spicule tip not everted. Anterior lip of

genital cone conical; posterior lip with paired

projections.

Female. Measurements of allotype. Total length

4.25; maximum width 0.21; buccal capsule 0.010

x 0.025; oesophagus 0.40; nerve ring to anterior

end 0.18; excretory pore to anterior end 0.29;

deirid to anterior end 0.11; tail 0.35; vulva to

posterior end 0.55; vagina 0.35.

Female tail elongate, conical; vulva

immediately anterior to anus; vagina short with

single convolution; egg not seen.

Remarks

Although described from only three specimens,

6 I. BEVERIDGE

FIGURES 14-23. Cloacina cunctabunda sp. nov., specimens from Dendrolagus mbaiso. 14. Anterior end, lateral

view. 15. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 16. Cephalic extremity, ventral view. 17.

Submedian cephalic papilla, lateral view. 18. Cephalic extremity, apical view. 19. Cephalic extremity, transverse

optical section through buccal capsule. 20. Bursa, apical view. 21. Gubernaculum, ventral view. 22. Female tail,

lateral view. 23. Female genital system, lateral view. Scale bars: 22, 23, 0.1 mm; 14-21 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 7

this species is sufficiently distinctive to allow

ready recognition, and comes from a host

restricted to isolated localities in Irian Jaya which

is unlikely to be sampled extensively in the future

(Flannery et al. 1995). Furthermore, this species

of Cloacina was not found in any other species of

tree kangaroo.

C. cunctabunda is distinguished from all

congeners except C. cretheis, C. caballeroi, C.

enyo, C. ips and C. syphax in possessing a simple

buccal capsule, an elongate, unornamented

oesophagus and the external branchlets of the

dorsal ray arising before the major bifurcation.

The position of the deirid anterior to the nerve

ring distinguishes the species from C. cretheis and

C. caballeroi, while the shape of the buccal

capsule and the submedian cephalic papillae

distinguish it from C. syphax. C. cunctabunda is

distinguished from C. enyo on the basis of spicule

lengths, which are 1.39-1.57 mm long in C.

cunctabunda compared with 0.68-0.88 mm in C.

enyo. C. cunctabunda therefore most closely

resembles C. ips, from which it is distinguished

by the number of leaf crown elements (six in C.

cunctabunda, eight in C. ips), in the absence of a

dorsal oesophageal tooth which projects

prominently into the buccal capsule of C. ips, in

the shape of the female tail which is slender and

elongate in C. cunctabunda but short and conical

in C. ips, and in the shape of the vagina which is

Straight in C. ips but convoluted in C.

cunctabunda.

Cloacina erigone sp. nov.

(Figs 24-36)

Types: From stomach of Dorcopsis hageni,

Usino, Madang, Papua New Guinea, coll. T.

Reardon, May 1987. Holotype 6, SAM AHC

31434; allotype 2, SAM AHC 31435. Paratypes:

43, 52, SAM AHC 31436, 16, 12, BMNH

2001.4.10.3-4, 1d, 12, USNPC 91134. Slide

preparations of apical views of mouth and bursa,

SAM AHC 28380.

Material examined: types.

Description

Small nematodes; cervical cuticle slightly

inflated to beyond level of excretory pore;

transverse cuticular annulations widely spaced,

0.015 apart. Submedian cephalic papillae

prominent, 0.013 long, projecting anteriorly

from peri-oral cuticle; distal segment ovoid,

0.005 long, shorter than cylindrical proximal

segment, 0.008 long. Buccal capsule shallow,

symmetrical in dorsal and lateral views, circular

in transverse section, wider than deep, wall

without striations. Leaf crown elements 8 in

number, slightly recurved at tips, margins

prominently thickened; peri-oral cuticle not

inflated into lobes attached to each element.

Dorsal sector of oesophagus bearing opening of

dorsal oesophageal gland, projecting

prominently into lumen of buccal capsule.

Oesophagus claviform, robust, slightly wider at

posterior end; lining with sclerotised bosses

extending from anterior end, halfway to nerve

ring; single dorsal denticle present in prominent

preneural swelling of oesophagus. Nerve ring in

mid-oesophageal region; deirids posterior to

nerve ring, almost at level of excretory pore;

excretory pore at level of oesophago-intestinal

junction.

Male. Measurements of 8 specimens, types.

Total length 2.88-5.11 (4.02); maximum width

0.18—0.31 (0.26); buccal capsule 0.007—0.010

(0.008) x 0.023—0.035 (0.032); oesophagus 0.26—

0.39 (0.34); nerve ring to anterior end 0.14—0.22

(0.18); excretory pore to anterior end 0.25—0.40

(0.33); deirid to anterior end 0.23-0.37 (0.32);

spicules 1.35-1.79 (1.59); gubernaculum 0.020—

0.030 (0.024) long.

Dorsal ray elongate, broad at origin; external

branchlets arise at 1/2 length, immediately before

major bifurcation; angle of bifurcation acute;

external branchlets shorter than internals, directed

laterally, not reaching margin of bursa; internal

branchlets originate immediately after externals,

directed postero-laterally, almost reaching margin

of bursa. Externo-dorsal ray not reaching margin

of bursa. Gubernaculum prominent, slightly wider

than long. Spicule tip blunt; ala terminates

abruptly anterior to spicule tip; anterior lip of

genital cone conical; posterior lip with paired

projections and cuticular inflation of internal

surface of bursa on either side.

Female. Measurements of 8 specimens, types.

Total length 3.56-4.81 (4.39); maximum width

0.31-0.48 (0.41); buccal capsule 0.008-0.010

(0.009) x 0.033—-0.045 (0.039); oesophagus 0.35—

0.44 (0.39); nerve ring to anterior end 0.16—0.23

(0.20); excretory pore to anterior end 0.21-0.43

(0.34); deirid to anterior end 0.26—-0.37 (0.32); tail

0.16-0.19 (0.17); vulva to posterior end 0.24—0.28

(0.26); vagina 0.53-0.99 (0.75); egg 0.055—0.070

(0.062) x 0.030—0.035 (0.031).

Female tail short, conical; vulva immediately

anterior to anus; vagina elongate, straight; egg

ellipsoidal.

8 I. BEVERIDGE

FIGURES 24-36. Cloacina erigone sp. nov., types. 24. Anterior end, lateral view. 25. Cephalic extremity, lateral

view, dorsal aspect on left-hand side. 26. Cephalic extremity, dorsal view. 27. Cephalic extremity, apical view. 28.

Cephalic extremity, transverse optical section through buccal capsule. 29. Submedian cephalic papilla, lateral view.

30. Preneural oesophageal swelling showing dorsal denticle, dorsal view. 31. Bursa, apical view. 32. Genital cone,

dorsal view. 33. Spicule tip, lateral view. 34. Gubernaculum, ventral view. 35. Female tail, lateral view. 36. Female

genital system, lateral view. Scale bars: 24, 31, 35, 36, 0.1 mm; 25-30, 32-34, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 9

Remarks

Cloacina erigone sp. nov. is characterised by

having submedian cephalic papillae with an

elongate proximal segment, a_ shallow,

unornamented buccal capsule, eight leaf crown

elements, deirid posterior to the nerve ring, a

dorsal ray with the external branchlets arising

before the main bifurcation and a straight vagina.

In these characters, C. erigone resembles C.

caballeroi, C. cretheis, C. enyo, C. ips and C.

syphax. It differs from all of these species,

however, in having bosses lining the anterior part

of the vagina and a single dorsal denticle. Species

with a symmetrical buccal capsule, oesophageal

bosses and a single dorsal denticle are C.

australis, C. dis Beveridge, 1998, C. hecuba

Beveridge, 1998, C. io Beveridge, 1998, C. laius,

C. leto Beveridge, 1998, C. minor (Davey &

Wood, 1938) and C. tyro Beveridge, 1998.

However, in none of these species is the deirid

posterior to the nerve ring, and in none do the

external branchlets of the dorsal ray arise before

the principal bifurcation.

This species has, thus far, been found only in

Do. hageni.

Cloacina eurynome sp. nov.

(Figs 37-49)

Types: From stomach of Dendrolagus dorianus,

Tembagapura, Irian Jaya, coll. T. Flannery,

19.v.1994, 23.v.1994. Holotype 6, SAM AHC

31437; allotype 2, SAM AHC 31438. Paratypes:

43,92, SAM AHC 31439, 1d, 22, SAM AHC

31440, 1d, 12, BMNH 2001.4.10.5-6, 1d, 12,

USNPC 91135. Slide preparations of spicules,

apical views of mouth and bursa, SAM AHC

28381.

Material examined: From Dendrolagus

dorianus: types. From Dendrolagus scottae: 4°,

Sweipini, Sandaun Province, Papua New Guinea,

coll. T. Flannery, 15.vi.1991, SAM AHC 31441.

Description

Small nematodes; cervical cuticle slightly

inflated to beyond level of excretory pore;

transverse cuticular annulations widely spaced,

0.030 apart. Submedian cephalic papillae

prominent, 0.018 long, projecting anteriorly from

peri-oral cuticle; distal segment ovoid, pointed

apically, 0.008 long, shorter than cylindrical

proximal segment, 0.010 long. Buccal capsule

shallow, symmetrical in dorsal and lateral views,

approximately octagonal in transverse section,

wider than deep, wall without striations. Leaf

crown elements 8 in number, slightly recurved at

tips; peri-oral cuticle not inflated into lobes

attached to each element. Dorsal sector of

oesophagus bearing opening of dorsal

oesophageal gland, not projecting into lumen of

buccal capsule. Oesophagus claviform, slender,

only slightly wider at posterior end; lining

without sclerotised bosses; three poorly

developed denticles, one dorsal and two

subventral, present in lumen of oesophagus

immediately anterior to nerve ring; preneural

swelling of oesophagus small. Nerve ring in mid-

oesophageal region; deirids at level of nerve

ring; excretory pore at level of oesophago-

intestinal junction.

Male. Measurements of 7 specimens, types.

Total length 8.75—-10.67 (9.53); maximum width

0.39-0.48 (0.45); buccal capsule 0.007—0.010

(0.008) x 0.027—-0.030 (0.029); oesophagus 0.57—

0.64 (0.59); nerve ring to anterior end 0.26—0.28

(0.27); excretory pore to anterior end 0.54—0.62

(0.58); deirid to anterior end 0.24—0.32 (0.28);

spicules 3.57—3.98 (3.77); gubernaculum 0.015—

0.020 (0.019) long.

Dorsal ray elongate, broad at origin; major

bifurcation occurs at 1/3 length; angle of

bifurcation acute; external branchlets arise mid-

way between major bifurcation and tip, much

shorter than internals, directed laterally or postero-

laterally, not reaching margin of bursa; internal

branchlets directed postero-laterally, almost

reaching margin of bursa. Externo-dorsal ray not

reaching margin of bursa. Gubernaculum

prominent, wider than long. Spicule tip blunt,

gently curved; ala terminates abruptly anterior to

spicule tip; anterior lip of genital cone conical;

posterior lip with paired projections and cuticular

inflation of internal surface of bursa on either

side.

Female. Measurements of 10 specimens, types.

Total length 10.19-13.28 (11.50); maximum

width 0.52—0.75 (0.60); buccal capsule 0.005—

0.013 (0.008) x 0.030—0.035 (0.031); oesophagus

0.61-0.73 (0.65); nerve ring to anterior end 0.25—

0.28 (0.27); excretory pore to anterior end 0.52—

0.71 (0.60); deirid to anterior end 0.18—0.34

(0.27); tail 0.22-0.29 (0.25); vulva to posterior

end 0.31-0.46 (0.39); vagina 1.57-1.85 (1.75);

egg 0.080—0.095 (0.086) x 0.040—0.050 (0.046).

Female tail slender, conical; vulva immediately

anterior to anus; vagina elongate, straight distally,

recurrent anterior to vestibule, recurrent section

twisted in characteristic figure of eight formation;

egg ellipsoidal.

10 I. BEVERIDGE

FIGURES 37-49. Cloacina eurynome sp. nov., types. 37. Anterior end, lateral view. 38. Cephalic extremity, lateral

view, dorsal aspect on right-hand side. 39. Cephalic extremity, ventral view. 40. Submedian cephalic papilla, lateral

view. 41. Cephalic extremity, apical view. 42. Cephalic extremity, transverse optical section through buccal capsule.

43. Preneural oesophageal swelling showing denticles, lateral view, dorsal aspect on right-hand side. 44. Preneural

oesophageal swelling showing denticles, ventral view. 45. Bursa, apical view. 46. Spicule tip, lateral view. 47.

Gubernaculum, genital cone and thickenings of spicule sheaths, dorsal view. 48. Female tail, lateral view. 49.

Female genital system, lateral view. Scale bars: 37, 45, 48, 49, 0.1 mm; 38-44, 46, 47, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 11

Remarks

The presence of three preneural denticles at

the same level in the oesophagus distinguishes

C. eurynome from all congeners except C. hera

Beveridge, 1998, C. hermes Beveridge, 1998

and C. hestia Beveridge, 1998, all of which are

parasites of grey kangaroos, Macropus

fuliginosus (Desmarest, 1817) and M. giganteus

Shaw, 1790 in eastern and southern Australia

(Beveridge 1998) and C. daveyi Mawson, 1977

in the wallaroo, M. robustus Gould, 1841. C.

eurynome is distinguished from the species in

grey kangaroos in having the deirid at the level

of the nerve ring rather than well anterior to it

and in having eight rather than six leaf crown

elements. It is distinguished from C. daveyi

which has cephalic papillae with large, globose,

medially directed distal segments. The denticles

in C. eurynome are vestigial, a feature found

only in C. hestia. C. eurynome is distinguished

from all congeners other than C. syphax, from

Dorcopsulus vanheurni, also from Papua New

Guinea, in the morphology of the vagina, which

exhibits a highly characteristic recurrent loop,

anterior to the vestibule, twisted in a figure of

eight formation. C. eurynome differs from C.

Syphax in possessing denticles, in having a

straight rather than a sinuous anterior margin to

the buccal capsule, and in having the external

branchlets of the dorsal ray arise after the major

bifurcation

Cloacina hecale sp. nov.

(Figs 50-61)

Types: From stomach of Dendrolagus

dorianus, Lake Trist, Papua New Guinea, coll. I.

Redmond, 1979. Holotype 36, BMNH

1981.4506; allotype 2, BMNH 1981.4507.

Paratypes: 49d, 882, BMNH 1981.4508-4535.

Slide preparations of apical views of bursa and

mouth, SAM AHC 28383.

Material examined: From Dendrolagus

dorianus: types.

Description

Robust nematodes; cervical cuticle slightly

inflated to beyond level of excretory pore;

transverse cuticular annulations widely spaced,

0.023 apart. Submedian cephalic papillae

prominent, 0.017 long, projecting anteriorly from

peri-oral cuticle; distal segment conical, 0.007

long, only slightly shorter than cylindrical

proximal segment, 0.010 long. Buccal capsule

shallow, symmetrical in dorsal and lateral views,

circular in transverse section, wider than deep,

wall without striations; anterior margin slightly

undulant. Leaf crown elements 8 in number,

slightly recurved at tips; peri-oral cuticle not

inflated into lobes attached to each element.

Dorsal sector of oesophagus bearing opening of

dorsal oesophageal gland, not projecting into

lumen of buccal capsule. Oesophagus claviform,

slender, slightly wider at posterior end; lining

without sclerotised bosses; denticles absent. Nerve

ring in anterior oesophageal region; deirids

anterior to nerve ring; excretory pore at level of

oesophago-intestinal junction.

Male. Measurements of 10 specimens, types.

Total length 9.44-13.86 (11.82); maximum width

0.50-0.67 (0.60); buccal capsule 0.010-0.013

(0.011) x 0.035—0.040 (0.039); oesophagus 0.76—

0.95 (0.87); nerve ring to anterior end 0.30—-0.32

(0.31); excretory pore to anterior end 0.52-0.98

(0.80); deirid to anterior end 0.17-0.26 (0.23);

spicules 3.90-5.05 (4.54); gubernaculum 0.040

long.

Dorsal ray broad at origin; major bifurcation

occurs at 1/2 length; external branchlets arise

immediately after major bifurcation, as long as

internals, though more robust, directed postero-

laterally, not reaching margin of bursa; internal

branchlets directed posteriorly, almost reaching

margin of bursa. Externo-dorsal ray not reaching

margin of bursa. Gubernaculum prominent, wider

than long. Spicule minutely bifid at tip; ala

terminates abruptly anterior to spicule tip; anterior

lip of genital cone conical; posterior lip with

paired projections and cuticular inflation of

internal surface of bursa on either side.

Female. Measurements of 5 specimens, types.

Total length 19.6-22.9 (21.3); maximum width

0.64-0.91 (0.81); buccal capsule 0.010 (0.010) x

0.040 (0.040); oesophagus 0.95-1.02 (0.97);

nerve ring to anterior end 0.25-0.29 (0.27);

excretory pore to anterior end 0.63—0.86 (0.70);

deirid to anterior end 0.19-0.29 (0.22); tail 0.13-

0.30 (0.21); vulva to posterior end 0.18-0.43

(0.28); vagina 2.22—2.57 (2.40); vestibule 0.27;

sphincter 0.22; infundibulum 0.17; egg 0.090—

0.110 (0.096) x 0.045-0.050 (0.048).

Female tail short, conical; vulva immediately

anterior to anus; vagina elongate, straight, extends

slightly anterior to vestibule with short recurrent

section; egg ellipsoidal.

Remarks

Cloacina hecale sp. nov. is a robust species

characterised by long spicules and an elongate

I. BEVERIDGE

FIGURES 50-61. Cloacina hecale sp. nov., types. 50. Anterior end, lateral view. 51. Cephalic extremity, lateral

view, dorsal aspect on right-hand side. 52. Cephalic extremity, dorsal view. 53. Submedian cephalic papilla, lateral

view. 54. Cephalic extremity, apical view. 55. Cephalic extremity, transverse optical section through buccal capsule.

56. Bursa, apical view. 57. Spicule tip, lateral view. 58. Gubernaculum, ventral view. 59. Genital cone, dorsal view.

60. Female tail, lateral view. 61. Female genital system, lateral view. Scale bars: 50, 56, 60, 61, 0.1 mm; 51-55,

57-59, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 13

vagina, slightly recurrent at its anterior extremity.

The length of the vagina relative to the sizes of

vestibule, sphincter and infundibulum are given

in the description in this case to emphasise the

extreme length of the vagina. The other features

of the species are unremarkable, with cephalic

papillae bearing a distal segment with an acute

tip, almost equal in length to the proximal

segment, eight leaf crown elements, a

symmetrical buccal capsule, a_ slender,

unornamented oesophagus, the deirid anterior to

the nerve ring and the external branchlets of the

dorsal ray arising after the principal bifurcation.

These features together with the length of the

spicules (> 3.0 mm) differentiate C. hecale from

congeners except C. clymene Beveridge, 1998,

C. curta Johnston & Mawson, 1938, C. liebigi

Johnston & Mawson, 1938, C. longispiculata

Johnston & Mawson, 1939, C. nike Beveridge,

1998, C. robertsi Johnston & Mawson, 1939, C.

smalesae Mawson, 1975 and C. solymus. C.

hecale is differentiated from C. clymene and C.

robertsi which have six elements to the leaf

crown and lip-like inflations of the peri-oral

cuticle attached to each element of the leaf

crown. C. curta, C. liebigi, C. longispiculata and

C. smalesae all have six leaf crown elements

rather than eight and the vagina is prominently

recurrent in these species rather than having a

short anterior recurrent section. In addition, C.

liebigi, C. longispiculata and C. smalesae have

the cervical cuticle inflated so as to form

‘shoulders’ in the oesophageal region. C. nike

has submedian cephalic papillae in which the

distal segment is much shorter than the proximal

segment and has a sinuous vagina and a longer,

slender tail in the female. C. solymus has a

robust subcylindrical oesophagus but the

submedian cephalic papillae are tiny and the

anterior margin of the buccal capsule is undulate.

Therefore, C. hecale is readily distinguishable

from all known congeners.

Cloacina hyperea sp. nov.

(Figs 62-74)

Types: From stomach of Dorcopsis hageni,

Usino, Madang, Papua New Guinea, coll. T.

Reardon, May, 1987. Holotype 6, SAM AHC

31442; allotype 2, SAM AHC 31443. Paratypes:

12d, 82, SAM AHC 31444, 1¢, 12, BMNH

2001.4.10.7-8, 1d, 12, USNPC 91136. Slide

preparations of apical views of mouth and bursa,

SAM AHC 28382.

Material examined: From Dorcopsis hageni:

types.

Description

Robust neniatodes; cervical cuticle slightly

inflated to beyond level of excretory pore;

transverse cuticular annulations widely spaced,

0.034 apart. Submedian cephalic papillae

prominent, 0.015 long, projecting anteriorly from

peri-oral cuticle; distal segment conical, 0.007

long, only slightly shorter than cylindrical

proximal segment, 0.008 long. Buccal capsule

shallow, symmetrically arched anteriorly in lateral

views; in dorsal views, buccal capsule arches

anteriorly over dorsal oesophageal gland; in

ventral views, curves posteriorly; oval and dorso-

ventrally elongate in transverse section, wider than

deep, wall with prominent striations; anterior

margin smooth, except on dorsal aspect. Leaf

crown elements 6 in number, slightly recurved at

tips; peri-oral cuticle slightly inflated into lobes

attached to each element. Dorsal sector of

oesophagus bearing opening of dorsal

oesophageal gland, projecting prominently into

lumen of buccal capsule. Oesophagus claviform,

robust; lining with sclerotised bosses extending to

level of nerve ring; single dorsal oesophageal

denticle present in prominent preneural swelling.

Nerve ring in mid-oesophageal region; deirids at

level of nerve ring; excretory pore at level of, or

slightly anterior to, oesophago-intestinal junction.

Male. Measurements of 10 specimens, types.

Total length 4.20—-5.21 (4.56); maximum width

0.26-0.40 (0.34); buccal capsule 0.013-0.020

(0.016) x 0.055—0.060 (0.056); oesophagus 0.42—

0.50 (0.47); nerve ring to anterior end 0.20-0.24

(0.22); excretory pore to anterior end 0.35-0.51

(0.43); deirid to anterior end 0.25—0.34 (0.31);

spicules 2.25-2.49 (2.37); gubernaculum 0.020—

0.030 (0.026) long.

Dorsal ray broad at origin; major bifurcation

occurs at 1/2 length; angle of bifurcation acute;

external branchlets arise after major bifurcation,

as long as internals, directed postero-laterally, not

reaching margin of bursa; internal branchlets

directed posteriorly, not reaching margin of bursa.

Externo-dorsal ray not reaching margin of bursa.

Gubernaculum prominent, slightly wider than

long. Spicule tip blunt; ala diminishes gradually

in width towards spicule tip; anterior lip of genital

cone conical; posterior lip with paired projections

and cuticular inflation of internal surface of bursa

on either side.

Female. Measurements of 6 specimens, types.

Total length 5.25-6.14 (5.67); maximum width

14 I. BEVERIDGE

SosO O08

oscars

2000090000

. “aN | |

FIGURES 62-74. Cloacina hyperea sp. nov., types. 62. Anterior end, lateral view. 63. Cephalic extremity, lateral

view, dorsal aspect on right-hand side. 64. Cephalic extremity, dorsal view. 65. Cephalic extremity, ventral view.

66. Submedian cephalic papilla, lateral view. 67. Cephalic extremity, apical view. 68. Cephalic extremity, transverse

optical section through buccal capsule. 69. Preneural swelling of oesophagus, dorsal view, showing denticle. 70.

Bursa, apical view. 71. Spicule tip, lateral view. 72. Gubernaculum and genital cone, dorsal view. 73. Female tail,

lateral view. 74. Female genital system, lateral view. Scale bars: 62, 70, 73, 74, 0.1 mm; 63-69, 71,72, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 15

0.32-0.45 (0.39); buccal capsule 0.013-0.018

(0.016) x 0.058—-0.060 (0.059); oesophagus 0.46—

0.52 (0.49); nerve ring to anterior end 0.20-0.23

(0.22); excretory pore to anterior end 0.35—0.50

(0.45); deirid to anterior end 0.24—0.33 (0.29); tail

0.19—-0.26 (0.23); vulva to posterior end 0.36—0.43

(0.39); vagina 0.96—-1.24 (1.07).

Female tail slender, conical; vulva immediately

anterior to anus; vagina elongate, sinuous, extends

slightly anterior to vestibule with short recurrent

section; egg not seen.

Remarks

The anteriorly arched buccal capsule wall

distinguishes C. hyperea sp. nov. from all

congeners except C. circe Beveridge, 1999 and C.

laius Beveridge, 1999, both of which occur in the

quokka, Setonix brachyurus (Quoy & Gaimard,

1830) in the south-west of Western Australia, and

from C. nephele sp. nov., described below. C.

hyperea is distinguished from C. circe in

possessing sclerotised bosses and a dorsal denticle

in the oesophagus. It is differentiated from C.

laius in having the buccal capsule less

prominently arched, in having submedian cephalic

papillae of a different shape, with both segments

of approximately equal length, whereas in C. laius

the proximal segment is almost twice as long as

the distal segment. In addition, the spicules are

2.25—2.49 mm long in C. hyperea compared with

1.50-1.97 mm in C. laius and the vagina is

commensurately longer, being 0.96-1.24 mm in

C. hyperea compared with 0.71-0.92 mm in C.

laius. In C. hyperea the internal and external

branchlets of the dorsal ray are of approximately

equal length whereas in C. laius the external

branchlets are much shorter than the internal

branchlets. C. nephele sp. nov. has lateral lips,

which are absent in C. hyperaea, and a bulbous

proximal segment to the cephalic papilla.

Cloacina nephele sp. nov.

(Figs 75-89)

Types: From stomach of Dorcopsis hageni,

Usino, Madang, Papua New Guinea, coll. T.

Reardon, May, 1987. Holotype d6, SAM AHC

31445; allotype 2, SAM AHC 31446. Paratypes:

3d, 22, SAM AHC 31447, 16, BMNH

2001.4.10.9, 16, USNPC 91137. Slide

preparations of apical views of mouth and bursa,

SAM AHC 28384.

Material examined: From Dorcopsis hageni:

types.

Description

Robust nematodes; cervical cuticle prominently

inflated to beyond level of excretory pore;

transverse cuticular annulations widely spaced,

0.040 apart. Submedian cephalic papillae

prominent, 0.020 long, projecting anteriorly from

inflated peri-oral cuticle; distal segment slender,

conical, 0.010 long, as long as robust,

asymmetrical proximal segment, 0.010 long.

Buccal capsule shallow, symmetrically arched

anteriorly in lateral views; dorsally, arches

anteriorly over dorsal oesophageal gland;

ventrally, curves posteriorly; oval and dorso-

ventrally elongate in transverse section, wider than

deep, wall with prominent striations; anterior

margin smooth. Leaf crown elements 6 in number,

recurved at tips; peri-oral cuticle inflated into lip-

like lobes attached to each element; extra dorsal

and ventral projections of peri-oral cuticle present,

separate from lateral arcades of amphids and

submedian papillae, giving the appearance of lips.

Dorsal sector of oesophagus bearing opening of

dorsal oesophageal gland, not projecting into

lumen of buccal capsule. Oesophagus claviform,

robust; lining without sclerotised bosses; denticles

absent in preneural region. Nerve ring in mid-

oesophageal region; deirids at level of nerve ring;

excretory pore between nerve ring and oesophago-

intestinal junction.

Male. Measurements of 5 specimens, types.

Total length 6.08—7.40 (6.66); maximum width

0.33-0.37 (0.35); buccal capsule 0.023-0.030

(0.026) x 0.090 (0.090); oesophagus 0.69-0.73

(0.71); nerve ring to anterior end 0.28—0.30

(0.29); excretory pore to anterior end 0.48-0.53

(0.51); deirid to anterior end 0.31-0.36 (0.34);

spicules 1.96-2.10 (2.03); gubernaculum 0.035—

0.040 (0.039) long.

Dorsal ray broad at origin, narrowing

posteriorly; major bifurcation occurs at 1/2 length;

angle of bifurcation acute; external branchlets

arise after major bifurcation, shorter than

internals, directed postero-laterally, not reaching

margin of bursa; internal branchlets directed

posteriorly, almost reaching margin of bursa.

Externo-dorsal ray not reaching margin of bursa.

Gubernaculum prominent, slightly wider than

long. Spicule tip blunt; ala diminishes gradually

in width towards spicule tip; anterior lip of genital

cone conical; posterior lip with paired projections

and cuticular inflation of internal surface of bursa

on either side.

Female. Measurements of 3 specimens, types.

Total length 5.94-7.87 (7.02); maximum width

0.34—0.63 (0.50); buccal capsule 0.025—0.030

16 I. BEVERIDGE

FIGURES 75-89. Cloacina nephele sp. nov., types. 75. Anterior end, lateral view. 76. Cephalic extremity, lateral

view, dorsal aspect on right-hand side. 77. Cephalic extremity, lateral view, dorsal aspect on right-hand side,

median optical section showing leaf crown elements and inflation of cephalic collar. 78. Cephalic extremity, dorsal

view. 79. Cephalic extremity, ventral view. 80. Cephalic extremity, ventral view, median optical section showing

leaf crown elements. 81. Submedian cephalic papilla, lateral view. 82. Cephalic extremity, apical view. 83. Cephalic

extremity, transverse optical section through buccal capsule. 84. Bursa, apical view. 85. Genital cone, dorsal view.

86. Gubernaculum, ventral view. 87. Spicule tip, lateral view. 88. Female tail, lateral view. 89. Female genital

system, lateral view. Scale bars: 75, 84, 88, 89, 0.1 mm; 76-83, 85-87, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 17

(0.027) x 0.100 (0.100); oesophagus 0.72-0.79

(0.76); nerve ring to anterior end 0,29-0.32

(0.30); excretory pore to anterior end 0.46-0.58

(0.52); deirid to anterior end 0.30-0.31 (0.31); tail

0.31-0.39 (0.35); vulva to posterior end 0.43-0.55

(0.48); vagina 0.51-0.66 (0.59); egg 0.080—0.085

(0.083) x 0.040-0.045 (0.043).

Female tail slender, conical; vulva immediately

anterior to anus; vagina elongate, sinuous, extends

slightly anterior to vestibule with short recurrent

section; egg ellipsoidal.

Remarks

C. nephele sp. nov. closely resembles C. circe,

C. laius and C. hyperea in having an anteriorly

arched buccal capsule. It differs from C. laius and

C. hyperea in lacking oesophageal bosses and

denticles, and differs from C. circe in having a

markedly inflated cervical cuticle, cephalic

papillae in which the distal segment is not

oriented medially, and a recurrent vagina. It

differs from all of these species in having a

swollen cephalic collar and lip-like inflations of

the peri-oral cuticle attached to each leaf crown

element. The dorsal and ventral ‘lips’ are unique

within the genus.

Cloacina oweni sp. nov.

(Figs 90-101)

Synonyms: Cloacina sp. nov. of Beveridge,

1998, p. 506 (Macropus agilis).

Types: From stomach of Macropus agilis, Bula

Plain, Bensbach, Papua New Guinea, coll. I.

Owen, May, 1998. Holotype ¢, SAM AHC

31448; allotype 9, SAM AHC 31449. Paratypes:

51d, 552, SAM AHC 31450, 1d, 12, BMNH

1998.9.28.11-12.

Material examined: From Macropus agilis:

types; 1d, 19, Dari, Bensbach, Papua New

Guinea, SAM AHC 11719.

Description

Robust nematodes; cervical cuticle slightly

inflated to level of excretory pore; transverse

cuticular annulations widely spaced, 0.030 apart.

Submedian cephalic papillae prominent, 0.013

long, projecting anteriorly from inflated peri-oral

cuticle; distal segment ovoid, 0.004 long, shorter

than cylindrical proximal segment, 0.008 long.

Buccal capsule shallow, symmetrical in lateral and

dorso-ventral views; octagonal in transverse

section, wider than deep, wall without prominent

Striations; anterior margin smooth. Leaf crown

elements 8 in number, recurved at tips; peri-oral

cuticle inflated into lip-like lobes attached to each

element. Dorsal sector of oesophagus bearing

opening of dorsal oesophageal gland, not

projecting into lumen of buccal capsule.

Oesophagus claviform, robust; lining without

sclerotised bosses; single dorsal denticle present

at level of nerve ring; preneural swelling absent.

Nerve ring in mid-oesophageal region; deirids in

anterior oesophageal region; excretory pore

between nerve ring and oesophago-intestinal

junction.

Male. Measurements of 10 specimens, types.

Total length 5.48-7.50 (6.73); maximum width

0.26-0.43 (0.34); buccal capsule 0.015-0.025

(0.019) x 0.045-0.070 (0.056); oesophagus 0.53-

0.65 (0.60); nerve ring to anterior end 0.25—0.32

(0.27); excretory pore to anterior end 0.26-0.53

(0.44); deirid to anterior end 0.10—0.16 (0.12);

spicules 2.19-2.67 (2.46); gubernaculum 0.025—

0.040 (0.031) long.

Dorsal ray broad at origin; major bifurcation

occurs at 1/3 length; angle of bifurcation obtuse;

external branchlets arise after major bifurcation,

near extremity of ray, slightly longer and more

robust than internals, directed postero-laterally,

not reaching margin of bursa; internal branchlets

directed postero-laterally, not reaching margin of

bursa. Externo-dorsal ray not reaching margin of

bursa. Gubernaculum prominent, slightly wider

than long. Spicule tip blunt, slightly recurved; ala

diminishes gradually in width towards spicule tip;

anterior lip of genital cone conical; posterior lip

with paired projections and cuticular inflation of

internal surface of bursa on either side.

Female. Measurements of 10 specimens, types.

Total length 6.70—9.36 (8.06); maximum width

0.30-0.61 (0.46); buccal capsule 0.015—0.025

(0.020) x 0.045-0.075 (0.057); oesophagus 0.55—

0.76 (0.64); nerve ring to anterior end 0,240.33

(0.28); excretory pore to anterior end 0.32-0.61

(0.48); deirid to anterior end 0.09-0.16 (0.11); tail

0.14—0.21 (0.17); vulva to posterior end 0.27—0.34

(0.31); vagina 0.45—0.70 (0.58); egg 0.090-0.100

(0.093) x 0.040—0.050 (0.048).

Female tail short, conical; vulva immediately

anterior to anus; vagina elongate, convoluted, not

recurrent; egg ellipsoidal.

Remarks

The prominent lips and symmetrical buccal

capsule distinguish C. oweni sp. nov. from all

congeners except C. artemis Beveridge, 1998, C.

caenis Beveridge, 1998, C. clymene, C.

dindymene Beveridge, 1998, C. hypsipyle, C.

18 I. BEVERIDGE

FIGURES 90-101. Cloacina oweni sp. nov., types. 90. Anterior end, lateral view. 91. Cephalic extremity, lateral

view, dorsal aspect on left-hand side. 92. Cephalic extremity, dorsal view. 93. Cephalic extremity, apical view. 94.

Cephalic extremity, transverse optical section through buccal capsule. 95. Submedian cephalic papilla, lateral view.

96. Preneural region of oesophagus, showing denticle, dorsal view. 97. Bursa, apical view. 98. Genital cone, dorsal

view and gubernaculum. 99. Spicule tip, lateral view. 100. Female tail, lateral view. 101. Female genital system,

lateral view. Scale bars: 90, 97, 100, 101, 0.1 mm; 91—96, 98, 99, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 19

linstowi Johnston & Mawson, 1940, C. maia

Beveridge, 1998, C. parva, C. robertsi, C.

smalesae, C. thetidis Johnston & Mawson, 1939

and C. wallabiae Johnston & Mawson, 1939. The

presence of a single dorsal denticle in the

oesophagus and the lack of oesophageal bosses

distinguishes it from all of these species except C.

dindymene. It differs from C. dindymene in the

shape of the submedian cephalic papillae which

have elongated distal segments in C. dindymene,

and in the shape of the dorsal ray, the bifurcations

of which form an acute angle in C. dindymene

rather than the obtuse angle seen in C. oweni. In

features of the head, C. oweni is most likely to be

confused with C. robertsi, found in rock wallabies

of the genus Petrogale Gray, 1837 in eastern

Australia. C. oweni is readily differentiated,

however, by the oesophageal denticle and the lack

of a prominently recurrent vagina as exhibited by

C. robertsi.

C. oweni is abundant in the stomachs of agile

wallabies in the Bula Plain region of Papua New

Guinea, but has not been found in the same host

species in northern Australia in spite of relatively

intensive examination of this host in all of the

northern states (Speare et al. 1983; Beveridge et

al. 1998). More surprising is the close

resemblance of this species to congeners

occurring in rock wallabies in Australia, a

relationship for which no simple explanation

currently exists.

Cloacina papuensis sp. nov.

(Figs 102-115)

Types: From stomach of Macropus agilis, Bula

Plain, Bensbach, Papua New Guinea, coll. I.

Owen, May, 1998. Holotype 6, SAM AHC

31451; allotype 2, SAM AHC 31452. Paratypes:

53, 62, SAM AHC 31453, 1¢, 12, BMNH

1998.9.28.13-14. Slide preparations of apical

view of mouth and bursa, SAM AHC 28386.

Material examined: From Macropus agilis:

types.

Description

Small nematodes; cervical cuticle not inflated;

transverse cuticular annulations widely spaced,

0.010 apart. Submedian cephalic papillae small,

conical, 0.006 long, projecting anteriorly from

peri-oral cuticle; distal segment ovoid, 0.002 long,

shorter than cylindrical proximal segment, 0.004

long. Buccal capsule shallow, symmetrical in

lateral and dorso-ventral views; circular in

transverse section, wider than deep, wall without

prominent striations; anterior margin smooth. Leaf

crown elements 8 in number, recurved at tips;

peri-oral cuticle not inflated into lip-like lobes

attached to each element. Dorsal sector of

oesophagus bearing opening of dorsal

oesophageal gland in prominent, bilobed, tooth-

like structure projecting into lumen of buccal

capsule; each ventral sector with 2 triangular

sclerotised projections. Oesophagus claviform,

slender; lining without sclerotised bosses; single

dorsal denticle present immediately anterior to

nerve ring; preneural swelling present. Nerve ring

in mid-oesophageal region; deirids in anterior

oesophageal region; excretory pore between nerve

ring and oesophago-intestinal junction.

Male. Measurements of 5 specimens, types.

Total length 3.18-5.09 (4.57); maximum width

0.18-0.32 (0.23); buccal capsule 0.007-—0.010

(0.008) x 0.020—0.026 (0.021); oesophagus 0.31-

0.41 (0.36); nerve ring to anterior end 0.16-0.17

(0.17); excretory pore to anterior end 0.20—0.34

(0.28); deirid to anterior end 0.08-0.11 (0.10);

spicules 1.84—2.26 (2.05); gubernaculum 0.025—

0.030 (0.029) long.

Dorsal ray broad at origin; major bifurcation

occurs at 1/2 length; angle of bifurcation acute;

external branchlets arise after major bifurcation,

shorter and more robust than internals, directed

postero-laterally, not reaching margin of bursa;

internal branchlets directed posteriorly, not

reaching margin of bursa. Externo-dorsal ray not

reaching margin of bursa. Gubernaculum

prominent, slightly wider than long. Spicule tip

bifid; ala diminishes gradually in width towards

spicule tip; anterior lip of genital cone conical;

posterior lip with paired projections and cuticular

inflation of internal surface of bursa on either

side.

Female. Measurements of 5 specimens, types.

Total length 5.14-6.05 (5.57); maximum width

0.25-0.36 (0.32); buccal capsule 0.005—0.008

(0.007) x 0.020—0.025 (0.023); oesophagus 0.35—

0.39 (0.38); nerve ring to anterior end 0.17-0.18

(0.17); excretory pore to anterior end 0.23-0.38

(0.33); deirid to anterior end 0.07-0.011 (0.09);

tail 0.14—0.22 (0.19); vulva to posterior end 0.30—

0.44 (0.39); vagina 0.47—-0.63 (0.53); egg 0.070—

0.085 (0.080) x 0.040—0.050 (0.046).

Female tail short, conical; vulva immediately

anterior to anus; vagina elongate, convoluted, not

recurrent; egg ellipsoidal.

Remarks

The simple, symmetrical buccal capsule, lack of

20 I. BEVERIDGE

FIGURES 102-115. Cloacina papuensis sp. nov., types. 102. Anterior end, lateral view. 103. Cephalic extremity,

lateral view, dorsal aspect on left-hand side. 104. Cephalic extremity, dorsal view. 105. Submedian cephalic papilla,

lateral view. 106. Cephalic extremity, apical view. 107. Cephalic extremity, transverse optical section through

buccal capsule. 108. Preneural region of oesophagus, showing denticle, lateral view, dorsal aspect on left-hand side.

109. Preneural region of oesophagus, showing denticle, dorsal view. 110. Spicule tips, ventral view. 111. Bursa,

ventral view. 112. Bursa, lateral view. 113. Female tail, lateral view. 114. Female genital system, lateral view. 115.

Genital cone, dorsal view. Scale bars: 113, 114, 0.1 mm; 102-104, 106-112, 115 0.01 mm; 105, 0.003 mm.

CLOACINA FROM PAPUA NEW GUINEA 21

prominent lips and unornamented oesophagus

with a single dorsal denticle differentiate C.

papuensis sp. nov. from all congeners except C.

cornuta, C. dirce, C. longispiculata and C. sciron.

C. papuensis is differentiated from C. sciron

primarily in having the deirid anterior to, rather

than at the level of, the nerve ring. In C. sciron,

the anterior margin of the buccal capsule arches

anteriorly and is sinuous. C. papuensis differs

from C. longispiculata in lacking the prominent

shoulder-like inflations of the cervical cuticle, in

having the excretory pore at the level of the

oesophago-intestinal junction rather than well

posterior to it as in C. longispiculata, and in

having eight leaf crown elements rather than the

six present in C. longispiculata. It differs from C.

dirce in lacking the prominent cervical inflation

of the cuticle, in the shape of the submedian

papillae which have an extended distal segment in

C. dirce, in spicule lengths (1.84—2.26 (2.05) mm

long in C. papuensis, 3.48-3.95 (3.70) mm long

in C. dirce), and in the shape of the vagina which

is longer and more convoluted in C. dirce.

Therefore, C. papuensis most closely resembles

C. cornuta, also a parasite of Macropus agilis, in

having a prominent dorsal oesophageal tooth. It

differs from C. cornuta in lacking a cervical

cuticular inflation, in the shape of the submedian

papillae which have elongate distal segments in

C. cornuta, in the shape of the dorsal ray which in

C. cornuta terminates in very short subequal

internal and external branchlets, in spicule length

(1.84-2.26 (2.05) in C. papuensis, 1.38-1.62

(1.51) mm in C. cornuta), and in the length and

shape of the female tail which is very short and

prominently swollen in C. cornuta.

Cloacina polymela sp. nov.

(Figs 116-128)

Types: From stomach of Dorcopsis hageni,

Usino, Madang, Papua New Guinea, coll. T.

Reardon, May, 1987. Holotype 6, SAM AHC

31454; allotype 2, SAM AHC 31455. Paratypes:

43, 82, SAM AHC 31456, 1d, 12, BMNH

2001.4,10.10-11, 1d, 12, USNPC 91138. Slide

preparations of apical views of mouth and bursa,

SAM AHC 28387.

Material examined: From Dorcopsis hageni:

types.

Description

Robust nematodes; anterior extremity deviated

dorsally; cervical cuticle inflated to level of

excretory pore; transverse cuticular annulations

widely spaced, 0.030—0.062 apart. Submedian

cephalic papillae prominent, 0.016 long,

projecting anteriorly from inflated peri-oral

cuticle; distal segment acute, conical, 0.010 long,

longer than cylindrical proximal segment, 0.006

long. Cephalic collar prominently inflated on

dorsal and ventral aspects, anterior margin of

collar rugose; amphids conical, projecting above

level of cephalic collar. Buccal capsule very

shallow, symmetrical, anterior margin highly

sinuous; due to dorsal deviation of anterior

extremity, buccal capsule appears to arch dorsally

in ventral views; oval and dorso-ventrally elongate

in transverse section, wider than deep, wall finely

striated. Leaf crown elements 8 in number,

recurved at tips; peri-oral cuticle inflated into lip-

like lobes attached to each element. Dorsal sector

of oesophagus bearing opening of dorsal

oesophageal gland, not projecting into lumen of

buccal capsule. Oesophagus claviform, robust,

anterior section of greater diameter than region

posterior to nerve ring; lining without sclerotised

bosses; denticles absent in preneural region. Nerve

ring posterior to mid point of oesophagus; deirids

slightly anterior to level of nerve ring; excretory

pore between nerve ring and oesophago-intestinal

junction.

Male. Measurements of 9 specimens, types.

Total length 5.86-6.03 (5.93); maximum width

0.29-0.46 (0.39); buccal capsule 0.020-—0.030

(0.025) x 0.115—0.150 (0.130); oesophagus 0.66—

0.79 (0.72); nerve ring to anterior end 0.44-0.53

(0.48); excretory pore to anterior end 0.55—0.70

(0.65); deirid to anterior end 0.36-0.45 (0.39);

spicules 2.61—2.71 (2.65); gubernaculum 0.035—

0.050 (0.045) long.

Dorsal ray broad at origin; major bifurcation

occurs at 1/2 length; angle of bifurcation acute;

external branchlets arise after major bifurcation,

much shorter than internals, directed postero-

laterally, not reaching margin of bursa; internal

branchlets elongate, directed posteriorly, almost

reaching margin of bursa. Externo-dorsal ray not

reaching margin of bursa. Gubernaculum

prominent, subcordate. Spicule tip blunt, slightly

recurved; ala diminishes gradually in width

towards spicule tip; anterior lip of genital cone

conical; posterior lip with paired projections and

cuticular inflation of internal surface of bursa on

either side.

Female. Measurements of 10 specimens, types.

Total length 5.45-8.22 (7.14); maximum width

0.33-0.59 (0.46); buccal capsule 0.015—0.030

(0.025) x 0.120—0.160 (0.150); oesophagus 0.70—

22 I. BEVERIDGE

124

FIGURES 116-128. Cloacina polymela sp. nov., types. 116. Anterior end, lateral view, showing dorsal deviation of

head. 117. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 118. Cephalic extremity, ventral view.

119. Cephalic extremity, dorsal view. 120. Cephalic extremity, apical view. 121. Cephalic extremity, transverse

optical section through buccal capsule. 122. Submedian cephalic papilla, lateral view. 123. Amphid, lateral view.

124. Bursa, ventral view. 125. Spicule tip, ventral view. 126. Gubernaculum and genital cone, dorsal view. 127.

Female tail, lateral view. 128. Female genital system, lateral view. Scale bars: 116, 124, 127, 128, 0.1 mm; 117-

123, 125, 126, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 23

0.94 (0.82); nerve ring to anterior end 0.45-0.61

(0.54); excretory pore to anterior end 0.55—0.85

(0.75); deirid to anterior end 0.31-0.50 (0.42); tail

0.21—0.30 (0.24); vulva to posterior end 0.33-0.50

(0.38); vagina 0.49-0.68 (0.60); egg 0.080-0.085

(0.082) x 0.040—-0.045 (0.042).

Female tail slender, conical; vulva immediately

anterior to anus; vagina elongate, sinuous, extends

slightly anterior to vestibule with short recurrent

section; egg ellipsoidal.

Remarks

Cloacina polymela sp. nov. is a highly

distinctive species which differs from all

congeners in having the anterior extremity

deviated dorsally. This feature appears in every

specimen and therefore is not likely to be a

fixation artefact. In addition, it differs from

congeners except C. dryope Beveridge, 1998 and

C. sappho in having an oesophagus in which the

preneural region is of greater diameter than the

posterior part. C. polymela is distinguished from

C. dryope in having eight rather than six leaf

crown elements, in having a more posterior deirid,

in having the excretory pore anterior to the

oesophago-intestinal junction rather than posterior

to it, and in the shape of the buccal capsule which

is extremely shallow in C. dryope and in which

the anterior margin has only slight saliences rather

than prominent undulations. C. polymela and C.

sappho are the only two members of the genus in

which the amphids form acutely pointed conical

projections above the cephalic collar. In all other

species, the amphids are dome-shaped and do not

project obviously beyond the collar. C. polymela

also resembles C. sappho in the shape of the

dorsal ray of the bursa and in possessing eight

leaf crown elements. However, it differs in having

prominent inflations of the cephalic collar on the

dorsal and ventral surface, a feature which

resembles the lip-like projections present in C.

nephele. C. polymela also differs from C. sappho

in spicule length (2.61—2.71 (2.65) mm in C.

polymela; 1.30—1.50 (1.38) mm in C. sappho) and

in the shape of the vagina which is recurrent in C.

polymela but short and straight in C. sappho.

Cloacina praxithea sp. nov.

(Figs 129-138)

Types: From stomach of Dorcopsis hageni,

Usino, Madang, Papua New Guinea, coll. T.

Reardon, May, 1987. Holotype 6, SAM AHC

31457. Paratypes: 23, SAM AHC 31458, 16,

BMNH 2001.4.10.12; 1d, USNPC 91139. Slide

preparations of apical views of mouth and bursa,

SAM AHC 28388.

Material examined: From Dorcopsis hageni:

types.

Description

Robust nematodes; cervical cuticle slightly

inflated to beyond level of excretory pore;

transverse cuticular annulations widely spaced,

0.015 apart. Submedian cephalic papillae

prominent, 0.019 long, projecting anteriorly from

peri-oral cuticle; distal segment ovoid, 0.008 long,

deviated medially, slightly shorter than

asymmetrical, cylindrical proximal segment, 0.011

long; submedian papillae situated anteriorly on

cephalic collar; amphids situated posteriorly,

below anterior margin of buccal capsule in lateral

views. Buccal capsule shallow, symmetrical in

lateral and dorso-ventral views, wall with faint

striations; anterior margin undulate, with anterior

projections associated with each leaf crown

element. Leaf crown elements 8 in number, not

recurved at tips; peri-oral cuticle inflated into

lobes attached to each element. Dorsal sector of

oesophagus bearing opening of dorsal

oesophageal gland, not projecting into lumen of

buccal capsule. Oesophagus subcylindrical, almost

claviform at base, robust; lining without

sclerotised bosses or denticles; preneural swelling

of oesophagus absent. Nerve ring in mid-

oesophageal region; deirids anterior to nerve ring;

excretory pore at level of, or slightly anterior to,

oesophago-intestinal junction.

Male. Measurements of 6 specimens, types.

Total length 3.71-5.34 (4.86); maximum width

0.32-0.38 (0.36); buccal capsule 0.015—0.018

(0.016) x 0.050—0.055 (0.054); oesophagus 0.66—

0.78 (0.71); nerve ring to anterior end 0.34-0.38

(0.36); excretory pore to anterior end 0.43-0.49

(0.46); deirid to anterior end 0.27-0.32 (0.30);

spicules 1.30-1.58 (1.43); gubernaculum 0.030—

0.040 (0.033) long.

Dorsal ray broad at origin; major bifurcation

occurs at 1/2 length; angle of bifurcation acute;

external branchlets arise immediately after major

bifurcation, much shorter than internals, directed

postero-laterally, not reaching margin of bursa;

internal branchlets directed posteriorly, not

reaching margin of bursa. Externo-dorsal ray not

reaching margin of bursa. Gubernaculum

prominent, subcordate, slightly wider than long.

Spicule tip blunt; ala diminishes gradually in

width then terminates abruptly anterior to spicule

tip; anterior lip of genital cone conical; posterior

24 I. BEVERIDGE

136

FIGURES 129-138. Cloacina praxithea sp. nov., types. 129. Anterior end, lateral view. 130. Cephalic extremity,

lateral view, dorsal aspect on right-hand side. 131. Cephalic extremity, dorsal view; arrows indicate amphids. 132.

Cephalic extremity, apical view; arrows indicate amphids. 133. Cephalic extremity, transverse optical section

through buccal capsule. 134. Submedian cephalic papilla, lateral view. 135. Gubernaculum, ventral view. 136.

Bursa, apical view. 137. Genital cone, dorsal view. 138. Spicule tip, lateral view. Scale bars: 129, 136, 0.1 mm;

130-135, 137, 138, 0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 25

lip with paired projections and cuticular inflation

of internal surface of bursa on either side.

Female. Not seen

Remarks

Cloacina praxithea sp. noy. is distinguished

from all congeners by the position of the amphids

on the cephalic collar. In other species the

amphids are at the same level as the submedian

papillae, anterior to the level of the buccal

capsule, while in C. praxithea the amphids, in

lateral view, are below the level of the anterior

margin of the buccal capsule (Fig. 130). C.

parxithea is characterised by a symmetrical, lobed

buccal capsule without bosses or denticles in the

oesophagus. As such, it has similarities with the

Australian species C. artemis, C. dryope, C. hebe,

C. hypsipyle, C. linstowi, C. maia, C. thetidis and

C. wallabiae, all of which differ from it in having

six rather than eight leaf crown elements. Among

the species known from New Guinea, the features

of the buccal capsule, as well as the presence of

eight leaf crown elements, indicate affinities with

C. syphax, C. solon, C. sappho and C. solymus. C.

solymus has tiny cephalic papillae, in striking

contrast to the prominent papillae of C. praxithea,

while C. sappho and C. polymela have the

amphids on or forming projections beyond the

level of the cephalic collar; C. solon has a greatly

inflated cervical cuticle and its spicules are 2.60—

2.95 (2.81) mm in length compared with 1.93-—

2.05 (1.99) mm in C. praxithea; C. syphax differs

in having the lateral branchlets of the dorsal ray

arise anterior to the principal bifurcation.

Therefore, C. praxithea is readily distinguishable

from all congeners even in the absence of females,

which are currently unknown.

Cloacina procris sp. nov.

(Figs 139-153)

Types: From stomach of Dorcopsis hageni,

Usino, Madang, Papua New Guinea, coll. T.

Reardon, May, 1987. Holotype ¢6, SAM AHC

31459; allotype 2, SAM AHC 31460. Paratypes:

16,22, SAM AHC 31461. Slide preparations of

apical views of mouth and bursa, SAM AHC

28389.

Material examined: From Dorcopsis hageni:

types.

Description

Robust nematodes; cervical cuticle slightly

inflated to level of excretory pore; transverse

cuticular annulations widely spaced, 0.020 apart.

Submedian cephalic papillae small, 0.0035 long,

projecting anteriorly from peri-oral cuticle; distal

segment ovoid, 0.0005 long, much shorter than

cylindrical proximal segment, 0.003 long. Buccal

capsule shallow, asymmetrical in lateral views,

deeper on ventral side, slight anterior arching of

capsule; dorsally, capsule arches anteriorly over

dorsal oesophageal tooth; ventrally, curves

posteriorly; oval and dorso-ventrally elongate in

transverse section, wider than deep, wall with

faint striations; anterior margin smooth. Leaf

crown elements 8 in number, not recurved at tips;

peri-oral cuticle very slightly inflated into lobes

attached to each element. Dorsal sector of

oesophagus bearing opening of dorsal

oesophageal gland, projecting prominently into

lumen of buccal capsule; each sector of

oesophagus bears single triangular tooth-like

structure projecting into lumen. Oesophagus

claviform, robust; lining with sclerotised bosses

extending to level of nerve ring; single dorsal

oesophageal denticle present in preneural region;

preneural swelling of oesophagus absent. Nerve

ring in mid-oesophageal region; deirids posterior

to nerve ring; excretory pore at level of, or slightly

anterior to, oesophago-intestinal junction.

Male. Measurements of 3 specimens, types.

Total length 3.25-4.52 (3.89); maximum width

0.17-0.32 (0.26); buccal capsule 0.015-0.018

(0.017) x 0.050—0.060 (0.054); oesophagus 0.40—

0.41 (0.40); nerve ring to anterior end 0.20-0.21

(0.21); excretory pore to anterior end 0.35—0.40

(0.38); deirid to anterior end 0.27-0.32 (0.30);

spicules 1.30-1.58 (1.43); gubernaculum 0.030

(0.030) long.

Dorsal lobe of bursa elongate; dorsal ray broad

at origin; major bifurcation occurs at 1/2 length;

angle of bifurcation acute; external branchlets

arise at level of major bifurcation, much shorter

than internals, directed almost laterally, not

reaching margin of bursa; internal branchlets

directed posteriorly, reaching margin of bursa.

Externo-dorsal ray not reaching margin of bursa.

Gubernaculum prominent, slightly wider than

long. Spicule tip blunt; ala diminishes gradually

in width then terminates abruptly anterior to

spicule tip; anterior lip of genital cone conical;

posterior lip with paired projections and cuticular

inflations of internal surface of bursa on either

side.

Female. Measurements of 3 specimens, types.

Total length 4.26-5.15 (4.64); maximum width

0.37-0.47 (0.42); buccal capsule 0.018-0.020

(0.019) x 0.060-0.065 (0.062); oesophagus

26 I. BEVERIDGE

FIGURES 139-153. Cloacina procris sp. nov., types. 139. Anterior end, lateral view. 140. Cephalic extremity,

lateral view, dorsal aspect on left-hand side. 141. Submedian cephalic papilla, lateral view. 142. Cephalic extremity,

dorsal view. 143. Cephalic extremity, ventral view. 144. Cephalic extremity, apical view. 145. Cephalic extremity,

transverse optical section through buccal capsule. 146. Transverse optical section through anterior extremity of

oesophagus showing bosses and three tooth-like projections in each of the oesophageal sectors. 147. Preneural

region of oesophagus, dorsal view, showing denticle. 148. Gubernaculum, ventral view. 149. Genital cone, dorsal

view. 150. Bursa, apical view. 151. Spicule tip, lateral view. 152. Female tail, lateral view. 153. Female genital

system, lateral view. Scale bars: 139, 152, 153, 0.1 mm; 140, 142-151, 0.01 mm; 141, 2.5 um.

CLOACINA FROM PAPUA NEW GUINEA 27

0.48-0.50 (0.49); nerve ring to anterior end

0.21-0.22 (0.22); excretory pore to anterior end

0.36—-0.40 (0.38); deirid to anterior end 0.29-

0.33 (0.31); tail 0.19-0.27 (0.23); vulva to

posterior end 0.35—0.47 (0.40); vagina 0.52—0.60

(0.53); egg 0.050—0.060 (0.055) x 0.030—0.035

(0.032).

Female tail slender, conical; vulva immediately

anterior to anus; vagina elongate, straight, turns

abruptly at entry to vestibule; egg ellipsoidal.

Remarks

Although described from only a small series

of specimens, C. procris sp. nov. is quite

distinctive morphologically. It is characterised

by very small submedian cephalic papillae, a

slightly asymmetrical, arched buccal capsule,

eight leaf crown elements, bosses lining the

anterior region of the oesophagus, a single,

small oesophageal denticle, deirid posterior to

the nerve ring, an elongate dorsal ray with the

external branchlets arising immediately after the

principal bifurcation, and a straight vagina.

These features distinguish the species from all

congeners.

The asymmetrical buccal capsule, oesophageal

bosses and dorsal denticle with the posterior

position of the deirid distinguish the species from

all others except C. eos Beveridge, 1998, C.

papillata Beveridge, 1979 and C. sterope. In C.

eos and C. papillata there are only six rather than

eight leaf crown elements and the vagina is

prominently recurrent. In C. sterope, the buccal

capsule exhibits a much greater degree of

asymmetry, the submedian cephalic papillae are

larger and more prominent and the spicules are

longer (1.67—2.07 (1.96) mm in C. sterope, 1.30-

1.58 (1.43) mm in C. procris).

Cloacina sterope Beveridge & Speare, 1999

Material examined: 33, 4°, from stomach of

Dorcopsis hageni, Usino, Madang, Papua New

Guinea, coll. T. Reardon, May, 1987, SAM AHC

31462.

Remarks

Cloacina sterope was described by Beveridge

and Speare (1999) from Dorcopsulus vanhearni

from the Chimbu Province of New Guinea.

Morphological features as well as measurements

of the new specimens from D. hageni agree well

with the original description. D. hageni represents

a new host record.

Cloacina syphax Beveridge & Speare, 1999

Material examined: 176, 92, from stomach of

Dorcopsis hageni, Usino, Madang, Papua New

Guinea, coll. T. Reardon, May, 1987, SAM AHC

31469; BMNH 2001.4.10.13.

Remarks

Cloacina syphax was described from

Dorcopsulus vanheurni by Beveridge and Speare

(1999). The current collection represents a new

host record.

Cloacina theope sp. nov.

(Figs 154-166)

Types: From stomach of Dendrolagus matschiei

Forster & Rothschild, 1907, Huon Peninsula,

Morobe Province, Papua New Guinea, coll. J.

Mayer, July 1999. Holotype ¢, USNPC 91140;

allotype 2, USNPC 91141. Paratypes: 2d, 49,

USNPC 91142; 1d, on slides, SAM AHC 28390.

Material examined: From Dendrolagus

matschiei: types. From Dendrolagus dorianus:

103, 5¢, Lake Trist, Papua New Guinea, coll. I.

Redmond, 1979, BMNH 1981. 4486-4505.

Description

Robust nematodes; cervical cuticle slightly

inflated; transverse cuticular annulations widely

spaced, 0.021—0.044 apart. Submedian cephalic

papillae prominent, 0.017 long, projecting

anteriorly from inflated peri-oral cuticle; distal

segment ovoid, 0.008 long, directed slightly

medially, slightly shorter than robust,

asymmetrical proximal segment, 0.009 long.

Buccal capsule shallow, symmetrical in lateral and

dorso-ventral views; circular in transverse section,

wall without prominent striations; anterior margin

smooth. Leaf crown elements 8 in number,

recurved at tips; peri-oral cuticle not inflated into

lip-like lobes attached to each element. Dorsal

sector of oesophagus bearing opening of dorsal

oesophageal gland, not projecting into lumen of

buccal capsule. Oesophagus claviform, slender;

lining without sclerotised bosses; denticles present

in preneural region; dorsal denticle prominent;

two subventral thickenings of oesophageal lining

giving appearance of denticles, at same level as

dorsal denticle. Nerve ring in mid-oesophageal

region; deirids at level of nerve ring; excretory

pore at level of oesophago-intestinal junction.

Male. Measurements of 4 specimens, types.

Total length 4.68-5.04 (4.88); maximum width

28 I. BEVERIDGE

FIGURES 154-166. Cloacina theope sp. nov., types. 154. Anterior end, lateral view. 155. Cephalic extremity,

lateral view, dorsal aspect on left-hand side. 156. Cephalic extremity, dorsal view. 157. Submedian cephalic papilla,

lateral view. 158. Cephalic extremity, apical view. 159. Cephalic extremity, transverse optical section through

buccal capsule. 160. Preneural region of oesophagus, lateral view, dorsal aspect on left-hand side, showing

denticles. 161. Preneural region of oesophagus, dorsal view, showing denticles. 162. Bursa, apical view. 163.

Gubernaculum, genital cone and spicule sheaths, dorsal view. 164. Spicule tip, lateral view. 165. Female tail, lateral

view. 166. Female genital system, lateral view. Scale bars: 154, 162, 165, 166, 0.1 mm; 155-161, 163-164,

0.01 mm.

CLOACINA FROM PAPUA NEW GUINEA 29

0.30-0.32 (0.31); buccal capsule 0.010 (0.010) x

0.030-0.033 (0.031); oesophagus 0.51-0.60

(0.55); nerve ring to anterior end 0.18-0.20

(0.19); excretory pore to anterior end 0.31-0.41

(0.35); deirid to anterior end 0.13-0.23 (0.18);

spicules 2.29-2.63 (2.52); gubernaculum 0.015—

0.020 (0.018) long.

Dorsal ray broad at origin; major bifurcation

occurs at 1/2 length; angle of bifurcation obtuse;

external branchlets arise immediately after major

bifurcation, shorter than internals, directed

laterally, not reaching margin of bursa; internal

branchlets directed postero-laterally, almost

reaching margin of bursa. Externo-dorsal ray not

reaching margin of bursa. Gubernaculum

prominent, slightly wider than long. Spicule tip

minutely bifid; ala diminishes gradually in width

towards spicule tip; anterior lip of genital cone

conical; posterior lip with paired projections and

cuticular inflations of internal surface of bursa on

either side.

Female. Measurements of 5 specimens, types.

Total length 6.09-6.71 (6.29); maximum width

0.39-0.43 (0.41); buccal capsule 0.008—0.010

(0.009) x 0.030-0.035 (0.032); oesophagus 0.52—

0.64 (0.59); nerve ring to anterior end 0.25 (0.25);

excretory pore to anterior end 0.30—0.51 (0.43);

deirid to anterior end 0.14—0.23 (0.20); tail 0.28—

0.36 (0.33); vulva to posterior end 0.49-0.68

(0.59); vagina 1.03-1.33 (1.19); egg 0.085-0.095

(0.089) x 0.045—0.050 (0.048).

Female tail slender, conical; vulva immediately

anterior to anus; vagina elongate, sinuous, extends

slightly anterior to vestibule with short recurrent

section; egg ellipsoidal.

Remarks

Cloacina theope sp. nov. differs from

congeners in having three oesophageal denticles

in the preneural region of the oesophagus which

are at the same level, but are of dissimilar sizes.

The dorsal denticle is well developed but the

subventral denticles are mere thickenings of the

oesophageal lining and may not warrant the

designation as denticles. In all other species with

three denticles at the same level in the

oesophagus, C. daveyi, C. eurynome, C. hera, C.

hermes, C. hestia, the denticles are equally

developed, although in C. hestia and C. eurynome

all three denticles may be weakly sclerotised. C.

theope differs from C. daveyi, which has

submedian cephalic papillae in which the distal

segment is large and directed medially, and from

all the remaining species, which have the deirid in

the anterior oesophageal position and six rather

than eight leaf crown elements. It is most similar

to C. eurynome from which it differs principally

in having shorter spicules (3.57-3.98 (3.77) mm

in C. eurynome, 2.29-2.63 (2.52) mm in C.

theope).

If the subventral denticles are excluded from

consideration, then the unornamented anterior

oesophagus, a symmetrical buccal capsule with a

smooth anterior margin, and the presence of a

dorsal denticle indicates similarities with C.

cornuta, C. dindymene, C. dirce, C. ernabella, C.

longispiculata and C. sciron. C. ernabella differs

in the shape of the cephalic papillae (globose,

medially directed distal segment in C. ernabella),

while all species except C. sciron differ in having

the deirid in the anterior oesophageal region rather

than at the level of the nerve ring. C. theope

differs from C. sciron, which has an anteriorly

arched buccal capsule, longer spicules (2.83-3.30

(3.00) mm in C. sciron, 2.29-2.63 (2.52) mm in

C. theope), a longer dorsal ray and a vagina which

is not recurrent. In C. theope the recurrent section

of the vagina is twisted around the ascending arm

in a manner similar to that found in C. syphax and

C. eurynome. C. theope differs from C. syphax in

the shape of the buccal capsule, which has an

undulating anterior margin in C. syphax but is

straight in C. theope. Therefore, C. theope is

clearly distinguishable from all congeners.

DIscussION

The new records and new species reported in

this paper suggest that a diverse array of species

of Cloacina exists in macropodid marsupials in

Papua New Guinea. Conclusions need to be

guarded as the current collections have been

obtained from a small number of individuals of

each host species, in some instances from a single

animal. As yet, there have been no comprehensive

surveys of New Guinean macropodids for

parasites, and several species of Thylogale,

Dorcopsis, Dorcopsulus and Dendrolagus have

apparently yet to be examined for helminths

(Spratt et al. 1991). In addition, Beveridge (1998)

and Beveridge and Speare (1999) have listed

museum records of additional undescribed species

from some of these hosts which cannot currently

be named due to lack of adequate material.

In spite of the relatively rudimentary knowledge

of the New Guinean parasite fauna, some

preliminary comparisons can be made with species

present in Australia. In terms of morphological

characters, the species of Cloacina present in New

30 I. BEVERIDGE

Guinea are virtually as diverse as those present in

Australia. Of the various morphological features of

the genus, only the spiral twisting of the proximal,

recurrent region of the vagina (C. syphax, C.

eurynome, C. theope) and the presence of amphids

on conical projections (C. praxithea, C. sappho)

appear to be restricted to species from New Guinea.

Two other apparently autapomorphic characters,

using Arundelia dissimilis (Johnston & Mawson,

1939) as an outgroup, are the posterior position of

the amphids in C. praxithea and the dorsal

deviation of the head in C. polymela. A. dissimilis

is the sole member of a closely related genus within

the tribe Cloacininea (see Beveridge 1987) and is

therefore considered to be an appropriate outgroup

for polarising morphological characters. Characters

such as the origin of the external branchlets of the

dorsal ray anterior to the major bifurcation are more

common in species from New Guinea, but are also

found in species from northern Australia. No

obvious patterns are detectable in the distribution

of other characters. While only tentative

conclusions can be drawn in the absence of a

formal phylogenetic analysis, there do not appear

to be any major distinctions which can be drawn

between species occurring in New Guinea

compared with those present in Australia.

In several instances (C. australis, C. cloelia, C.

cornuta, C. cybele, C. dahli), the same species of

Cloacina are found in Australia and in New

Guinea, a phenomenon explicable in terms of host

distribution. C. australis and C. cornuta are both

parasites of M. agilis, a wallaby which is common

in northern Australia (Strahan 1995) and which

invaded southern New Guinea relatively recently

from Australia (Flannery 1995). Similarly, C.

cloelia, C. cybele and C. dahli occur in pademelons

of the genus Thylogale, and at least one species, T.

stigmatica, is thought to be a recent arrival in New

Guinea (Winter 1997). However, the occurrence of

these same nematodes in species of Thylogale

restricted to New Guinea suggests that they have

been present for a longer period of time than the

recent invasion of T. stigmatica.

The remaining species of Cloacina currently

described from New Guinea occur in hosts which

are endemic. Species of scrub wallabies Dorcopsis

and Dorcopsulus do not occur in Australia, while

the tree kangaroos, Dendrolagus spp., have

apparently radiated in New Guinea and invaded

north-eastern Australia secondarily (Flannery et al.

1996). The recent arrival of the genus

Dendrolagus in Australia may have contributed to

the absence of species of Cloacina in D. lumholtzi

and D. bennettianus.

While the records of species of Cloacina from

New Guinea are still fragmentary, it appears that a

substantial New Guinean fauna exists, and its

relationships with its hosts and the comparisons

that can be made with Australian representatives

may provide insights into the evolution and hence

the diversity of this nematode genus.

KEY TO SPECIES OF CLOACINA IN MACROPODIDS

FROM PAPUA NEW GUINEA

Denticles present in preneural region of

oesophageal lumen... eeeeeeeeeeeeeeee 2

— Denticles absent... eee 13

Paired ventral oesophageal denticles

anterior to dorsal denticle ......... C. dahli

— Single dorsal denticle, or three denticles at

same level in oesophagus .............00 3

Anterior part of oesophagus lined with

SClerotised BOSSES oo... eeeeeeeeeeeteeereeeeens 4

— Anterior oesophagus without sclerotised

DOSSES sire ..- 02 saceste cots sutoacesntcenons dlusecrsurehs 8

Anterior oesophageal bosses much larger

than other bosses; deirid anterior to nerve

ring; female tail swollen ..... C. australis

— Oesophageal bosses of similar size; deirid

at level of nerve ring or posterior to it;

female tail not swollen «0.0... eee 5)

Buccal capsule symmetrical in lateral view

BEST OM tos daa. Se. ALAS eee 2 ee C. erigone

— Buccal capsule asymmetrical in lateral

view, more shallow on dorsal aspect ... 6

Buccal capsule prominently arched

anteriorly 2.8, ents ef, oe C. hyperea

— Buccal capsule not prominently arched

ANLETIOTLY reeiceesk filecseatebiccessteveteettaessls 7

Spicules 1.30-1.58 mm; distal segment of

submedian papilla less than 1/4 length of

proximal segment ................ C. procris

— Spicules 1.67—2.07 mm; distal segment of

submedian papilla only slightly shorter

than proximal segment ......... C. sterope

Dorsal denticle and 2 vestigial subventral

denticles present ...... ce eeeeceeeeeteeeeeeeees 9

— Single dorsal denticle present ............ 10

Spicules 3.57—3.98 mm .....C. eurynome

— Spicules 2.29-2.63 mm .......... C. theope

10. — Prominent lips present ............. C. oweni

== ‘Lips absent: 4. shen ld aanieuace 11

11. — _ Deiridatlevel of nerve ring; anterior margin

CLOACINA FROM PAPUA NEW GUINEA 31

of buccal capsule arched......... C. sciron

— Deirid in anterior oesophageal region,

anterior margin of buccal capsule

NOtzZontale A. ace iba esa 12

12. — Cervical cuticle and female tail inflated,

distal segment of submedian papilla almost

as long as proximal segment, spicules

1.38-1.62 MM... eee C. cornuta

— Cervicalcuticle and female tail not inflated,

distal segment of submedian cephalic

papilla much shorter than proximal

segment, spicules 1.84—2.26 mm............

ee tint de Mawngrdedideigereetchs C. papuensis

13. — Anteriormargin of buccal capsule undulate

ogedeusecersih Mlanadooowadesepagensedcbaregethyivpectesens 14

— Anterior margin of buccal capsule not

MNGUIALC 2.0, astsies AO ME Mae eel 18

14. — Amphids on or forming conical elevations

projecting beyond cephalic collar...... 15

— Amphidsembedded within cephalic collar

aptacabng db guy ke EOE 16

15. — Head deviated dorsally, cephalic collar

elevated between adjacent submedian

papillae 2s nce a cere C. polymela

— Head directed anteriorly, cephalic collar

depressed between adjacent submedian

PAPAS. is sivscsospeedyeetesueideseter C. sappho

16. — External branchlets of dorsal ray arise

before major bifurcation ........ C. syphax

— External branchlets of dorsal ray arise

after major bifurcation ..........c cee 17

17. — Cervical cuticle prominently inflated,

submedian papillae project prominently

beyond cephalic collar .............. C. solon

— Cervical cuticle not inflated, submedian

papillae very small, barely project above

cephalic collar... C. solymus

18. — External branchlets of dorsal ray arise

before major bifurcation ...............0.. 19

— External branchlets of dorsal ray arise

after major bifurcation .........0.ccee 21

REFERENCES

BEVERIDGE, I. 1987. The systematic status of

Australian Strongyloidea (Nematoda). Bulletin du

Muséum National d’Histoire Naturelle, Paris, 4éme

série 9: 107-126.

BEVERIDGE, I. 1998. Taxonomic revision of the genus

Cloacina von Linstow (Nematoda: Strongyloidea)

19.

20.

21.

22.

23.

24.

25.

Deirid anterior to nerve ring ..............006

pabsstbesebadedosecdeaeaenegengyyeied C. cunctabunda

Deirid posterior to nerve ring ............ 20

Spicule length 2.34-2.97 mm.............

Spicule length 1.23-1.45 mm...............

pbabbdeecvdiaretdacsacsssouns tha tgllenay C. caballeroi

Buccal capsule sinuous in apical views,

appearing as extra thickenings of wall in

lateral or dorso-ventral ViewS ...........006.

sendepeh dnesiesnsyasoatemhadcernaced edeetiv diy seieiodarged 22

Buccal capsule prominently arched

AMLETIOTL YE ceil. pereredeteabteceed C. nephele

Buccal capsule not prominently arched .

Pighiethdrgr brig tadessegdepateenescude ry hs itteaesedy 23

Amphids posterior to anterior margin of

buccal capsule 0. C. praxithea

Amphids anterior to anterior margin of

buccal capsule wo... cee eseeeeteeeeeteees 24

Oesophagus with sclerotised bosses,

spicule tip foot-shaped .......... C. cloelia

Oesophagus lacking bosses, spicule tip

SUIMPIC: ssh s.2-,escaedencsecenacuecenrssdicenedevidvass 25

Spicule length 3.90-5.05 mm, spicule tip

SIMPIC ts. 50, cihersseere?Bepugenctetesteers C. hecale

Spicule length 2.30-2.90 mm, spicule tip

with enlarged flange, ala terminating

abruptly anterior to spicule tip ...............

ACKNOWLEDGMENTS

Sincere thanks are due to Tim Flannery, Terry

Reardon, Ian Redmond, Joerg Mayer and Ifor Owen for

collecting the material, sometimes under trying

circumstances, on which this paper is based and to

Eileen Harris and Janet Martin for making the material

available for study. Rosemary Harrigan is thanked for

excellent technical assistance and Dave Spratt for

comments on a draft of the manuscript.

from macropodid marsupials. Invertebrate Zoology

12: 273-508.

BEVERIDGE, I. 1999. New species of Cloacina von

Linstow, 1898 (Nematoda: Strongyloidea) parasitic

in the stomach of the quokka, Setonix brachyurus

(Marsupialia: Macropodidae), from Western

32 I. BEVERIDGE

Australia. Transactions of the Royal Society of South

Australia 123: 17-30.

BEVERIDGE, I. & SPEARE, R. 1999. New species of

parasitic nematodes from Dorcopsulus vanheurni

(Marsupialia: Macropodidae) from Papua New

Guinea. Transactions of the Royal Society of South

Australia 123: 85-100.

BEVERIDGE, I., CHILTON, N. B., JOHNSON, P. M.,

SPEARE, R., SMALES, L. R. & SPRATT, D. M.

1998. Helminth parasite communities of kangaroos

and wallabies (Macropus spp. and Wallabia bicolor)

from north and central Queensland. Australian

Journal of Zoology 46: 473-495.

FLANNERY, T. F. 1995. ‘Mammals of New Guinea’.

Reed Books: New South Wales.

FLANNERY, T. F., BOEADI, & SZALAY, A. L. 1995.

A new tree kangaroo (Dendrolagus: Marsupialia)

from Irian Jaya, Indonesia, with notes on

ethnography and the evolution of tree kangaroos.

Mammalia 59: 65-84.

FLANNERY, T. F., MARTIN, R. & SZALAY, A. L.

1996. ‘Tree kangaroos: a curious natural history’.

Reed Books: Melbourne.

LEWIN, R. A. 1999. ‘Merde. Excursions into scientific,

cultural and sociohistorical coprology’. Aurum Press:

London.

MAWSON, P. M. 1977. Cloacina cornuta (Davey & ~

Wood) and C. caballeroi sp. nov. (Nematoda :

Cloacininae) from macropods from Papua. Excerta

Parasitologica en memoria del Doctor Eduardo

Caballero y Caballero. Instituto de Biologia,

Publicaciones Especiales 4: 455-458.

SPEARE, R., BEVERIDGE, I. & JOHNSON, P. M.

1983. Parasites of the agile wallaby, Macropus agilis

(Marsupialia). Australian Wildlife Research 10: 89-

96.

SPRATT, D. M., BEVERIDGE, I. & WALTER, E. L.

1991. A catalogue of Australasian monotremes and

marsupials and their recorded helminth parasites.

Records of the South Australian Museum,

Monograph Series, 1: 1-105.

STRAHAN, R. 1995. ’The mammals of Australia’.

Reed Books, Chatswood.

WINTER, J. W. 1997. Responses of non-volant

mammals to late Quaternary climatic changes in the

wet tropics region of north-eastern Australia. Wildlife

Research 24: 493-511.

DARWIN FORMATION (EARLY CRETACEOUS, NORTHERN

TERRITORY) MARINE REPTILE REMAINS IN THE SOUTH

AUSTRALIAN MUSEUM

BENJAMIN P. KEAR

Summary

Marine reptile remains from Early Cretaceous (Aptian/Albian) deposits of the Darwin Formation,

Northern Territory, are described from the collections of the South Australian Museum. The

material includes vertebra and limb girdle elements from ichthyosaurs and plesiosauroids, as well as

the first described remains from the Northern Territory. The pliosaurid specimen (a single femur) is

attributed to the small-bodied (up to 3 m in length) freshwater/near-shore marine taxon

Leptocleidus, supporting interpretation of the Darwin Formation as representing a near-shore

marine depositional environment. A comparison of the Darwin Formation marine reptile fauna with

those known from Early Cretaceous deposits elsewhere in Australia, indicates greatest

compositional similarity to the Hauterivian-Barremian Birdrong Sandstone fauna of Western

Australia. A potential link between the distribution of some marine reptile taxa (eg pliosaurids) and

the prevailing depositional environment is also suggested.

DARWIN FORMATION (EARLY CRETACEOUS, NORTHERN TERRITORY) MARINE

REPTILE REMAINS IN THE SOUTH AUSTRALIAN MUSEUM

BENJAMIN P. KEAR

KEAR, B. P. 2002. Darwin Formation (Early Cretaceous, Northern Territory) marine reptile

remains in the South Australian Museum. Records of the South Australian Museum 35(1): 33-47.

Marine reptile remains from Early Cretaceous (Aptian/Albian) deposits of the Darwin

Formation, Northern Territory, are described from the collections of the South Australian

Museum. The material includes vertebra and limb girdle elements from ichthyosaurs and

plesiosauroids, as well as the first described pliosaurid remains from the Northern Territory. The

pliosaurid specimen (a single femur) is attributed to the small-bodied (up to 3m in length)

freshwater/near-shore marine taxon Leptocleidus, supporting interpretation of the Darwin

Formation as representing a near-shore marine depositional environment. A comparison of the

Darwin Formation marine reptile fauna with those known from Early Cretaceous deposits

elsewhere in Australia, indicates greatest compositional similarity to the Hauterivian—Barremian

Birdrong Sandstone fauna of Western Australia. A potential link between the distribution of some

marine reptile taxa (eg pliosaurids) and the prevailing depositional environment is also suggested.

Benjamin P. Kear [kear.ben@saugov.sa.gov.au], *South Australian Museum, North Terrace,

Adelaide, South Australia 5000; and Vertebrate Palaeontology Laboratory, School of

Biological Sciences, University of New South Wales, UNSW Sydney, New South Wales.

2052; *address for correspondence. Revised manuscript received | February 2002.

Sometime in the 1960s (exact date unknown), a

small collection of fossil marine reptile remains

from the Early Cretaceous (Aptian/Albian)

deposits of Casuarina Beach, Darwin Formation,

Northern Territory (see Murray 1987, Fig. 1, p. 95

for locality map) was donated to the South

Australian Museum. In 1994 several additional

Darwin Formation marine reptile specimens were

acquired as part of an exchange with the Museum

and Art Gallery of the Northern Territory. A

recent examination of these remains, which

include isolated elements from both ichthyosaurs

and plesiosauroids, has also revealed the presence

of a pliosaurid femur. This specimen is

attributable to the genus Leptocleidus Andrews,

1922 and represents the first documented pliosaur

material from the Northern Territory. This paper

describes Darwin Formation marine reptile

remains currently housed in the South Australian

Museum and provides a brief palaeoecological

analysis based on faunal comparison with other

known Early Cretaceous marine reptile bearing

deposits.

Marine reptile remains were first reported from

the Darwin Formation by an anonymous author

(1924) who described a fragmentary ichthyosaur

skeleton from Fannie Bay near Darwin. This

specimen was later attributed to Ichthyosaurus

australis (Tiechert & Matheson 1944); however,

Murray (1985) re-assigned the material, along

with other ichthyosaur remains from Nightcliff,

north of Fannie Bay, to Ichthyosauria gen. et sp.

indet. A more recent examination by Wade (1990)

suggested that all ichthyosaur material from the

Darwin Formation might be synonymous with

Platypterygius longmani from the Aptian—Albian

of Queensland. Murray (1987) described isolated

ichthyosaur and elasmosaurid remains from

Casuarina Beach, assigning the material to

Platypterygius sp. and Elasmosauridae gen. et sp.

indet., respectively. The presence of elasmosaurid

and pliosaurid remains in the Casuarina Beach

deposits was also noted by Cruickshank et al.

(1999) but no descriptions were provided.

A comparison of the Darwin Formation marine

reptile fauna with those known from Early

Cretaceous deltaic-marine deposits elsewhere in

Australia (Wallumbilla Formation [Doncaster

Member], Allaru Mudstone, Toolebuc Formation

and Mackunda Formation, Queensland; Bulldog

Shale, South Australia; Wallumbilla Formation

{Doncaster Member], New South Wales; Birdrong

Sandstone and Barrow Group, Western Australia)

indicates greatest compositional similarity to the

Hauterivian—Barremian Birdrong Sandstone

fauna, with ichthyosaurs, elasmosaurids and the

small-bodied pliosaurid Leptocleidus represented.

A potential link between the distribution of some

34 B. P. KEAR

marine reptile taxa (eg pliosaurids) and the

prevailing depositional environment is also

suggested. The presence of Leptocleidus, a

pliosaurid genus known otherwise from

freshwater and near-shore marine deposits

(Andrews 1911, 1922; Strémer 1935; Cruickshank

1997; Cruickshank & Long 1997; Cruickshank et

al. 1999) supports interpretation of the Darwin

Formation as representing a shallow near-shore

marine depositional environment (sensu Smart &

Senior 1980; Murray 1985; Henderson 1998).

MATERIALS AND METHODS

Institutional abbreviations used: SAM, South

Australian Museum, Adelaide; NTM, Museum

and Art Gallery of the Northern Territory, Darwin;

AM, Australian Museum, Sydney.

Lithostratigraphic nomenclature follows Mory

(1988) for Early Cretaceous deposits of the

Darwin area and Burton & Mason (1998) for the

opal-bearing sediments of White Cliffs, New

South Wales. Systematic terminology follows

Montani (1999) for Ichthyopterygia and Brown

(1981) for Plesiosauria except in the subdivision

of Pliosauroidea, in which Polycotylidae is

retained as a separate family (sensu Williston

1925: White 1940; Persson 1963; Welles 1962;

Carpenter 1996, 1997). Functional sub-division of

the vertebral column in ichthyopterygians follows

Buchholtz (2001), with designation of structural

units as neck, trunk, tail stock and fluke.

Interpretation of plesiosaur pelvic and femoral

musculature follows Robinson (1975) and

Lingham-Soliar (2000). All measurements were

taken using callipers and are in millimetres (mm),

GEOLOGICAL SETTING AND PRESERVATION

The marine reptile-bearing deposits of the

Darwin Formation are typically characterised by

glauconitic sandstone and radiolarian mudstone

with basal quartzose conglomerate and localised

nodular phosphorite horizons (Henderson 1998).

The unit forms part of the Bathurst Island Group,

which rests nonconformably on Precambrian

basement within the Money Shoals Platform (see

Fig. 1, p. 117 and Fig. 6, p. 124 of Henderson

1998 for locality and stratigraphic setting). At its

base, the Bathurst Island Group is composed of

the laterally equivalent Darwin Formation and

Marligar Formation, which are overlain in turn by

the Wangarlu Mudstone and Moonkinu

Formation.

The Darwin Formation has historically been

considered Albian in age on the basis of its

macroinvertebrate (Day 1969; Skwarko 1966,

1968; Henderson 1990) and palynomorph

(Morgan 1980) assemblages; however, more

recent analyses (Henderson 1998; D. Megirian

pers. comm. 2000) suggest an upper Aptian age.

The Marligar Formation has been dated as

Neocomian/Aptian by palynological studies

(Burger 1978) and upper Aptian because of its

laterally equivalent relationship with the Darwin

Formation (Henderson 1998). The overlying

Wangarlu Mudstone is regarded as upper Albian—

Cenomanian (possibly with a basal lower Albian

component, Henderson 1998) on the basis of

ammonite (Henderson 1990) and palynological

(Norvick & Burger 1975; Burger 1978) data. The

uppermost Moonkinu Formation is considered

Cenomanian—Turonian on account of its

ammonite (Wright 1963) and bivalve/gastropod/

scaphopod (Skwarko 1983) faunas.

The glauconitic sandstones and predominant

radiolarian mudstones of the Darwin Formation

reflect a near-shore marine/paralic depositional

environment (Smart & Senior 1980; Henderson

1998). The presence of a limited benthic

invertebrate fauna also suggests unfavourable,

possibly anoxic bottom conditions (Henderson

1998).

The Darwin Formation marine reptile remains

generally occur as heavily weathered isolated

elements, although several fragmentary skeletons

have been recovered (Murray 1985; D. Megirian

pers. comm. 2000). The disarticulated nature of

the remains led Murray (1985) to speculate that an

inshore and/or tidal depositional environment may

have facilitated stranding of decaying carcasses

on mudflats prior to burial.

SYSTEMATICS

DIAPSIDA Osborn, 1903

ICHTHYOPTERYGIA Owen, 1860

ICHTHYOSAURIA de Blainville, 1835

OPHTHALMOSAURIA Appleby, 1956 (sensu

Montani 1999)

Platypterygius von Huene, 1922

cf. Platypterygius longmani

DARWIN FORMATION MARINE REPTILE REMAINS

FIGURE 1. SAM P35039, ef. Platpterygius longmani dorsal vertebra in (A) anterior and (B) lateral views. Scale

bar is 40 mm.

Material trunk centra; SAM P35427 (Table 1), three

SAM P35039 (Figs 1A, B; Table 1), four associated anterior tail stock centra; SAM P35429

associated mid-posterior trunk vertebral centra; (Table 1), two associated sections of eight and

SAM P35426 (Table 1), two articulated anterior four articulated anterior? tail stock centra; SAM

FIGURE 2, SAM P35430, cf. Platpterygius longmani caudal vertebra in (A) anterior, (B) lateral, (C) ventral and

(D) dorsal views. Scale bar is 20 mm.

36 B. P. KEAR

TABLE 1. Measurements (mm) of cf. Platypterygius

longmani vertebral centra from the Darwin Formation,

Northern Territory.

Specimen Length Width Height

SAM P35039 39.6 91.2 98.9

33.1 88.9 90.2

30.2 80.1 89.2

32.3 62.6 66.1

SAM P35426 40.6 85.5 88.9

37.0 86.1 89.7

SAM P35427 40.7 94.0 96.4

30.1 86.2 92.4

25.8 102.8 109.6

*SAM P35429 24.9 81.6 69.9

25.8 83.1 75.2

26.4 86.4 79.1

25.2 64.8 83.9

20.5 - 78.2

21.6 58.3 60.1

21.1 58.9 50.8

22.3 - 57.2

17.4 - 55.1

18.9 - 46.6

14.1 - 46.9

SAM P35430 20.8 68.9 59.7

20.8 61.1 62.1

18.7 55.7 56.1

14.6 53.2 50.4

15.3 47.9 48.3

15.3 42.8 42.6

15.2 43.6 45.0

12.9 41.5 41.3

* some centra represented by fragments only

P35430 (Figs 2A, B, C, D; Table 1), nine

associated anterior tail stock centra (juvenile).

Locality

Casuarina Beach, north of Darwin, Northern

Territory. Darwin Formation, Aptian or Albian.

Description

All of the preserved centra are disc-like and

anteroposteriorly compressed, with the dorsal

length subequal to the ventral. Tail stock centra

(Figs 2A, B, C, D) exhibit a more marked degree

of anteroposterior compression relative to those

from the trunk region (Figs 1A, B) and are slightly

ovoid in outline, with height being approximately

1.5 times the transverse width. The articular

surfaces of all centra are deeply amphicoelous,

with a distinct central notochordal pit. The neural

canal is generally broad and shallow, becoming

narrower in centra from the tail stock region. The

neural arch facets are narrow and rectangular in

all centra. Where preserved, both the diapophyses

and parapophyses of the trunk centra are distinctly

raised and subcircular. This is unlike centra from

the tail stock region, in which only the ovoid

diapophysis is present and positioned low on the

lateral centrum surface. The ventral surfaces of at

least two of the recovered centra from the tail

stock region show distinct haemal arch facets.

These are slightly concave and saddle-shaped in

outline with low ridge-like margins. Medially, the

haemal arch facets border the very weakly

concave, spool-shaped depression for the haemal

canal.

Remarks

All of the ichthyosaur vertebral centra examined

can be distinguished from those of basal

ichthyopterygians by their disc-like shape and

strong anteroposterior compression (the more

plesiomorphic Grippidia having cylindrical centra,

Montani 1999). The absence of any further

diagnostic features restricts attribution of the

specimens to Ichthyosauria gen. et sp. indet.

However, tentative assignment of the remains to

cf. Platypterygius longmani is justifiable on the

basis of: 1) overall similarity to P. longmani

vertebral material described from the Allaru

Mudstone and Toolebuc Formation, Queensland

(Wade 1990) and Darwin Formation (Murray

1985, 1987 sensu Wade 1990); and 2) current

recognition of P. longmani as the only valid

ichthyosaur species from the Aptian—Albian of

Australia (Wade 1990; Bardet et al. 1994;

Arkhangelsky 1998; Montani 1999).

Buchholtz (2001) discussed differentiation of

structural units within the vertebral column of

Jurassic ichthyosaurs with relation to variation in

swimming styles between taxa. Following on from

this study, it can be suggested that differences

observed in the Darwin Formation centra (relating

to regionalised morphological variation along the

vertebral column) can be used to predict preferred

swimming style in the Cretaceous P. longmani.

The marked increase in height proportionate to

width, and anteroposterior compression of the tail

stock relative to trunk centra, in the Darwin

Formation specimens is comparable to the

condition recorded in derived thunniform-model

DARWIN FORMATION MARINE REPTILE REMAINS 37

ichthyosaurs such as Ophthalmosaurus icenicus

and Stenopterygius quadrissicus (Buchholtz

2001). Wade (1990) and Broili (1907) noted that

the anterior fluke centra in P. longmani and P.

platydactylus were distinctly higher than wide, a

feature also conforming to the predicted

thunniform condition. Taken together therefore,

these characteristics may suggest that

Platypterygius spp. employed an axial oscillatory

swimming mode and probably shared many other

key traits (see Montani 1999; Buchholtz 2001)

with derived thunniform-model ichthyosaurs.

SAUROPTERYGIA Owen, 1860

PLESIOSAURIA de Blainville, 1835

PLESIOSAUROIDEA (Gray, 1825) Welles, 1943

Plesiosauroidea gen. et sp. indet.

Material

SAM P35431 (Figs 3A, B, C), incomplete and

badly weathered element tentatively identified as

the ventral plate portion of a left ischium.

Orientation is based on position of the lateral

margin and its attitude relative to the horizontal

axis; SAM P35434 (Figs 4A, B, C), right ischium

lacking part of anteromedial margin.

Locality

Casuarina Beach, north of Darwin, Northern

Territory. Darwin Formation, Aptian or Albian.

Description

Dimensions of remains are: SAM P35431, total

length of 96.7 mm; total width is 98.9 mm;

maximum height is lateral extremity is 22.5 mm;

maximum length of lateral extremity is 55.3 mm;

maximum height of ventral plate is 45.9 mm.

SAM P35434, total length is 74.6 mm; total width

FIGURE 3. SAM P35431, Plesiosauroidea gen. et sp. indet. partial ischium in (A) dorsal, (B) medial and (C)

posterior views. Scale bar is 40 mm.

38 B. P. KEAR

FIGURE 4. SAM P35434, Plesiosauroidea gen. et sp. indet. ischium in (A) dorsal, (B) posterior and (C) lateral

views. Scale bar is 40 mm.

is 97.4mm; maximum height of the of lateral

articular surface is 52.1 mm; maximum length of

lateral articular surface is 46.1 mm; maximum

height of ventral plate is 10.8 mm. Both SAM

P35431 and SAM P35434 are hatchet-shaped in

dorsal outline with an anteroposteriorly expanded

ventral plate. The lateral articular extremity of

SAM P35434 is massive but has been lost in SAM

P35431. Medially, the ventral plate portion of

both elements is dorsoventrally compressed with

an inclined medial edge for contact with the

opposing ischium. This surface is oriented

approximately 10° ventral to a horizontal plane

through the lateral articular extremity, and is

deeply pitted, suggesting the presence of cartilage.

Both the dorsal and ventral surfaces of the ventral

plate in SAM P35434 are rugose, possibly for

attachment of the m. puboischiofemoralis internus

and m. p. externus, respectively. A weak ridge on

the posteromedial margin of the ventral plate may

also have supported slips of the m.

ischiotrochantericus. The ventral plate of both

specimens is not greatly anteroposteriorly

expanded (with total length being just slightly less

than the total transverse width of each element)

and is separated from the lateral articular

extremity by a constricted waist-like midsection.

This is ovoid to circular in cross-section,

becoming increasingly dorsoventrally thickened

towards the lateral articular surface (not preserved

in SAM P35431). The articular surface itself is

convex, with the rectangular anterior pubis facet

DARWIN FORMATION MARINE REPTILE REMAINS 39

offset approximately 195° relative to the

longitudinal axis of the articular surface. The

pubis facet is separated posteriorly from the

rectangular median acetabular facet, and in turn

from the lobate posterior-most ilial facet (offset

approximately 150° relative to the longitudinal

axis of the articular surface) by weak parallel

ridges. The articular surface margin is produced

into a crennate rim, which is continuous ventrally,

with a low rugose tuberosity possibly marking part

of the m. puboischiofemoralis internus

attachment.

Remarks

Despite being heavily weathered, SAM

P35431 is identified as the ventral plate portion

of a plesiosaurian left ischium on the basis of its

hatchet-shaped outline and general morphology.

Nevertheless, the specimen is unusual in its

markedly dorsoventrally deep, medial articular

margin. SAM P35431 was initially registered as

an ichthyosaurian humerus belonging to

Platypterygius sp. However, such an

identification is unlikely as the humeri of

Platypterygius spp. are much more robust, with

rectangular to spool-shaped dorsal outline and

prominent ridge-like tuberosities on both the

dorsal and ventral surfaces (eg Kiprijanoff 1881;

Broili 1907; Kuhn 1946; McGowan 1972; Wade

1984; Murray 1987; Choo 1999). SAM P35431

also differs from the larger girdle elements of

Platypterygius spp., in which the coracoid is

subcircular in outline with dorsoventrally deep

glenoid/intercoracoid facets (Broili 1907; Wade

1984), and both the scapula and ischiopubis are

strap-like with predominantly narrow, ridge-like

articular surfaces (Broili 1907; Nace 1939;

Murray 1987).

SAM P35434 probably represents a juvenile

individual because of its small size and poor

ossification of the articular surfaces. Both it and

SAM P35431 can be assigned to Plesiosauroidea

on the basis of their anteroposteriorly short ventral

plates. This differs from the condition in

pliosauroids in which the ventral plate of the

ischium bears a marked posterior expansion (Mehl

1912; Andrews 1913; Tarlo 1959, 1960; Brown

1981). The presence of a relatively weak posterior

expansion in Eurycleidus (Cruickshank 1994) and

Leptocleidus (pers. obs, AM F99374 from the

Aptian—Albian Bulldog Shale of Coober Pedy,

South Australia), however, suggests that this

character state may not be universal for the group.

SAM P35434 is unusual in its proportionately

massive articular head relative to the ventral plate.

The phylogenetic significance of this feature, if

any, is uncertain because of its immature stage of

development.

ELASMOSAURIDAE Cope, 1869

Elasmosauridae gen. et sp. indet.

Material

SAM P35432 (Figs SA, B, C, D), a single

weathered posterior cervical vertebral centrum.

Locality

Casuarina Beach, north of Darwin, Northern

Territory. Darwin Formation, Aptian or Albian.

Description

The surface of the centrum is badly weathered

with very little periosteal bone remaining. Total

length is 72.8mm; width of anterior surface is

109.5 mm; depth of anterior surface is 85.2 mm.

In general proportions, the centrum is

anteroposteriorly short relative to both the height

and width of the anterior articular surface. Both

the lateral and ventral surfaces are shallowly

concave, with the ventral surface being pierced by

two large foramina. The articular surfaces are

poorly preserved but clearly platycoelous, with a

raised convex rim. The neural arch facets are

indistinct but prominent bosses high on the lateral

surface of the centrum indicate positioning of the

rib facets.

Remarks

SAM P35432 is tentatively assigned to

Elasmosauridae on the basis of its platycoelous

articular surfaces. Murray (1987) reported an

elasmosaurid cervical with very shallowly concave

articular facets from the Casuarina Beach locality.

Brown (1981, 1993) and Bardet et al. (1999)

regarded the presence of platycoelous articular

surfaces on the cervical centra to be a potential

synapomorphy for Elasmosauridae. The character

state has also been recorded in the Maastrichtian

cryptoclidid Morturneria (Chattergee & Small

1989), though Bardet et al. (1991) suggested that

this taxon may represent a derived elasmosaurid.

The proportionately short centrum length

relative to height of SAM P35432 differs from the

common condition in elasmosaurids (such as that

described by Murray 1987), in which the cervical

centra are markedly elongate (Brown 1993; Bardet

et al. 1999). The presence of relatively short

cervical centra is considered plesiomorphic among

40 B. P. KEAR

FIGURE 5. SAM P35432, Elasmosauridae gen. et sp. indet. posterior cervical centrum in (A) dorsal, (B) ventral,

plesiosauroids (Brown 1993; Bardet ef al. 1999).

However, the feature has been described in the

potential elasmosaurid taxa (sensu Bardet et al.

1991) Aristonectes (Cabrera 1941) and

Morturneria (Chattergee & Small 1989) as well

as in indeterminate elasmosaurid posterior

cervicals from unspecified (probably Albian)

deposits near Oodnadatta (Freytag 1964) and the

Neales River region (SAM P6181, Persson 1960),

South Australia; Toolebuc Formation, Queensland

(Kear 2001); and Molecap Greensand, Western

Australia (Long & Cruickshank 1998). The

presence of proportionately elongate cervicals

may therefore not necessarily be universal

amongst elasmosaurids, with some taxa exhibiting

relatively short centra, particularly in the posterior

cervical region.

SAM P35432 is notable for its possession of a

raised convex rim surrounding the central

concavity of the articular surface. Brown et al.

(1986) and Brown (1993) considered this a

plesiomorphic condition commonly associated

with members of the Cryptoclididae and

Plesiosauridae. In contrast, elasmosaurids,

including basal taxa such as Occitanosaurus

(Bardet et al. 1999) and Muraenosaurus (Andrews

1910; Brown 1981, 1993), generally exhibit a

more open V-shape, in which the articular surface

margins form an abrupt angle with the centrum

sides. A similar trait is evident in elasmosaurid

material from the Toolebuc Formation of

Queensland (Kear 2001), and is indicated (but not

described) in the figured Darwin Formation

specimen (NTM P8727-70, Fig. 1, p. 97) of

Murray (1987). Carpenter (1996), however,

considered articular rim morphology to be a

variable character amongst plesiosaurs and of

questionable diagnostic value.

DARWIN FORMATION MARINE REPTILE REMAINS 41

PLIOSAUROIDEA Seeley, 1874

PLIOSAURIDAE Seeley, 1874

Leptocleidus Andrews, 1922

Leptocleidus sp.

Material

SAM P35053, cast of NTM P913-5 (Figs 6A,

B, C) right femur.

Locality

Casuarina Beach, north of Darwin, Northern

Territory. Darwin Formation, Aptian or Albian.

Description

A near complete right femur lacking part of

posterodistal margin. Total length is 260.8 mm;

FIGURE 6. SAM P35053 (cast of NTM P913-5),

Leptocleidus sp. femur in (A) anterior, (B) dorsal and

total proximal length is 47.9 mm; total proximal

height (including trochanter) is 51.5 mm; width

across base of dorsal trochanter is 37.2 mm; total

distal length (not including missing posterior

margin) is 91.6 mm; total distal height is

25.9 mm. The anterior edge of the femur is weakly

sigmoidal, with a strongly convex distal extremity.

The posterior edge is uniformly concave along its

entire length. Both the dorsal and ventral surfaces

are shallowly concave along their longitudinal

axis; however, the ventral surface becomes

convex and upturned towards the distal extremity.

Proximally, the femur is greatly dorsoventrally

expanded and separated from its distal section by

a constricted neck. The dorsal trochanter is robust

and anteroposteriorly constricted at its base. It is

separated from the ovoid glenoid articular surface

by a shallow groove. The surfaces of both the

dorsal trochanter and glenoid articulation are

deeply pitted, suggesting the presence of cartilage.

The femoral shaft is weakly anteroposteriorly

constricted and ovoid in cross-section, with its

posterior margin bearing a raised rugose

tuberosity, possibly for attachment of the m.

caudifemoratis. A second low tuberosity is present

on the median ventral surface and may represent

part of the m. puboischiofemoralis insertion. The

distal femoral extremity is fan-shaped in

dorsoventral outline, with a weakly projecting

anterodistal margin. The distal articular surface is

narrow and elliptical, with a raised median ridge

separating the very large radial facet from the

remains of the ulnar facet.

Remarks

Despite the poor existing knowledge of femoral

morphology in Leptocleidus spp., SAM P35053

(NTM P913-5) can be readily distinguished from

the proportionately shorter and stouter propodials

of elasmosaurids (Brown 1981; Murray 1987) by

its anteroposteriorly broad, flat femoral shaft,

greatly expanded distal extremity and sigmoidal

anterior margin. These characteristics are very

similar to those of the fragmentary femora of L.

capensis (Andrews 1911) and well-preserved

material (AM F99374, SAM P15980) from the

Bulldog Shale of South Australia (Figs 7A, B).

The femur of L. clemai (Cruickshank & Long

1997) differs in its possession of a concave

anterior margin. This feature, along with a

proportionately more slender shaft, is also present

in the femora of polycotylids and most other

pliosauroids (Andrews 1913; Welles 1943, 1962;

Brown 1981; Storrs 1999). A sigmoidal anterior

femoral and humeral margin has, however, been

42 B. P. KEAR

TABLE 2. Marine reptile groups from Early Cretaceous deltaic-marine deposits of Australia. Pliosauroids are

separated into small-bodied (eg Leptocleidus) and large-bodied (eg Kronosaurus) forms because of their differing

ecological roles. The poorly known taxon ‘Cimoliasaurus maccoyi’ is recorded separately because of its unknown

affinity. Indeterminate plesiosaur material from the marine Bungil Formation, Queensland and freshwater deposits

of the Griman Creek Formation, New South Wales, and Strzeleki and Otway Groups, Victoria is excluded pending

a more thorough examination of the remains. Taxa from the opal bearing deposits of White Cliffs, New South

Wales are placed within the Wallumbilla Formation in accordance with the lithostratigraphic nomenclature of

Burton & Mason (1998). Source texts include: Etheridge (1904)', Longman (1924), Tierchert & Matheson (1944)3,

Persson (19604, 19635, 1982°), Ludbrook (1966)’, Condon (1968)’, Pledge (1980), Smart & Senior (1980)!,

Gaffney (1981)"', Molnar (1982!7, 199119), Wade (1984"4, 1990!5), Murray (1985!°, 19877), Hocking et al. (1987)!8,

Burger (1988)!°, Bardet (1992), Thulborn & Turner (1993)?!, Krieg & Rodgers (1995), Cruickshank & Long

(1997)3, Henderson (1998)4, Long (1998), Long & Cruickshank (1998), Cruickshank et al. (1999)?’, Choo

(1999)*8, Kear (2001).

Darwin Formation Wallumbilla Formation

(Money Shoals Platform)

Aptian/Albian

(Eromanga Basin)

Aptian—Albian

Ichthyosauria*!3!5.16.17 Ichthyosauria!3

Elasmosauridae™3!” Elasmosauridae**!2:!325

Small pliosaurid”’ Large pliosaurid !245:12.1321.25

Polycotylidae**3

*C. maccoyi’'43

Shallow near-shore Coastal / offshore shallow

marine™ / paralic!? / marine!®!9

possibly tidal'®

cited as a potentially diagnostic character state for

the polycotylid genus Polycotylus (Storrs 1999).

DIscussION

The Darwin Formation sediments have been

interpreted as representing a shallow near-shore

marine/paralic (Smart & Senior 1980; Henderson

1998) to possibly tidal (Murray 1985) depositional

environment. The presence of the small pliosaurid

taxon Leptocleidus, known elsewhere from

freshwater and near-shore marine deposits

(Andrews 1911, 1922; Str6mer 1935; Cruickshank

1997; Cruickshank & Long 1997; Cruickshank et

al. 1999), is consistent with this interpretation.

A comparison (Table 2) of the Darwin

Formation marine reptile fauna with those known

from Early Cretaceous deltaic-marine deposits

elsewhere in Australia (Wallumbilla Formation

[Doncaster Member], Allaru Mudstone, Toolebuc

Formation and Mackunda Formation, Queensland;

Bulldog Shale, South Australia; Wallumbilla

Formation [Doncaster Member], New South

Wales; Birdrong Sandstone and Barrow Group,

Allaru Mudstone

(Eromanga Basin)

Albian

Toolebuc Formation

(Eromanga Basin)

Albian

Ichthyosauria!*:!3-14.15.25

Elasmosauridae**!?:13.25

Chelonioidea’3*5

Ichthyosauria!2!314.15.25

Elasmo sauridae*>®! 2,13,21,25,29

Large pliosaurid?*5!2.3

Polycotylidae?!?527

Chelonioidea!!:!2.13.25

Shallow marine!” Shallow marine’®

Western Australia) indicates greatest

compositional similarity to the Hauterivian—

Barremian Birdrong Sandstone fauna, with

ichthyosaurs, elasmosaurids and the small-bodied

pliosaurid Leptocleidus represented. Throughout

the Early Cretaceous marine deposits,

ichthyosaurs show a wide distribution (occurring

in all currently recognised deposits except the

Albian Mackunda Formation and unspecified

Berriasian subsurface sediments of the Barrow

Group), as do elasmosaurids, which have been

recorded from all localities except those of the

Albian Mackunda Formation and Berriasian

Barrow Group.

Cruickshank et al. (1999) indicated the possible

presence of cryptoclidids in the Aptian—Albian

Bulldog Shale. This is supported by more recent

observations of the South Australian fauna (Kear

unpubl.) but as yet the group has not been

recorded from any other Australian deposit.

The distribution of small- and large-bodied

pliosaurid taxa in the Australian Early Cretaceous

is variable, with smaller forms (represented by at

least two species of Leptocleidus, Cruickshank et

al. 1999) confined to the Darwin Formation,

TABLE 2. (cont.)

Mackunda Formation

(Eromanga Basin)

Albian

Polycotylidae’

DARWIN FORMATION MARINE REPTILE REMAINS

Bulldog Shale

(Eromanga Basin)

Aptian—A Ibian

Birdrong Sandstone

(Carnarvon Basin)

Hauterivian—Barremian

Ichthyosauria®'3° Ichthyosauria*!378

Barrow Group

(Carnarvon Basin)

Berriasian

Small pliosaurid?¢

Elasmosauridae®”’

Small pliosaurid®?5?7

Large pliosaurid’

Cryptoclididae?”’

Paralic’® Shallow marine? /

offshore’

Bulldog Shale and Birdrong Sandstone. Larger

pliosaurid remains occur only in the Aptian—

Albian Wallumbilla Formation (Doncaster

Member), Albian Toolebuc Formation and

Bulldog Shale deposits. This division may be

linked to the different habitat preferences and

ecological roles of small- and large-bodied

pliosaurid taxa, with offshore environments

preferentially supporting larger-bodied forms. An

exception is seen in the shallow offshore marine

(Ludbrook 1966; Krieg & Rogers 1995) Bulldog

Shale, which includes both small- and large-

bodied pliosaurid remains (the latter being

represented by an isolated tooth, SAM P22525,

from the opal-bearing sediments of Coober Pedy,

figured by Pledge 1980, p. 8).

Distribution of polycotylids in the shallow

marine (Smart & Senior 1980; Burger 1988)

Wallumbilla Formation, Toolebuc Formation and

paralic (Smart & Senior 1980) Mackunda

Formation may, as with large-bodied pliosaurids,

be related to a preference for predominantly

offshore marine environments. However,

individuals also appear to have readily

Elasmosauridae”©??

Small pliosaurid?3??

Coastal near-shore Fluviatile-deltaic'®

shallow marine!®/

paralic®

frequented more inshore habitats, as suggested

by their occurrence in the Mackunda Formation

deposits.

Restriction of chelonioids to the upper Albian

Allaru Mudstone and Toolebuc Formation may be

the product of both the group’s temporal

distribution (the earliest-known chelonioid being

recorded from the upper Aptian — lower Albian

Santana Formation of Brazil, Hirayama 1997,

1998) as well as environmental factors such as

prevailing water temperatures. Studies of

sedimentary sequences (Frakes & Francis 1988;

Frakes & Krassay 1992; Frakes et al. 1995;

Constantine et al. 1998), climatic modelling

(Barron & Washington 1982) and isotope data

(Gregory et al. 1989; Pirrie et al. 1995) have

suggested that strongly seasonal climates with

winter freezing and at least seasonal sea ice

characterised high latitudes in the Early

Cretaceous of Australia. Such conditions may well

have been unfavourable to chelonioids and

provided an effective barrier to the group’s

dispersal into the Australian region prior to the

upper Albian.

44 B. P. KEAR

FIGURE 7. SAM P15980 juvenile Leptocleidus sp. humerus (A) and femur (B) from Andamooka, South Australia.

Scale bar is 20 mm.

ACKNOWLEDGMENTS

I wish to thank Dirk Megirian of the Museum and

Art Gallery of the Northern Territory for assistance

with locating literature on the Darwin Formation

deposits, and generous provision of NTM P913-S for

study. Tim Flannery, Neville Pledge, Jo Bain and Ben

Head of the South Australian Museum provided

support, enthusiasm, preparation facilities and

materials. This manuscript benefited greatly from the

comments of Dirk Megirian and Arthur Cruickshank

of the Leicester City Museums Service, New Walk

Museum. Origin Energy, The Advertiser, the

Waterhouse Club and the Coober Pedy Tourism

Association provided financial support for this

research,

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TENTACULARIID CESTODES OF THE ORDER TRYPANORHNCHA

(PLATYHELMINTHES) FROM THE AUSTRALIAN REGION

H. W. PALM & 1. BEVERIDGE

Summary

The present study summarises information on tentaculariid trypanorhynchs from Australian waters.

A total of 19 species from the genera Nybelina Poche, 1926, Heteronybelinia Palm, 1999,

Mixonybelinia Palm, 1999, Kotorella Euzet & Radujkovic, 1989 and Kotorelliella gen. nov. were

identified: N. aequidentata (Shipley & Hornell, 1906); Nybelina africana Dollfus, 1960; N.

hemipristis sp. nov.; N. jayapaulazariahi Reimer, 1980; N. mehlhorni sp. nov.; N. schmidti Palm,

1999; N. scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996); N. strongyla Dollfus,

1960; N. thyrsites Korotaeva, 1971; N. victoriae sp. nov.; Heteronybelinia australis sp. nov.; H.

estigmena (Dollfus, 1960); H. pseudorobusta sp. nov.; Mixonybelinia beveridgei (palm, Walter,

Schwerdtfeger & Reimer, 1997); M. cribbi sp. nov.; M. edwinlintoni (Dollfus, 1960); M. southwelli

Palm & Walter, 1999; Kotorella pronosoma (Stossich, 1901) and Kotorelliella jonesi gen. et sp.

nov. The new genus Kotorelliella is characterised by a homeocanthous, heteromorphous metabasal

armature and a basal armature with additional interpolated hooks on the external surface of the

tentacle, thus appearing heterocanthous. The new species appears to be closely related to Kotorella

pronosoma. The tentaculariid trypanorhynch fauna in Australian waters is pecies rich, with 22

(48%) of the total of 46 known species occurring in waters of the region. Eleven new locality, and

20 new host records are reported.

TENTACULARIID CESTODES OF THE ORDER TRYPANORHYNCHA

(PLATYHELMINTHES) FROM THE AUSTRALIAN REGION

H. W. PALM & I. BEVERIDGE

PALM, H. W. & BEVERIDGE, I. 2002. Tentaculariid cestodes of the order Trypanorhyncha

(Platyhelminthes) from the Australian region. Records of the South Australian Museum 35(1):

49-78.

The present study summarises information on tentaculariid trypanorhynchs from Australian

waters. A total of 19 species from the genera Nybelinia Poche, 1926, Heteronybelinia Palm,

1999, Mixonybelinia Palm, 1999, Kotorella Euzet & Radujkovic, 1989 and Kotorelliella gen.

nov. were identified: N. aequidentata (Shipley & Hornell, 1906); Nybelinia africana Dollfus,

1960; N. hemipristis sp. nov.; N. jayapaulazariahi Reimer, 1980; N. mehlhorni sp. nov.; N.

schmidti Palm, 1999; N. scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996); N.

strongyla Dollfus, 1960; N. thyrsites Korotaeva, 1971; N. victoriae sp. nov.; Heteronybelinia

australis sp. nov.; H. estigmena (Dollfus, 1960); H. pseudorobusta sp. nov.; Mixonybelinia

beveridgei (Palm, Walter, Schwerdtfeger & Reimer, 1997); M. cribbi sp. nov.; M. edwinlintoni

(Dollfus, 1960); M. southwelli Palm & Walter, 1999; Kotorella pronosoma (Stossich, 1901)

and Kotorelliella jonesi gen. et sp. nov. The new genus Kotorelliella is characterised by a

homeoacanthous, heteromorphous metabasal armature and a basal armature with additional

interpolated hooks on the external surface of the tentacle, thus appearing heteroacanthous, The

new species appears to be closely related to Kotorella pronosoma. The tentaculariid

trypanorhynch fauna in Australian waters is species rich, with 22 (48%) of the total of 46

known species occurring in waters of the region. Eleven new locality, and 20 new host records

are reported.

H, W. Palm, Institut fiir Zoomorphologie, Zellbiologie und Parasitologie, Heinrich-Heine-

Universitat Diisseldorf, UniversitatsstraBe 1, D-40225 Diisseldorf, Germany, current address:

Centre for Coastal and Marine Resource Studies, Faculty of Fisheries and Marine Sciences,

Bogor Agricultural University, 16680 Bogor, Indonesia; email: hpalm@indo.net.id,

hpalm@ gmx.net; and I. Beveridge, Department of Veterinary Science, University of Melbourne,

Parkville, Victoria 3052, Australia. Manuscript received 20 November 2000.

Palm (1999), Palm et al. (1997), Palm and

Overstreet (2000) and Palm and Walter (1999,

2000) partially revised the trypanorhynch cestode

family Tentaculariidae Poche, 1926 on the basis

of material deposited in museums in London,

Paris, Vienna and the United States. Following

the erection of two new genera, Heteronybelinia

Palm, 1999 and Mixonybelinia Palm, 1999 by

Palm (1999), Palm and Walter (2000) gave a

summary of the current state of knowledge within

the tentaculariid trypanorhynch genera. Thus, to

date, a total of 39 species of Kotorella, Nybelinia,

Heteronybelinia and Mixonybelinia are considered

valid (Palm & Walter 2000).

There is littke known about the tentaculariid

trypanorhynchs from Australian coastal waters.

The first collections were made by a French

scientific expedition under the command of

Nicolas Baudin between 1801 and 1803

(Beveridge & Campbell 1996), but the cestodes,

Tentacularia coryphaenae Bosc, 1797 and an

unidentified species of Nybelinia were not

described until 1942 (Dollfus 1942). Korotaeva

(1971) named the Nybelinia species of Dollfus

(1942) N. thyrsites and subsequently (Korotaeva

1974) reported additional unidentified Nybelinia

spp. from other fishes from southern Australia.

Lester et al. (1988) and Sewell and Lester (1988)

reported Tentacularia sp. from orange roughy,

Hoplostethus atlanticus Collett, 1889 off the

coasts of South Australia and Tasmania; while

Sewell and Lester (1995) found Nybelinia sp. in

jewfish, Rexea solandri (Cuvier & Valenciennes,

1832) from southern Australian waters. Beveridge

and Campbell (1996) recorded Tentacularia

coryphaenae Bosc, 1797, Nybelinia thyrsites

Korotaeva, 1971 and N. sphyrnae Yamaguti, 1952

from Australian fishes; while Jones and Beveridge

(1998) added N. queenslandensis from a

carcharhinid shark as a new species. Palm (1999)

recorded Nybelinia lingualis (Cuvier, 1817) from

southern Australian waters and Speare (1999)

50 H. W. PALM & I. BEVERIDGE

reported Nybelinia sp. from sailfish, [stiophorus

platypterus (Shaw & Nodder, 1792) off the coast

of Queensland. Thus, a total of five tentaculariid

species has been described from Australia.

Last and Stevens (1994) summarised

knowledge of the Australian chondrichthyan

fauna, the definitive hosts of trypanorhynchan

cestodes, as being extremely rich and consisting

of 296 species, 54% of them endemic. In terms of

host-specificity and zoogeography' of

trypanorhynch cestodes, the Australian fauna

might therefore harbour a high number of

endemic and locally distributed tentaculariid

trypanorhynchs, such as was observed within the

related family Eutetrarhynchidae by Beveridge

(1990). Palm (1999) and Palm and Walter (2000)

suggested that many tentaculariids, for example

Tentacularia coryphaenae and Nybelinia

lingualis, exhibit a cosmopolitan or circumtropical

distribution pattern. However, records from the

south-west Pacific and south-east Indian Ocean

are scarce. For example, the tentaculariids

Tentacularia coryphaenae, Nybelinia africana

and N. scoliodoni were only recently recorded

from Indonesian coastal waters (Palm 2000).

Other records which might confirm a wide

distribution and possibly a low degree of host

specificity for these trypanorhynchs are still

lacking.

The present study was carried out to examine

the species of Nybelinia, Heteronybelina,

Mixonybelinia and Kotorella deposited in

museum collections in Australia as well as in the

collection of the junior author. Where necessary,

the specimens are described and illustrated as

emendations to currently available descriptions.

Apart from the establishment of new host and

locality records, species identifications provide

further insight into the zoogeographical

distribution of tentaculariid cestodes. The

comparison of the specimens with earlier

descriptions allows comments to be made on the

extent of intraspecific morphological variability

within tentaculariid trypanorhynchs. Together

with the study of deposited tentaculariids from

other collections (Palm 1999; Palm & Overstreet

2000; Palm and Walter 1999, 2000), the present

study summarises the current state of knowledge

on the genera of tentaculariid cestodes.

MATERIAL AND METHODS

Specimens were examined from the Australian

Helminthological Collection (AHC) housed in the

South Australian Museum (SAM), Adelaide, as

well as the Queensland Museum (QM), Brisbane.

Special attention was given to unidentified

specimens deposited simply as Nybelinia sp.

Unmounted specimens were stained in Celestine

Blue, dehydrated in ethanol, cleared in methyl

salicylate and mounted in Canada balsam. Pieces

of strobilae were embedded in paraffin, serial

sections were cut at a thickness of 5S fim and

stained with haematoxylin and eosin. Drawings

were made using an Olympus BH2 microscope

with an ocular micrometer and drawing tube.

Terminology for morphological features peculiar

to the Trypanorhyncha follows Dollfus (1942) and

Campbell and Beveridge (1994).

The following measurements were made: scolex

length (SL), scolex width at level of pars

bothridialis (SW), pars bothridialis (pbo), pars

vaginalis (pv), pars bulbosa (pb), pars postbulbosa

(ppb), velum (vel), appendix (app), bulb length

(BL), bulb width (BW), bulb ratio (BR), scolex

proportions of pbo:pv:pb (SP), tentacle width

(TW) and tentacle sheath width (TSW). If

possible, the tentacle length (TL) was estimated.

The tentacular armature was described as follows:

armature homeomorphous or heteromorphous,

number of hooks per half spiral row (hsr), total

hook length (L) and the total length of the base of

the hooks (B). The abbreviation ‘nm’ (not

measured) indicates that no measurement was

taken. All measurements are given in micrometers

unless otherwise indicated. Specimens belonging

to the same species from different hosts or

localities were measured in the same order as cited

in the material examined. If more than three

measurements were taken from a single host

species, the mean is given with the range in

parentheses. Because tentaculariid cestodes are

morphologically uniform and the genera are well

defined, the species descriptions presented here

are restricted to key differential features.

Illustrations are provided if useful for future

species identification; otherwise the reader is

referred to illustrations given by other authors.

The classification utilised follows Palm (1995,

1997). Host identity follows Last and Stevens

(1994) for the elasmobranchs and FishBase 1998

(Froese & Pauly 1998) for the teleosts.

SYSTEMATICS

A total of 19 species were identified, and 11

new locality and 20 new host records were

established. Seven new species are described and

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 51

a new tentaculariid genus is erected: Nybelinia

hemipristis sp. nov., N. mehlhorni sp. nov., N.

victoriae sp. nov., Heteronybelinia australis sp.

nov., H. pseudorobusta sp. nov., Mixonybelinia

cribbi sp. nov. and Kotorelliella jonesi gen. et sp.

nov. Detailed information on individual

specimens with comments on their taxonomy and

distribution are provided below.

Order TRYPANORHYNCHA Diesing, 1863

Superfamily TENTACULARIOIDEA Poche,

1926

Family TENTACULARIIDAE Poche, 1926

Genus Nybelinia Poche, 1926

Nybelinia aequidentata Shipley & Hornell, 1906

(Figs 1-2)

Material examined

From Dendrochirus zebra (Cuvier, 1829): 1

postlarva, Noumea, New Caledonia, Nov. 1997,

coll. S. Pichelin (QM G 218031).

Description

SL=7900 (Fig. 1); SW =1775; pbo = 2975;

pv = 4950; pb = 1575; vel = 925; app = 1400;

BL = 1417 (1400-1450); BW = 280 (270-290);

BR =5.1:1; SP = 1.9:3.1:1. Basal tentacular

swelling absent. TW basal = 100-110, TW

metabasal = 110-120. Tentacle sheaths straight,

TSW = 80-100, prebulbar organs and muscular

rings around basal part of tentacle sheaths not

visible. Retractor muscles originate in basal parts

of bulbs. Tentacular armature homeoacanthous,

homeomorphous (Fig. 2); distinctive basal

armature absent. Hooks falciform, with slender

shaft, stout base and strongly recurved tip,

L = 57.5-65.0; B = 17.7—22.5. Hooks decrease in

size towards apical region and towards base

L = 35.0-57.5; B = 12.5-17.5. Smallest hook

measured L = 17.5; hsr = 7-8.

Remarks

The present specimen has one of the largest

scoleces among Nybelinia species. In addition, the

postlarva is characterised by falciform hooks over

60 in length and a bulb ratio greater than 4. The

postlarva is most similar to Nybelinia

aequidentata Shipley & Hornell, 1906, described

by Shipley and Hornell (1906), Pintner (1927) and

Palm (1999). Shipley and Hornell (1906) recorded

the type specimen as having a scolex 4-5 mm

long. The hook shape of the present specimen

corresponds with those of the type, though the

hooks in the present specimen are larger (57-65

compared with 49 in the type according to Shipley

and Hornell (1906) or 33-38 reported by Palm

(1999). Pintner (1927) redescribed the type and

added information on the bulb ratio (4.3:1), hook

length (48) and scolex measurements (SW = 1900,

pbo = 1500, pb = 1600 and vel = 600). He also

remarked that the hooks were not uniform in size

but increased from the base towards the metabasal

region and then decreased in the apical region of

the armature. Thus, although the scolex and hook

sizes of the present specimen are distinctly larger

than those described from the type specimen, it

has been identified as N. aequidentata. The

present finding represents a new host and locality

record.

Nybelinia syngenes Pintner, 1929 is the only

other species of Nybelinia which has been

described as having tentacular hooks with a length

greater than 60 and with a similar hook form and

armature pattern. However, the scolex of this

species, described originally from Sphyrna

zygaena (Linnaeus, 1758), is distinctly smaller.

Whether N. aequidentata exhibits variation in

scolex size, as described for N. lingualis (Cuvier,

1817) and Heteronybelinia yamaguti (Dollfus,

1960) (see Dollfus 1942; Palm et al. 1997; Palm

1999; Palm & Walter 2000) is not known. Palm

(1999) has suggested that this might be the case

for a postlarva from Lepturacanthus savala

(Cuvier, 1829) with a scolex length of 3400, a

bulb ratio of 3.3:1 and slender falciform hooks

33-38 in length.

Nybelinia africana Dollfus, 1960

(Figs 3-5)

Material examined

From unidentified shark: 1 adult, Queensland,

coll. J. C. Pearson (QM G 218169).

Description

SL = 1780 (Fig. 3); SW = 1134; pbo = 930;

pv = 840; pb = 523; vel =417; BL = 439 (423-

459); BW=163 (157-171); BR=2.7:1;

SP = 1.8:1.6:1. TW metabasal = 47-50. Basal

tentacular swelling absent. Tentacle sheaths

spirally coiled; TSW = 65-70. Prebulbar organs

absent, muscular rings around basal part of

tentacle sheaths not seen. Retractor muscles

52 ‘ H. W. PALM & I. BEVERIDGE

FIGURES 1-2. Nybelinia aequidentata Shipley & Hornell, 1906 from Dendrochirus zebrae. QM G 218031. 1.

Scolex. 2. Homeomorphous armature with falciform hooks. Scale bars: Figure 1, 500 pm; Figure 2, 100 pm.

FIGURES 3-5. Nybelinia africana Dollfus, 1960 from an unidentified shark. QM G 218169. 3. Scolex. 4. Basal

and metabasal armature. 5. Mature proglottid. Scale bars: Figure 3, 200 um; Figure 4, 20 um; Figure 5, 100 pm.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 53

originate in basal parts of bulbs. Tentacular

armature consists of homeomorphous hooks along

tentacle (Fig. 4). Tentacles not completely

evaginated; metabasal armature differs distinctly

from basal armature. Basal 2—3 rows of rosethorn-

shaped hooks with distinct anterior extension of

base, L = 14-16, 16-17; metabasal hooks larger,

falciform with small base, strongly recurved at tip,

L = 26-27; B = 10-13. Strobila 57 mm long, with

about 215 acraspedote segments which enlarge in

size towards end of strobila, maximum width 1.48

mm; velum straight or very slightly scalloped; first

segments 15-30 x 600-675, mature segments

(Fig. 5) wider than long, 220-470 x 650-970

(n=4). Pre-gravid segments (with some eggs)

440-670 x 1150-1480 (n = 5); terminal pre-gravid

segment with rounded end. In mature segments,

genital pore ventro-submarginal, in anterior half

or anterior third of segment; genital pore to

anterior end 90-150 in mature and 180-200 in

pre-gravid segments; genital pores alternate

irregularly. Cirrus sac thin-walled, elongate, 370—

480 x 70-100 (n=5), length:width ratio 4.5:1

(3.7-5.0:1), distal pole directed anteromedially,

nearly reaching anterior end of proglottid, cirrus

unarmed; internal and external seminal vesicle

absent. Vas deferens greatly coiled, extends to

midline of segment, then posteriorly towards

female genital complex. Testes of varying shape,

60-90 in diameter, arranged in single layer

centrally and in double layer peripherally; testis

number 50-71 per segment, between 5 and 8

testes anterior to cirrus (n= 5). Ovary follicular,

in centre of segment, bilobed, each lobe 190-260

x 140-220, increasing in size along strobila.

Vitelline follicles encircle medulla, follicles 30-

50 in diameter. Ventral and dorsal osmoregulatory

canals 30 in diameter, testes extend external to

ventral canal but not beyond dorsal canal.

Remarks

The present specimen closely resembles N.

africana Dollfus, 1960 as described by Dollfus

(1960) and Palm (1999) in having rosethorn-shaped

basal hooks, changing to falciform metabasal hooks

of similar size and shape. Additionally, the segment

morphology with number and size of segments,

size of cirrus sac, testes and vitelline follicles, as

well as arrangement of the genital complex is

similar. However, some differences were observed

between the present specimen and WN. africana. The

specimen from Australia differs in having 2 to 3

rows of basal hooks with a distinct anterior

extension of the base (Fig. 4), which has not been

reported in other specimens of N. africana.

However, Dollfus (1960, figs 14, 17) illustrated

comparable basal hooks, some with an anterior

extension of the base. The scolex is larger in the

Australian specimen (1780) compared with other

material (1118-1568) (Palm et al. 1997) and the

testis number is smaller. However, as most other

morphological characters correspond, the specimen

is identified as N. africana and the observed

differences are considered to be due to intraspecific

morphological variability. The present finding is a

new locality record for the species.

Nybelinia hemipristis sp. nov.

(Figs 6-9)

Types

Holotype from stomach of Hemipristis elongata

Klunzinger, 1871, Balgal, Queensland, coll. B. G.

Robertson, 16.ix.1985 (SAM AHC 28309).

Paratypes: 3 specimens, Hemipristis elongatus,

Marchinbar Island, Northern Territory, coll. G.

Cuthbertson, 25.v.1985 (SAM AHC 28310).

Material examined

Types.

Description

SL = 2030 (Fig. 6); SW = 1230; pbo = 1000;

pv = 860; pb = 560; vel = 630; BL=553 (550-

560); BW=218 (210-220); BR = 2.5:1;

SP = 1.8:1.5:1. Tentacles short, massive,

decreasing in diameter towards apical region;

basal tentacular swelling absent. TW basal and

metabasal = 75-85, TW apical = 37.5-62.5.

Tentacle sheaths straight (TSW = 50-60),

prebulbar organs and muscular rings around basal

part of tentacle sheaths not seen. Retractor

muscles originate in basal parts of bulbs.

Tentacular armature homeoacanthous,

homeomorphous (Fig. 7); distinctive basal

armature absent (Fig. 8). Hooks slender,

rosethorn-shaped with anterior elongation of base,

increasing in size from basal (L = 15.5-30.0;

B = 16.3-20.0) towards metabasal regions

(L = 35.0-40.0; B = 25.0-27.5), then decreasing

in size towards apical (L = 27.5-32.5; B = 15.0—

20.0) region; hsr = 6. Strobila acraspedote, velum

straight; 33 mm long, maximum width 1030,

number of segments 300. Mature segments (Fig.

9) wider than long, 600-670 x 750-920. Genital

pore submarginal, 200 from anterior end,

alternating irregularly. Cirrus sac short and stout,

180-210 x 50-100 (n=5S), length:width ratio

2.9:1 (2.0-3.6:1), distal pole directed

54 H. W. PALM & I. BEVERIDGE

SAN ae fe weay

a Les Boy

< es

FIGURES 6-9. Nybelinia hemipristis sp. nov. from Hemipristis elongata. Holotype, SAM AHC 28309. 6. Scolex.

7. Metabasal armature. 8. Basal armature. 9. Mature proglottid. Note short cirrus sac, the uterine duct (ud), and the

Mehlis’ gland (mg). Scale bars: Figure 6, 200 um; Figures 7-8, 40 um; Figure 9, 100 pm. FIGURES 10-11.

Nybelinia jayapaulazariahi Reimer, 1980 from a ‘sole’. QM G 207318. 10. Metabasal armature. 11. Basal

armature. Scale bars: Figures 10-11, 10 um.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 55

anteromedially, not reaching anterior end of

segments, cirrus unarmed; internal and external

seminal vesicle absent. Vas deferens coiled,

extends to midline of segment, then posteriorly

towards female genital complex. Testes 84-97 in

number, 55-85 in diameter centrally and 40-60

peripherally, distributed in single layer; arranged

in 2 lateral groups, confluent posterior to ovary;

extend between ovarian lobes, about 6—7 testes

anterior to cirrus sac. Ovary in centre of segment,

ovarian lobes 200-260 x 120-160. Vagina ventral

to cirrus sac. Vitelline follicles encircle medulla;

follicles 2540 in diameter.

Remarks

The present specimens, with rosethorn-shaped

hooks diminishing in size towards the apical

armature and a strobila consisting of acraspedote

segments, resemble N. anthicosum Heinz &

Dailey, 1974. However, the species differ in

having different scolex forms, tentacles, tentacular

armatures and sizes. N. anthicosum has an

elongated prominent velum, long tentacles and

tentacular hooks which are distinctly spaced (see

Heinz & Dailey 1974). The largest hooks occur in

the 7th to 9th rows. N. hemipristis sp. nov. has a

massive scolex with a shorter velum (Fig. 6), short

tentacles and tentacular hooks which are tightly

spaced along the tentacle. The largest hooks are in

the 10th to 14th rows. In the last few hook rows,

the size and number of hooks per half spiral row

decrease rapidly (Fig. 7). In N. anthicosum the

hook size diminishes gradually towards the apical

part of the tentacle. As with N. anthicosum, N.

hemipristis is a species without a characteristic

basal armature and with a smaller basal than

metabasal hook size, therefore belonging to

species subgroup ‘Aa’ of Palm (1999).

The description of the strobilar characters of N.

hemipristis demonstrates that this species has a

very characteristic, short, stout cirrus sac, with a

length: width ratio of 2 in some segments, a feature

unusual in most species of Nybelinia. However,

this character might prove to be of taxonomic

significance in the future.

The new species is named after

elasmobranch host genus, Hemipristis.

the

Nybelinia jayapaulazariahi Reimer, 1980

(Figs. 10-11)

Material examined

From ‘sole’, either Synaptura nigra Macleay,

1880 or Aseraggodes macleayanus (Ramsay,

1881): 1 postlarva, Moreton Bay, Queensland,

coll. J. C. Pearson, 1968 (QM G 207318).

Description

SL = 1060; SW = 700; pbo = 470; pv = 450;

pb = 365; BL =323 (320-325); BW=119 (85-

135); BR =2.7:1; SP = 1.3:1.2:1. TW = 32.5-35;

hsr = 6. Basal tentacular swelling absent. Retractor

muscles originate in basal parts of bulbs. Tentacular

armature consists of homeomorphous slender

uncinate hooks, L = 15.0-16.3, B = 11.2-13.7 (Fig.

10). Size of hooks diminishes towards basal part of

tentacle, L = 11.2-13.7, B = 8.7-11.2 (Fig. 11).

Remarks

N. jayapaulazariahi was originally described

from Cynoglossus sp. from the Bay of Bengal,

India by Reimer (1980). Palm (1999) redescribed

the species from another host, Harpodon nehereus

(Hamilton-Buchanan, 1822), also from India. The

tentacular armature has characteristic, slender,

regularly curved hooks which increase in size

towards the metabasal part of the tentacle. The

present specimen, also from a ‘sole’, has a similar

hook shape (compare Fig. 4 with Fig. 5 of Palm

1999) and bulb ratio, while it differs in having a

larger scolex (1060 in the present specimen

compared with 530 described previously) and

hook size (11.2-16.3 in the current specimen

compared with 5.6—11.2 in previous descriptions).

However, the present specimen was in a poor state

of preservation, and the larger scolex might be

related to the slightly larger hook size. A small

scolex of about 0.5-1.0 mm, together with the

uncinate hook form, is characteristic for N.

jayapaulazariahi, and soles seem to be important

intermediate hosts. The present finding represents

a new locality record for the species.

Nybelinia mehlhorni sp. nov.

(Figs 12-14)

Types

Holotype from stomach of Hemigaleus

microstoma Bleeker, 1852, Heron Island,

Queensland, coll. P. McBoarman, 20.xii.1995

(QM G 218032); paratype, same data (QM G

218033).

Material examined

Types.

Description

SL = 560 (Fig. 12); SW = 400; pbo = 330;

56 H. W. PALM & I. BEVERIDGE

FIGURES 12-14. Nybelinia mehlhorni sp. nov. from Hemigaleus microstoma. Types, QM G 218032-3. 12.

Scolex. 13. Metabasal and apical armature. 14. Mature proglottid. Note the uterine duct (ud) and the seminal

receptacle (sr). Scale bars: Figure 12, 50 ym; Figure 13, 10 ym; Figure 14, 50 pm. FIGURES 15-16. Nybelinia

scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996) from Diodon hystrix. QM G 218035-7. 15.

Scolex. 16. Basal and metabasal armature, external surface. Scale bars: Figure 15, 150 ym; Figure 16, 25 um.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 57

pv = 125; pb = 250; vel = 200; BL= 178 (150-

200); BW=58 (50-70); BR=3.1:1;

SP = 1.3:0.5:1. Tentacles short; basal tentacular

swelling absent; TW = 22.5-27.5. Tentacle

sheaths sinuous to spirally coiled (TSW = 17.5—

20.0); muscular rings around basal part of tentacle

sheaths seen. Retractor muscles originate in basal

parts of bulbs. Tentacular armature

homeoacanthous, homeomorphous (Fig. 13);

distinctive basal armature absent. Hooks

falciform, with stout base, elongated handle and

strongly recurved tip. Hooks increase in size from

basal (L = 3.5-7.5, B = 2.5—5.0) to metabasal

region (L = 15.0-17.5, B = 6.3-8.7) then decrease

towards apex (L = 11.3-13.8, B = 2.5-4.3);

hsr = 7. Small worms, length 25 (16) mm,

maximum width 950 (700) with 135 (90)

segments. Strobila acraspedote, velum straight;

pre-mature (testes visible) proglottids (Fig. 14)

80-280 long x 560-790 wide; mature segments

(female genitalia developed) wider than long,

380-600 x 600-900; terminal segment 830 x 800.

Genital pore almost lateral, 130-200 from anterior

end (190 in terminal segment); pores alternate

irregularly. Cirrus sac elongate, 170-220 x 40-60

(n=5) in mature segments, length:width ratio

4.1:1 (2.8-5.5:1), distal pole directed

anteromedially, not reaching anterior end of

segments; cirrus unarmed, internal and external

seminal vesicles absent. Vas deferens greatly

coiled in midline of segments, extending

posteriorly towards female genital complex.

Testes 103-120 in number, 80 x 95 in size,

smaller testes peripheral, 35-40 in size,

distributed in single layer; testes in 2 lateral

groups confluent posterior to ovary; 10-16 testes

anterior to cirrus sac. Vagina tubiform, 30 in

diameter, ventral to cirrus sac, passes

anteromedially to midline, then posteriorly to

ovary; seminal receptacle present, 60-90 x 25-50.

Ovary in centre of segment, ovarian lobes 250—

300 x 160-210. Vitelline follicles encircle

medulla, follicles 30-45 in diameter. Uterine pore

absent.

Remarks

Following Palm (1999), N. mehlhorni sp. nov.

belongs to Nybelinia species without a

characteristic basal armature and a basal hook size

smaller than the metabasal one, thus belonging to

the subgroup ‘Aa’.

On the basis of the hook shape, the species

resembles N. aequidentata (Shipley & Hornell,

1906) and N. goreensis Dollfus, 1960. While the

former is much larger than the present specimen,

the latter species has different scolex proportions

as well as a different hook arrangement, as

illustrated recently by Palm and Walter (2000).

N. mehlhorni sp. nov. also resembles N.

bengalensis Reimer, 1980 in its falciform hooks

and in hook size (Reimer 1980). However, N.

bengalensis can be distinguished by having a

distinctly different scolex form and size, a bulb

ratio of about 2, long and slender tentacle sheaths

and tentacles with spaced falciform hooks (Reimer

1980). By contrast, N. mehlhorni sp. nov. has

short tentacles with hooks tightly spaced along the

tentacle. The scolex (0.56 mm in total length) is

much smaller.

The new species is named in honour of Prof.

Heinz Mehlhorn, Heinrich-Heine-University,

Diisseldorf.

Nybelinia schmidti Palm, 1999

Material examined

From /surus oxyrhinchus Rafinesque, 1810: 1

adult, Bicheno, Tasmania, coll. B. G. Robertson,

24.iv.1987 (SAM AHC 28313).

Description

SL = 1600; SW = 850; pbo = 1040; pv = 680;

pb = 340; vel = 600; BL = 298 (270-330);

BW = 131 (110-150); BR = 2.3:1; SP = 3.0:2.0:1.

TW basal = 45-48, TW metabasal = 30.0-32.5;

hsr = 5-6. Basal tentacular swelling absent.

Tentacle sheaths spirally coiled, TSW = 25—40.

Prebulbar organs absent, muscular rings present

around the tentacle sheaths. Retractor muscles

originate in basal part of bulbs. Tentacular

armature homeoacanthous, homeomorphous,

consisting of massive rosethorn-shaped hooks

with anterior extension of base, size in metabasal

region L = 13.8-15.0, B = 11.3—12.5, decreasing

towards base, L=8.8-11.3, B = 6.2-8.8.

Characteristic basal armature absent. Strobila

immature with few segments, acraspedote;

segments wider than long (660-690 x 15-60).

Remarks

The present specimen closely resembles that

of the type, also from Jsurus oxyrhinchus (syn.

Tsurus glaucus), in having similar scolex

proportions and almost identical hook sizes.

Other species with a similar tentacular armature

are Nybelinia strongyla Dollfus, 1960 and

Heteronybelinia australis sp. nov. (vide infra).

While the latter species differs in having

homeomorphous hooks of different sizes on the

58 H. W. PALM & I. BEVERIDGE

internal and external tentacle surfaces as well as

smaller basal hooks, N. strongyla differs in

scolex proportions and larger hook sizes. The

present record extends the range of distribution

from the South African to the southern

Australian coast.

Nybelinia scoliodoni (Vijayalakshmi,

Vijayalakshmi & Gangadharam, 1996)

(Figs 15-16)

Material examined

From Diodon hystrix Linnaeus, 1758: Heron

Island, Queensland, 2 postlarvae, coll. J. Sakanari,

23.viii.1986 (QM G 218034); 8 postlarvae, coll.

M. K. Jones, 6.vii.1995 (QM G 218035-218037);

8 postlarvae, coll. T. H. Cribb, Jan. 1997 (QM G

218038-218041).

From Makaira indica (Cuvier, 1832): 1

postlarva, Cape Bowling Green, Queensland, coll.

P. Speare, 18.ix.1987 (SAM AHC 21351).

From Istiophorus platypterus (Shaw & Nodder,

1792): Whitsunday Island, Queensland, coll. P.

Speare, 5.xii.1988 (SAM AHC 21351).

From Carcharhinus limbatus (Valenciennes,

1839): 1 adult, stomach, Bremer Island, Northern

Territory, coll. J. Stevens, 29.v.1985 (SAM AHC

28314).

Description

SL = 1350, 1115 (Fig. 15); SW = 860, 800;

pbo = 670, 520; pv = 230, 210; pb = 300, 320;

vel = 330, 330; app = 310, 200; BL = 288 (280—

300), 282 (260-300); BW = 115 (110-120), 125

(120-130); BR =2.5:1, 2.3:1; SP = 2.2:0.8:1,

1.6:0.7:1. Tentacles nearly completely evaginated,

TL = 390, 375; basal tentacular swelling absent.

TW at basal armature 35, 30; TW at metabasal

armature 50, 45; TW at apex 30, 25. Tentacle

sheaths sinuous, TSW = 30, 40; prebulbar organs

and muscular rings around basal part of tentacle

sheaths not visible. Retractor muscles originate in

basal parts of bulbs. Metabasal armature

homeoacanthous, homeomorphous; distinctive

basal armature present (Fig. 16). Basal armature

consists of about 10 rows with compact rosethorn-

shaped hooks, increasing in size from L = 4.5,

B =3.9 (row 1) to L= 19.5, B=11.7 (row 10).

From row 11, hook form changes to slender

spiniform, L = 22.7, B = 13.0. Metabasal and

apical hooks L = 29.9, with small base, B= 11.

Number of hooks per half row (hsr) 6-7 in basal

region, decreasing to 5 in metabasal and apical

regions.

Remarks

Nybelinia scoliodoni is a widely distributed

tentaculariid and is here recorded for the first time

from Australian waters. The species is easily

recognised by its rosethorn-shaped basal armature

of about 11 rows which is followed by long,

spiniform metabasal hooks. The present

specimens correspond with the description given

by Palm (1999) in having a similar armature and

bulb ratio. They differ in having a larger scolex

and hook size. However, they correspond closely

to the original description of Nybelinia

(= Tentacularia) scoliodoni of Vijayalakshmi et

al. (1996). The present findings constitute three

new host records. C. limbatus has been recorded

previously as a definitive host for N. scoliodoni

by Palm (1999). The specimens described here are

the same as those reported by Speare (1999).

Nybelinia strongyla Dollfus, 1960

(Figs 17-20)

Material examined

From Argyrosomus hololepidotus Lacépéde,

1802: 2 postlarvae from stomach, Murray Mouth,

South Australia, coll. M. G. O’Callaghan, May

1992 (SAM AHC 28345).

From Johnius vogleri (Bleeker, 1853): 1

postlarva from body cavity, Heron Island,

Queensland, coll. J. Reddich, Jan 1997 (QM G

218109).

Description

Postlarvae from A. hololepidotus: SL = 1303,

1566 (Fig. 17); SW = 648, 796; pbo = 700, 781;

pv = 647, 764; pb = 292, 355; vel = 332, 326;

app = 374, 429; BL = 286 (244-292), 355 (334—

380); BW = 86 (82-90), 98 (92-107); BR = 3.3:1,

3.6:1; SP = 2.4:2.2:1, 2.2:2.2:1. Tentacles not

completely evaginated; basal tentacle swelling

absent. TW = 50-58, 43-45. Tentacle sheaths

sinuous, TSW = 36-40, 36-40; prebulbar organs

and muscular rings around basal part of tentacle

sheaths not visible. Retractor muscles originate in

basal part of bulbs. Tentacle armature

homeoacanthous, homeomorphous (Fig. 18);

distinctive basal armature absent. Hooks slender,

rosethorn-shaped, increasing in size from basal

towards metabasal part of tentacle. Metabasal

hooks L = 17.5—20.0; B = 13.8-16.3; basal hooks

L= 10.0-12.5; B = 10.0-12.5; hsr = 7.

Postlarva from J. vogleri: SL = 1880;

SW = 1660; pv = 680; pb = 440; BL = 308 (300-

325); BW = 129 (115-150); BR = 2.4:1;

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 59

FIGURES 17-18. Nybelinia strongyla Dollfus, 1960 from Argyrosomus hololepidotus. SAM AHC 28345. 17.

Scolex. 18. Metabasal armature. Scale bars: Figure 17, 150 um; Figure 18: 15 um. FIGURES 19-20. Nybelinia

strongyla from Johniops vogleri. QM G218109. 19. Metabasal armature. 20. Basal armature. Scale bars: Figure

19-20: 10 um. FIGURES 21-24. Nybelinia thyrsites Korotaeva, 1971 from Trachurus declivis. QM G 214194-S.

21. Scolex. 22. Basal armature. 23. Metabasal armature with falciform hooks. 24. Apical armature with rosethorn-

shaped hooks. Scale bars: Figure 21, 200 um; Figures 22—24, 25 um.

60 H. W. PALM & I. BEVERIDGE

SP = -:1.5:1. TW. basal = 37-40, TW

metabasal = 30-33. Basal tentacular swelling

absent. Tentacle sheaths straight; TSW = 20-25.

Prebulbar organs absent, muscular rings around

basal part of tentacle sheaths absent. Retractor

muscles originate in basal part of bulbs. Armature

homeoacanthous, homeomorphous (Fig. 19);

characteristic basal armature absent (Fig 20).

Hooks rosethorn-shaped with anterior extension

of basal plate; hooks in basal part of tentacle

smaller (L = 7.5-10.0; B = 7,.5—9.5) than in

metabasal (L = 13.8-16.3; B = 10.0—12.5)

armature. Number of hooks per half spiral (hsr)

diminishes from 6—7 in metabasal region to 5-6

in apical part of tentacle.

Remarks

Nybelinia strongyla Dollfus, 1960 was

described from a postlarva from the west coast of

Africa, and is characterised by the

homeoacanthous tentacular armature with slender

rosethorn-shaped hooks, reaching a size of up to

16 in the metabasal armature. Palm and Walter

(2000) described adult specimens of N. strongyla

from Africa. The present specimens correspond

closely in their tentacular armature as well as in

scolex proportions to those specimens described

by Dollfus (1960) and Palm and Walter (2000).

Minor variations in the hook pattern are observed

within N. strongyla but are difficult to define, and

many scolex characters within tentaculariids

appear to be variable (Palm 1999, Palm and

Walter 2000). Therefore, the postlarvae described

above are identified as N. strongyla. However,

some doubt surrounding the identification

remains. Further studies are needed to determine

whether postlarvae with rosethorn-shaped hooks

as described above are conspecific with the

African material, or whether they belong to a new

species of Nybelinia. Argyrosomus hololepidotus

is a new host record, and the distribution of the

species is extended to the southern Australian

coast.

The specimen from J. vogleri, with rosethorn-

shaped hooks, resembles N. lingualis, N. schmidti,

N. sphyrnae, N. strongyla and N. thyrsites. It

differs from N. lingualis in having basal hooks

with an anterior extension of the basal plate, and

from N. thyrsites in having differently shaped

metabasal hooks and a larger bulb ratio (see

below). N. schmidti differs in having smaller

hooks and different scolex proportions. N.

sphyrnae differs in having slender, more uncinate

hooks in the metabasal armature, while the hooks

are more massive rosethorn-shaped in N.

strongyla. In the present specimen, the basal hook

length is slightly smaller than described by Palm

and Walter (2000) for NV. strongyla. However, the

metabasal hooks are the same size. Although the

scolex proportions were difficult to measure in

the present specimen, it is also identified as N.

strongyla, representing a new host and locality

record.

Nybelinia thyrsites Korotaeva, 1971

(Figs 21-24)

Material examined

From Trachurus declivis Jenyns, 1841: 2

postlarvae from stomach wall, Crayfish Bay,

Tasmania, coll. K. B. Sewell, 11.v.1987 (QM G

214194, 212145).

From Carcharhinus brachyurus (Giinther,

1870): 1 adult from stomach, Goolwa, South

Australia, coll. R. Martin, 28.x.1985 (SAM AHC

28312).

From Mustelus antarcticus Giinther, 1870: 1

adult from stomach, Goolwa, South Australia,

coll. R. Martin, 28.x.1985 (SAM AHC 28311).

Description

SL = 2032, 1550 (Fig. 21); SW = 1520, 700;

pbo = 1024, 610; py = 640, 500; pb = 368, 520;

vel = 992, 490; app = 496; BL = 358 (355-365),

495 (490-500); BW = 117 (114-120), 155 (140-

170); BR=3.1:1, 3.2:1; SP =2.8:1.7:1, 1.2:1:1,

Tentacle sheaths spirally coiled; TSW = 51-57,

35-45. Prebulbar organs absent, muscular rings

around basal part of tentacle sheaths present.

Retractor muscles originate in basal part of bulbs.

Tentacles long (TL = 832) and slender, basal

tentacular swelling absent; TW basal = 38-44,

25-30; TW metabasal = 57.0, 57.7-62.5; TW

apical 32.0, 27.5. Characteristic basal armature

present (Fig. 22), consisting of about 7-8 rows of

homeomorphous, rosethorn-shaped hooks with

slight anterior extension, increasing in size

towards metabasal armature (lst row hooks:

L = 9.5-11.0, B = 8.0-11.0, L = 7.5-8.7, B = 6.3-

7.5; 8th row: L= 15.8-17.4, L = 16.2-17.5,

B = 11.0-12.6, B = 10.0-12.5). Metabasal

armature of about 12-14 rows of hooks, with

largest hooks in rows 12-17. Metabasal armature

with slender falcate to falciform hooks (L = 21.7—

26.9, B = 8.7-11.0; L = 21.3-24.0, B = 11.3-

12.5) (Fig. 23). Apical hooks rosethorn-shaped,

with hook form changing from slender rosethorn-

shaped with slight anterior extension to rosethorn-

shaped with distinct anterior extension (Fig. 24).

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 61

Hooks diminish in size towards apical region

(34th row of hooks: L= 14.2-15.8, B= 11.0-

14.2); hsr basal: 6—7; hsr metabasal and apical: 6.

Remarks

N. thyrsites was redescribed in detail by

Beveridge and Campbell (1996). The present

material corresponds with the redescription given.

In addition, the basal hook form differs from the

metabasal and apical hooks; thus, the species

belongs to subgroup ‘Ba’ of Palm (1999). The

present specimens from Trachurus declivis

provide a further example in which the postlarvae

can be larger than adult worms, and also show

variable scolex measurements depending on the

state of contraction. Trachurus declivis and

Carcharhinus brachyurus are new hosts for N.

thyrsites.

Nybelinia victoriae sp. nov.

(Figs 25-26)

Types

Holotype from body cavity of Lepidotrigla

modesta Waite, 1899, Port Phillip Bay, Victoria,

coll. R. Norman, 17.iii1.1989 (SAM AHC 28343);

paratype, same data (SAM AHC 28344).

Material examined

Types.

Description

SL = 1030, 1050 (Fig 25); SW=700, 650;

Pbo = 570, 450; Pv = 350, 320; Pb =315, 320;

BL = 310 (300-315), 310 (305-320); BW = 124

(115-135), 116 (110-125); BR =2.5:1, 2.7:1;

SP = 1.8:1.1:1, 1.4:1.0:1. Tentacles elongate;

basal tentacular swelling absent. TW = 40.0-50.0,

45.0-S0.0. Tentacle sheaths sinuous to spirally

coiled (TSW = 40.0-S0.0, 35.0-40.0); prebulbar

organs and muscular rings around basal part of

tentacle sheaths not seen. Retractor muscles

originate in basal part of bulbs. Armature

homeoacanthous, homeomorphous (Fig. 26);

distinctive basal armature absent. Hooks

falciform, with stout base, elongated handle and

strongly recurved tip. Hooks increase in size from

basal (L = 7.5-10.0, B = 6.3-8.8) to metabasal

region (L = 12.5—15.0, B = 7.0-8.0) and decrease

towards apical (L = 11.3-12.5, B = 6.3-8.0) part

of tentacle; hsr = 7.

Remarks

This species, with a tentacular armature

consisting of slender falciform hooks, increasing

in size towards the metabasal region and

decreasing in size towards the apical region,

resembles Nybelinia aequidentata Shipley &

Hornell, 1906; N. syngenes Pintner, 1927; N.

anantaramanorum Reimer, 1980; and N.

bengalensis Reimer, 1980. While the former two

species differ in having different scolex

proportions and distinctly larger hooks (see

remarks on N. aequidentata), the present

specimens are most similar to N. bengalensis and

N. anantaramanorum. N. anantaramanorum has

larger hooks and was considered a species

inquirenda by Palm and Walter (2000). N.

bengalensis differs in having different scolex

proportions, including very short bulbs (BR < 2)

(Reimer 1980). Thus, the present specimens

represent a new species here named Nybelinia

victoriae sp. nov., the specific epithet being

derived from the Australian state from which

samples were collected. With hook size smaller in

the basal rather than the metabasal region, the

species is considered to belong to subgroup ‘Aa’

of Palm (1999).

Genus Heteronybelinia Palm, 1999

Heteronybelinia australis sp. nov.

(Figs 27-30)

Types

Holotype from stomach of Carcharhinus

amboinensis (Miiller & Henle, 1839), St

Lawrence, Queensland, coll. B. G. Robertson,

29.x.1985 (SAM AHC 28315); paratype, same

data (SAM AHC 28316).

Material examined

From Carcharhinus amboinensis: types.

From Carcharhinus brachyurus: 1 specimen,

Goolwa, South Australia, coll. R. Martin,

28.x.1985 (SAM AHC 28317).

Description

SL = 1190, 1210 (Fig. 27); SW = 660, 680;

pbo = 650, 700; pv = 530, 650; pb = 350, 310;

vel = 220, 190; BL = 313 (305-320), 308 (290—

340); BW = 138 (125-150), 119 (110-130);

BR = 2.3:1, 2.6:1; SP = 1.9:1.7:1, 2.3:2.1:1.

Tentacle sheaths spirally coiled. TSW = 27.5-—

32.5, 25.0-35.0. Tentacles robust, TL = 255-290,

increasing in width towards apex. TW basal and

metabasal 30.0—32.5; basal tentacular swelling

absent. Prebulbar organs absent; muscular rings

62 H. W. PALM & I. BEVERIDGE

FIGURES 25-26. Nybelinia victoriae sp. nov. from Lepidotrigla modesta. Types, SAM AHC 28343-4. 25. Scolex.

26. Basal and metabasal armature. Scale bars: Figure 25, 100 pm; Figure 26, 20 um. FIGURES 27-30.

Heteronybelinia australis sp. nov. from Carcharhinus amboinensis. Types, SAM AHC 28315-6. 27. Scolex. 28.

Metabasal armature. 29. Basal armature. 30. Mature proglottid. Note the uterine pore. Scale bars: Figure 27, 100

um; Figures 28-29, 10 um; Figure 30, 50 um.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 63

around basal parts of tentacle sheaths present.

Retractor muscles originate at base of bulbs.

Tentacular armature homeoacanthous,

heteromorphous, with hooks tightly spaced along

tentacle (Fig. 28); characteristic basal armature

absent (Fig. 29). Hooks massive, rose-thorn

shaped, with slight anterior extension of base,

increase in size from basal towards metabasal

armature. Hook size differs on bothridial/

antibothridial (L = 12.5-15.0, B = 11.3-13.7) and

antibothridial/bothridial (L = 11.3-12.5, B = 7.0—

8.7) tentacle surfaces (Fig. 28), with two tentacles

having largest hooks on bothridial surfaces and

two other tentacles having largest hooks on

antibothridial surfaces. Basal hooks L = 5.0-7.5,

7.5-8.7; B =8.7-10.0 (internal) and L = 5.0-6.3,

5.0-7.5; B = 3.8-5.0, 5.0-6.3 (external); hsr = 6—

7. Strobila only very slightly craspedote, velum

scalloped; total length 18.0 and 15.0 mm,

maximum width 640 and 730, with 125 and 155

segments. In holotype, terminal segment pre-

mature (320 x 550), segments wider than long,

231 x 630. Genital pores ventro-submarginal, 60

from anterior margin; alternate irregularly. Cirrus

sac of pre-mature segments elongate and slender,

210-230 x 35-50 (n = 5), length:width ratio 5.3:1

(4.6-6.0:1), with distal pole directed

anteromedially, reaching anterior end of segment

(Fig. 30); cirrus unarmed; internal and external

seminal vesicles absent. Vas deferens coiled,

extends posteriorly from cirrus sac towards female

genital complex. Testes 106-130 in number,

smallest testes at margin of medulla, distributed in

1-2 layers; in 2 lateral groups, confluent posterior

to ovary; extend between ovarian lobes; 8-10

testes anterior to cirrus sac. Ventral

osmoregulatory canal 15 in diameter, internal to

sinuous dorsal canal, 5 in diameter; 24 prominent

bands of longitudinal muscles on each side of

strobila. Segments in paratype mature, wider than

long (240 x 700 to 380 x 720). Genital pores 110

from anterior margin. Cirrus sac of mature

segments (Fig. 30) elongate and slender, 260-320

x 50-60 (n=5), length:width ratio 5.2:1 (4.3-

5.8:1). Central testes 45-65 in diameter, smallest

testes peripheral, 30-50 in diameter, distributed in

1-2 layers; ovarian lobes 110-200 x 80-110,

seminal receptacle SO-65 x 25-30.

Remarks

Heteronybelinia australis sp. nov. has a unique

armature. The tentacular hooks are tightly spaced

and appear homeomorphous along the tentacle.

However, the hook sizes differ on the bothridial

and antibothridial tentacular surfaces. In addition,

two tentacles have the largest hooks on the

bothridial surfaces while the other two tentacles

have the largest hooks on the antibothridial

surfaces. Whether this armature pattern is unique

or whether it occurs in other tentaculariid species

remains to be determined.

The specific epithet indicates the occurrence of

the species in Australian waters.

Heteronybelinia estigmena (Dollfus, 1960)

(Figs 31-36)

Material examined

From Carcharhinus limbatus Valenciennes,

1839: 16 adults from spiral valve, Geraldton,

Western Australia, coll. B. G. Robertson,

27.xi.1986 (SAM AHC 21352, 28318); 4 adults

from spiral valve, Nickol Bay, Western Australia,

coll. B. G. Robertson, 11.xi.1986 (SAM AHC

28319); 3 adults from spiral valve, Darwin

Harbour, Northern Territory, coll. B. G.

Robertson, 28.viii.1986 (SAM AHC 28320); 6

adults from spiral valve, Fog Bay, Northern

Territory, coll. B. G. Robertson, 4.x.1986 (SAM

AHC 28329).

From Carcharhinus amblyrhynchoides

(Whitley, 1934): 2 adults from spiral valve,

Tommy Cut Mouth, Northern Territory, coll. B.

G. Robertson, 10.ix.1986 (SAM AHC 28321).

From Carcharhinus sp: 3 adults from spiral

valve, Queensland, coll. B. G. Robertson, Oct.

1985 (SAM AHC 18322).

From Sarda australis (Macleay, 1881): 9

postlarvae from gill arches, Heron Island,

Queensland, coll. M. K. Jones, Jan. 1991 (QM G

218042-218046).

Description

Adult: SL = 1650, 1500 (Fig. 31); SW = 790,

770; pbo = 820, 820; pv = 690, 610; pb = 480,

430; ppb = 110, 80; vel = 330, 320; BL = 455

(440-480), 425 (420-430); BW = 133 (130-

140), 120 (118-122); BR =3.4:1, 3.6:1;

SP = 1.7:1.4:1, 1.9:1.4:1. Tentacles long, slender

(TL = 520, 560), TSW = 42.5-47.5, 45.0-55.0;

TW basal 45-50, 42.5-45; TW apical 25-30,

27.5-32.5; basal tentacular swelling absent.

Prebulbar organs absent, muscular rings around

basal part of tentacle sheaths visible in one

specimen. Retractor muscles originate at bases

of bulbs. Tentacular armature homeoacanthous,

heteromorphous, no characteristic basal armature

present (Figs 32-33). Hooks rosethorn-shaped,

increase in size towards metabasal part of

64 H. W. PALM & I. BEVERIDGE

FIGURES 31-34. Heteronybelinia estigmena (Dollfus, 1960) from Carcharhinus limbatus. SAM AHC 28318-20.

31. Scolex. 32. Metabasal armature. 33. Basal armature. 34. Mature proglottid. Scale bars: Figure 31, 150 ym;

Figures 32-33, 15 um; Figure 34, 100 um. FIGURES 35-36. H. estigmena from Sarda australis. QM G 218042-

46. 35. Scolex. 36. Basal armature. Scale bars: Figure 35, 150 ym; Figure 36, 15 um.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 65

tentacle. Hook form varies from compact and

rosethorn-shaped (bothridial) to smaller

rosethorn-shaped hooks with elongated base

(antibothridial). Hook size, metabasal, L = 11.3-

12.5; B = 9.3-10.8 (bothridial) and L = 9.3-10.8;

B = 10 (antibothridial), with largest hooks

L = 13.8-15.0; B = 10.0-12.5 (bothridial); hooks

diminish in size towards apical part of tentacle.

Hook size in basal region L = 7.5-8.8; B = 6.3-

10.0 (bothridial) and L =5.0-7.5; B = 6.3-7.5

(antibothridial); hsr = 6-7. Strobila 56 mm long,

maximum width 1250, about 280 acraspedote

segments (Fig. 34). First segments behind velum

short, enlarge in size, mature segments wider

than long, from 500 x 840 to 700 x 1030. In

mature segments, genital atrium ventro-

submarginal, in anterior third of segment; genital

pores alternate irregularly. Cirrus sac elongated,

300-450 x 80-110 (n=5), length:width ratio

3.8-4.1:1, directed anteromedially, sac thin-

walled; cirrus unarmed and coiled within sac.

Ovary median, follicular, ovarian lobes 200—230

(length) x 110-170 (width), Mehlis’ gland 80 in

diameter, seminal receptacle 110 x 70; testes of

variable shape, 60-80 x 30-60 (n=5) in

diameter, arranged in single layer; testes number

106-126 (n=5) per segment, encircle female

genital complex and occupy entire medulla

except for region of female genital complex and

anterior to it; 7-9 testes anterior to cirrus sac.

Vitelline follicles 20-40 in diameter; egg

diameter 20-25; ventral osmoregulatory canals

along margins of cortex, 6 in diameter.

Postlarva: SL = 1650, 1500 (Fig. 35);

SW = 790, 770; pbo = 820, 820; py = 690, 610;

pb = 480, 430; ppb= 110, 80; vel = 330, 320;

BL = 460 (430-470), 445 (440-450); BW = 115

(110-120), 120 (118-122); BR =4.0, 3.7;

SP = 1.7:1.4:1, 1.9:1.4:1. Tentacles long, slender

(TL = 550-580), TSW = 40-45; TW basal = 35—

40, TW metabasal and apical = 30-40; basal

tentacular swelling absent. Prebulbar organs

absent, muscular rings around basal part of

tentacle sheaths present. Retractor muscles

originate at base of bulbs. Tentacular armature

homeoacanthous, heteromorphous; characteristic

basal armature absent. Hook shape varies from

compact, rosethorn-shaped (bothridial) to smaller

rosethorn-shaped hooks with elongated base

(antibothridial); hooks increase in size towards

metabasal part of tentacle on bothridial and

antibothridial surface (Fig. 36). Hook size,

metabasal, L=12.5-15.0, B= 11.8-13.3

(bothridial) and L = 11.3-12.5; B = 7.5-10.0

(antibothridial); basal L = 7.5—-10.0; B = 8.8-11.3

(bothridial) and L = 5.0-7.5; B =6.3-8.8

(antibothridial); hsr = 6-7.

Remarks

Heteronybelinia estigmena (Dollfus, 1960) is a

well-described tentaculariid. Palm (1995) reported

this cestode as Nybelinia alloiotica Dollfus, 1960

from Carcharhinus limbatus in the Atlantic, the

same host species as in the present study. Palm

(1999) and Palm and Walter (2000) provided

additional information on the species. H.

estigmena resembles H. perideraeus (Shipley &

Hornell, 1906) and H. elongata (Shah & Bilquees,

1979) known from the coast of India. However,

the hook size increases on both the bothridial and

antibothridial tentacle surfaces in H. estigmena. In

H. elongata, the hook size increases on the

antibothridial tentacle surface only (Palm &

Walter 1999), while in H. perideraeus, the basal

and metabasal hook sizes are the same (Palm

1999). The present study represents the first record

of the cestode from Australian waters.

Carcharhinus amblyrhynchoides and Sarda

australis represent new hosts for the species.

Heteronybelinia pseudorobusta sp. nov.

(Figs 37-41)

Types

Holotype from gills of Lepidotrigla modesta

Waite, 1899, Port Phillip Bay, Victoria, coll. R.

Norman, 17.iii.1989 (SAM AHC 28341); 3

paratypes, same data (SAM AHC 28342).

Material examined

Types.

Description

Small form (holotype and one paratype):

SL = 960 (Fig. 37), 730; SW = 660, 550;

pbo = 570, 450; pv = 420, 270; pb = 285, 290;

app = 220, 160; vel = 70, 60; BL = 280 (275-

285), 298 (275-315); BW = 150 (145-155), 139

(110-165); BR = 1.9:1, 2.1:1; SP = 2.0:1.5:1,

1.6:0.9:1. Tentacle sheaths sinuous to spirally

coiled; TSW = 45.0-47.5, 45.0-50.0; TW basal

45.0—-50.0, metabasal 40.0-45.0, basal tentacular

swelling absent. Prebulbar organs absent,

muscular rings around basal part of tentacle

sheaths not seen, retractor muscles originate at

base of bulbs. Tentacular armature

homeoacanthous, heteromorphous, characteristic

basal armature absent (Fig. 38). Hook form

changes from compact, rosethorn-shaped

H. W. PALM & I. BEVERIDGE

FIGURES 37-41. Heteronybelinia pseudorobusta sp. nov. from Lepidotrigla modesta. 37. Scolex of small form

SAM AHC 28314. 38. Basal armature. 39. Metabasal armature. 40. Scolex of large form SAM AHC 28342. 41.

Metabasal armature. Scale bars: Figure 37, 100 ym; Figures 38-39, 20 um; Figure 40, 100 um; Figure 41, 15 um.

FIGURES 42-44. Mixonybelinia cribbi sp. nov. from Platycephalus arenarius. Holotype, QM G 218047. 42.

Scolex. 43. Metabasal armature. 44. Basal armature. Scale bars: Figure 42, 200 um; Figures 43-44, 20 pm.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 67

(bothridial) to more slender hooks with stout base

(antibothridial) (Fig. 39). Hook size in metabasal

armature ranged between L = 13.8-16.3;

B = 11.3-13.8 (bothridial) and L = 16.3-17.5;

B = 10,0—11.3 (antibothridial), hooks of basal part

of tentacle smaller, between L = 10.0-11.0;

B = 11.3-9.0 (bothridial) and L = 11.3-12.5;

B = 8.8-10.0 (antibothridial), continuously

increasing towards the tip; hsr = 6-7.

Large form (2 paratypes): SL = 1310 (Fig. 40);

1310; SW = 600, 620; pbo = 810, 780; pv = 580,

500; pb = 390, 380; app = 330, 330; vel = 280,

310; BL = 376 (360-390), 321 (310-330);

BW = 124 (115-140), 121 (105-135);

BR =3.0:1; SP = 2.1:1.5:1. Tentacle sheaths

sinuous to spirally coiled; TSW = 40-45, 42.5-

47.5; TW basal 45.0-47.5, metabasal 30.0-35.0,

30.0-35.0. Basal tentacular swelling absent,

prebulbar organs absent, muscular rings around

basal part of tentacle sheaths not seen. Retractor

muscles originate at base of bulbs. Tentacular

armature homeoacanthous, heteromorphous,

characteristic basal armature absent. Hooks

change from compact and rosethorn-shaped

(bothridial) to more slender hooks with a stout

base (antibothridial) (Fig. 41). Hook size in

metabasal armature between L = 12.5-14.5;

B = 11.3-12.5 (bothridial) and L = 14.5—15.5;

B = 8.8-10.0 (antibothridial), hooks of basal part

of tentacle small, between L = 8.8-10; B = 7.5—

9.0 (bothridial) and L = 6.3-8.8; B = 6.3-7.5

(antibothridial), increasing in size towards tip;

hsr = 6-7.

Remarks

Heteronybelinia pseudorobusta sp. nov. is

characterised by a heteromorphous tentacular

armature consisting of small basal hooks

increasing in size towards the metabasal region.

The hook shape varies from rosethorn-shaped to

slender hooks with a stout base on different

surfaces of the tentacle. Small basal hooks which

gradually increase in size towards the metabasal

armature are characteristic of Heteronybelinia

robusta (Linton, 1890). However, in contrast to

H. pseudorobusta sp. nov., H. robusta has minute

basal hooks, less than 5 in length, and the hook

form is uniform along the tentacle. In the present

specimens, the basal hook size is larger and the

hook form is rosethorn-shaped. All other

Heteronybelinia species differ in having a

different hook shape or arrangement. Thus, the

present specimens belong to a new species, H.

pseudorobusta sp. nov., the name being derived

from the similarity in hook arrangement to H.

robusta (Linton, 1890). H. pseudorobusta is a

species without a characteristic basal armature and

with basal hook sizes smaller than metabasal hook

sizes. It therefore belongs to the Heteronybelinia

species subgroup ‘Aa’ in Palm (1999).

Heteronybelinia pseudorobusta sp. nov.

occurred in two different size groups, those

smaller than 1 mm and those larger than 1.3 mm,

respectively. Thus, the present description is an

example of intraspecific morphological variability

within species of Heteronybelinia, apparently due

to scolex contraction during fixation.

Genus Mixonybelinia Palm, 1999

Mixonybelinia beveridgei (Palm, Walter,

Schwerdtfeger & Reimer, 1997)

Material examined

From Macruronus novaezelandiae (Hector,

1871): 1 postlarva, west coast of Tasmania, coll.

K. Sewell, June 1986 (QM G 218067).

Description

SL = 3200; SW = 2750; pbo = 1750; pv = 850;

pb = 1300; ppb = 80; vel = 950; app = 730;

BL = 1226 (1150-1310); BW = 290 (240-320);

BR = 4.2:1; SP = 1.3:0.7:1. Tentacle sheaths

sinuous to spirally coiled; TSW = 100-130.

Prebulbar organs absent, muscular rings around

basal part of tentacle sheaths not seen. Retractor

muscles originate in basal parts of bulbs. Basal

tentacular swelling absent; TW basal = 110-120,

TW metabasal = 115-135. Metabasal armature

homeoacanthous, heteromorphous, characteristic

homeomorphous basal armature consisting of

about 6-7 rows of slender hooks with elongated

shaft, stout base and strongly recurved at tip

(L = 31.3+40.0, B = 18.8—27.5). Metabasal hooks

strongly recurved, rosethorn-shaped with large

base on antibothridial surface: L = 50.0-52.5,

B = 37.5-42.5; slender falcate hooks with stout

base on bothridial surface: L = 55.0-60.0,

B = 27.5-30.0; hsr = 6.

Remarks

The present specimen is characterised by a

homeomorphous basal armature of 6-7 rows of

hooks and a heteromorphous metabasal armature

of massive hooks, thus corresponding in form and

size with specimens of M. beveridgei described

from African waters by Palm et al. (1997). Other

features are the large scolex size as well as TSW

and TW. However, due to the contracted nature of

68 H. W. PALM & I. BEVERIDGE

the scolex of the current specimen, values for

scolex and bulb proportions as well as the tentacle

sheaths differed, indicating the variability of these

characters due to the degree of contraction of the

scolex. The present finding represents a new host

and locality record for M. beveridgei.

Mixonybelinia cribbi sp. nov.

(Figs 42-44)

Types

Holotype, postlarva from Platycephalus

arenarius Ramsay & Ogilby, 1886, Heron Island,

Queensland, coll. J. Reddick, Jan. 1997 (QM G

218047).

Material examined

Types.

Description

Scolex craspedote, SL = 2920 (Fig. 42);

SW = 2200; pbo = 1700; pv = 1020; pb = 730;

ppb = 220; vel = 760; app = 780; BL = 615 (600—

630); BW = 232 (220-240); BR = 2.7:1;

SP = 2.3:1.4:1. Tentacle sheaths short;

TSW = 50-65. Prebulbar organs absent, muscular

rings around basal part of tentacle sheaths not

seen. Retractor muscles originate in basal part of

bulbs. Basal tentacular swelling absent; TW

basal = 70-75, TW _ metabasal = 65-75.

Homeoacanthous, heteromorphous metabasal

armature consisting of rosethorn-shaped hooks on

bothnidial and more slender hooks with stout base

on antibothridial surface (Fig. 43); characteristic

homeomorphous basal armature (Fig. 44)

consisting of about 4 rows of slender hooks with

enlarged base and strongly recurved at tip

(L = 12.5-17.5, B = 10.0-15.0 to 8.8-11.3).

Metabasal hooks strongly recurved, rosethorn-

shaped with large base on antibothridial surface:

L= 20.0—22.5, B = 18.8—21.3; slender falcate with

stout base along bothridial surface: L = 21.3—23.7,

B = 15.0-17.5; hsr = 6-7.

Remarks

M. cribbi sp. nov. resembles M. beveridgei in

scolex size as well as in tentacular armature. Both

species have a homeomorphous basal and a

heteromorphous metabasal armature as well as a

large scolex about 3 mm in length. However, M.

cribbi sp. nov. is clearly distinguishable from M.

beveridgei by having only 4 rows of basal hooks

compared with 6 in M. beveridgei. In addition, the

hooks have a distinctly enlarged base in M. cribbi

and all hooks are smaller (20-24 in M. cribbi

compared with 46-68 in M. beveridgei, see Palm

et al. 1997). The new species was named after Dr

T. H. Cribb, Department of Parasitology,

University of Queensland, Australia.

Mixonybelinia edwinlintoni (Dollfus, 1960)

(Figs 45-53)

Material examined

From Sphyrna lewini (Griffith & Smith, 1843):

10 adults from stomach, Flat Top Island,

Queensland, coll. B. G. Robertson, 23.x.1985

(SAM AHC 28324); 1 adult, Geraldton, Western

Australia, coll. B. G. Robertson, 27.xi.1986 (SAM

AHC 28323).

From Rhynchobatus djiddensis (Forsskal,

1775): 9 postlarvae from spiral valve, Moreton

Bay, Queensland, coll. S. Butler, 17.iv.1980 (QM

G 218048-218060).

From Carcharhinus melanopterus (Quoy &

Gaimard, 1824): 1 postlarva, (QM G 4813).

Description

Adults: SL = 1780, 1620 (Fig. 45); SW = 1210,

1160; pbo = 1000, 1000; pv = 760, 690; pb = 620,

530; ppb = 60, 30; vel = 340, 330; BL = 585, 520,

520, 530, 490, within single specimen 580-600;

BW = 235 (230-240); BR = 2.5:1; SP = 1.6:1.2:1.

Tentacles robust; tentacle sheaths straight,

TSW = 50-90. Prebulbar organs absent, muscular

rings around basal part of tentacle sheaths visible

in some specimens. Retractor muscles originate in

basal part of bulbs: basal tentacular swelling

absent; TW basal = 60-65; TW metabasal = 70-

75. Metabasal armature homeoacanthous,

heteromorphous (Fig. 46); characteristic

homeomorphous basal armature consisting of

about 10 rows of slender falciform hooks (Fig.

46). Metabasal hooks strongly recurved along

antibothridial surface: L = 20.0-25.0; B = 17.8—

21.3; slender falcate hooks along bothridial

surface: L = 32.5-35.0; B = 15.0-17.5. Basal

hook size: L = 17.5-20.0, B = 10.0-13.8; hsr

basal: 7-8; hsr metabasal: 8-9.

Largest cestode 53 mm, maximum width 880;

560 segments; strobila craspedote, velum

irregularly scalloped, up to 50 wide; pre-mature

(no ovary) segments (Fig. 47) wider than long,

100 x 1020 to 120 x 1020. Genital pores

submarginal, in first third of segment; alternate

irregularly. Cirrus sac elongate, 280-330 x 45-60

(n=5) in mature segments, length:width ratio

5.7:1 (4.7-7.1:1), distal pole directed

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 69

= : ‘j Y “\

\ Sees

SLR RUS

4 Coarse aks RSE

—_

FIGURES 45-47. Mixonybelinia edwinlintoni (Dollfus, 1960) from Sphyrna lewini. SAM AHC 28324. 45. Scolex.

46. Basal and metabasal armature. 47. Pre-mature proglottid. Scale bars: Figure 45, 150 pm; Figure 46, 25 um;

Figure 47, 60 um.

anteromedially, nearly reaching anterior end of

segment; cirrus unarmed; internal and external

seminal vesicles absent. Vas deferens coiled,

extends to midline of segment, then posteriorly

towards female genital complex. Testes in two

groups, not confluent posterior to ovary, 80

(poral) and 116 (antiporal) (estimated), 30-40 in

size (central), smallest testes at margin of medulla,

external to osmoregulatory canal (15-25),

distributed in 2—3 layers, extend anterior to cirrus

sac; seminal receptacle present. Small vitelline

follicles encircle medulla, between 24 longitudinal

muscle bands. Osmoregulatory canals 5 and 20 in

diameter.

70 H. W. PALM & I. BEVERIDGE

Hor,

wee:

GES

SRe2277

SD 19 GO,

Abs —_

ORS

FIGURES 48-52. Mixonybelinia edwinlintoni (Dollfus, 1960) from Rhynchobatus djiddensis. QM G 218048-60.

48. Scolex. 49. Basal armature. 50. Metabasal armature, 20 rows from the base. 51-52. Bulb with tentacle sheath.

Scale bars: Figure 48, 150 um; Figures 49-50, 25 um; Figures 51-52, 50 um. FIGURE 53. M. edwinlintoni from

Carcharhinus melanopterus. QM G 4813. Scolex. Scale bar: 100 um. FIGURE 54. Kotorella pronosoma (Stossich,

1901) from Dasyatis fluviorum. QM G 218063. Mature proglottid. Scale bar: 50 ym.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 71

Postlarvae (from R. djiddensis): SL = 1600,

1500 (Fig. 48); SW = 1000, 1180; pbo = 1020,

950; pv = 680, 600; pb = 420, 470; ppb = 140,

120; vel = 280, 250; app= 280, 250; BL = 390

(370-410), 430 (410-440); BW = 188 (180-190),

163 (150-180); BR = 2.1:1, 2.6:1; SP = 2.4:1.6:1;

2.0:1.3:1. Tentacle sheaths straight, TSW = 40-

85. Prebulbar organs absent, muscular rings and

thickenings around basal part of tentacle sheaths

visible in some specimens (Figs 51, 52). Retractor

muscles originate in basal part of bulbs (Fig. 48);

basal tentacular swelling absent; TW basal = 80,

80; TW metabasal = 70, 80. Homeomorphous

basal armature of about 10 rows of slender

falciform hooks (Fig. 49), metabasal armature

homeoacanthous, heteromorphous (Fig. 50).

Metabasal hooks strongly recurved along

antibothridial surface: L = 22.5—25.0; B = 17.5-

20.0; slender falcate hooks along bothridial

surface: L = 23.8—26.3; B = 12.5-15.0. Basal

hook size: L = 17.5-20.0, B = 12.5—13.8; hsr

basal: 7-8; hsr metabasal: 8-9.

Postlarva (from C. melanopterus): SL = 1648

(Fig. 53); SW = 848; pbo = 912; pv = 768;

pb = 432; ppb = 96; app = 320; vel = 240;

BL = 429 (411-443); BW =171 (158-177);

BR = 2.5:1; SP = 2.1:1.8:1. Tentacles massive

(TL = 630), TSW = 68-70; TW basal 77-80, TW

metabasal 85-89, basal tentacular swelling absent.

Prebulbar organs absent; muscular rings around

basal part of tentacle sheaths not visible; retractor

muscles originate at base of bulbs. Tentacular

armature homeoacanthous, heteromorphous;

characteristic basal armature absent. Hooks

diminish in size towards base; hook shape varies

from compact rosethorn-shaped (antibothridial) to

slender rosethorn-shaped hooks with stout base

(bothridial). Hook size, metabasal, rows 16-17,

L = 23.7-25.2; B = 14.2-15.8 (antibothridial) and

L = 25.2—26.8; B =9.5—-12.0 (bothridial); basal

L = 17.4-19.0; B = 14.2-15.8 (antibothridial) and

L = 17.4-19.0; B = 12.6-14.2 (bothridial);

hsr = 7-8.

Remarks

Palm and Walter (2000) redescribed

Mixonybelinia edwinlintoni (Dollfus, 1960)

illustrating a distinctly different basal and

metabasal armature within the species. While the

metabasal armature has different hooks on the

bothridial and antibothridial tentacle surface, the

hooks of the basal armature appear to be

homeomorphous. The arrangement of the hooks,

together with their characteristic shape, and the

scolex shape, indicate that the present specimens

belong to M edwinlintoni. Rhynchobatus

djiddensis and Carcharhinus melanopterus

represent two new hosts for the species.

Prior to the re-description of M. edwinlintoni,

Palm (1999) erected Heteronybelinia

heteromorphi, a species which is also very similar

to the present material. In H. heteromorphi the

basal armature is heteromorphous, while in M.

edwinlintoni it is homeomorphous. In addition,

the type material of H. heteromorphi was

described as being acraspedote while M.

edwinlintoni is craspedote (Fig. 47). Interestingly,

both species occur within the same host genus,

Sphyrna. Other characters such as the massive

scolex form, the heteromorphous metabasal

armature, and the testis sizes are similar in both

species. The type material of H. heteromorphi

needs to be re-examined to confirm the

differences between the species.

Mixonybelinia southwelli (Palm & Walter,

1999)

Material examined

From Makaira indica (Cuvier, 1832): 1

postlarva, Cape Bowling Green, Queensland, coll.

P. Speare, 18.ix.1987 (QM G 218061).

From Chaerodon venustus (DeVis, 1885): 1

postlarva, Heron Island, Queensland, coll. R.

Bray, 30.1.1991 (QM G 218062).

Description

SL = 2000, 1550; SW=1050, 690;

pbo = 1030, 890; pv = 850, 660; pb = 440, 590;

ppb = 40; vel = 450, 220; app = 670, 270;

BL = 411 (400-425), 548 (520-580); BW = 153

(140-165), 153 (140-170); BR =2.7:1, 3.6:1;

SP = 2.3:1.9:1, 1.5:1.1:1. Tentacle sheaths

spirally coiled; TSW = 55-60, 50-60. Prebulbar

organs absent, muscular rings around the basal

part of tentacle sheaths not seen; retractor

muscles originate in basal part of bulbs;

tentacular swelling absent; TW basal = 50.0—

55.0, 47.5-55.0, TW metabasal = 35-47, 30-—

35. Armature homeoacanthous,

heteromorphous; characteristic basal armature

consisting of about 14-16 rows. Antibothridial

metabasal hooks massive, rosethorn-shaped,

L = 15.0-17.5, 15.0-16.3, B = 13.7-15.0, 15.0-

17.5; bothridial hooks more slender and slightly

curved with stout base, L = 18.7-20.0, 17.5—

20.0, B = 11.3-12.5, 11.3-13.8. Basal armature

homeomorphous, hooks falciform with stout

base and strongly recurved tip, L = 15.0—22.5,

72 H. W. PALM & I. BEVERIDGE

18.8-21.3; B = 7.5-8.7, 7.0-8.0; hsr basal = 6—

7, hsr metabasal = 5-6.

Remarks

The postlarva from Chaerodon venustus,

collected at Heron Island, is similar in scolex size

and proportions, armature pattern and hook sizes

to those described for M. southwelli by Palm and

Walter (1999). Differences in scolex

measurements of the specimen from Makaira

indica are probably due to the shrunken condition

of that specimen. The change in armature pattern

from falciform basal hooks to heteromorphous

rosethorn-shaped metabasal hooks is characteristic

for M. southwelli. The distribution of the species

is extended to eastern Australian waters, and

Makaira indica and Chaerodon venustus represent

new hosts for the species.

Genus Kotorella Euzet & Radujkovic, 1989

Kotorella pronosoma (Stossich, 1901)

(Fig. 54)

Material examined

From Dasyatis fluviorum Ogilby, 1908: 2

adults, Moreton Bay, Queensland, coll. S. Butler,

26.ix.1980 (QM G 218063-218064).

Description

SL = 570, 660; SW = 200, 200; pbo = 400, 390;

pv = 385, 510; pb = 80, 85; vel=110, 80;

BL=71 (70-73), 78 (71-88); BW = 39.4 (37.5—

40.0), 40.0 (35.3-45.0); BR = 1.8:1, 2.0:1;

SP = 5.0:4.8:1, 4.6:6.0:1. TW basal = 17.5, 22.5.

Basal tentacular swelling absent. Tentacle sheaths

straight; TSW = 11.3—15.0, 5.5—15.0. Prebulbar

organs and muscular rings around basal part of

tentacle sheaths absent; retractor muscles originate

in basal part of bulbs. Tentacular armature

homeoacanthous, heteromorphous; basal hooks on

bothridial surface L = 6.3-7.5, 6.3-7.5, B =5.0-

6.0, 5.0-6.0, smallest hooks on antibothridial

surface L=2, increasing in size towards

metabasal region. Small, incomplete worms to

15 mm long, maximum width 570; 90 segments.

Strobila acraspedote; pre-mature segments 30 x

110 — 260 x 360, mature segments longer than

wide or wider than long, 480 x 430 to 550 x 620

(Fig. 54). In second specimen, mature segments

500 x 440 to 550 x 650, and pre-gravid (with

some eggs) segments 320 x 710 to 750 x 620.

Genital pores submarginal, 210 from anterior

margin; alternate irregularly. Cirrus sac elongate,

150-250 x 30-60 (n=5) in mature segments,

length:width ratio 4.4:1 (4.0-5.0:1); cirrus

unarmed; internal and external seminal vesicles

absent. Vas deferens in large coils, extends to

midline of segment, then posteriorly towards

female genital complex. Testes 86 (81-100) in

number, 45-85 in size (central), smallest testes

20-50 (peripheral), distributed in single layer,

confluent posterior and anterior to ovary. Ovary

median, ovarian lobes 110-200 x 100-150;

Mehlis’ gland small, situated between ovarian

lobes. No vitelline follicles in centre of the

segment, follicles 33 (25-45) in diameter. No

uterus observed, spherical eggs appear in

parenchyma, egg diameter 22 (15-40).

Remarks

The morphology of Kotorella pronosoma

(Stossich, 1901) is summarised by Euzet and

Radujkovic (1989), Palm and Walter (1999) and

Palm and Overstreet (2000). The present

specimens from Moreton Bay correspond in

scolex size and proportions as well as in the

tentacular armature with the other described

specimens. The sole difference noted was in testis

number which exceeded 80 per segment in the

specimens described. The species has previously

been found in rays such as Aetobatus narinari

(Euphrasen, 1790 ) (see Palm & Overstreet 2000),

Himantura imbricata (Bloch & Schneider, 1801),

Rhynchobatus djiddensis (see Shipley & Hornell

1906; Palm & Walter 1999) and Dasyatis

pastinaca (Linnaeus, 1758) (see Euzet &

Radujkovic 1989; Palm & Walter 2000). Dasyatis

fluviorum represents a new host record, and the

range of distribution is extended to eastern

Australian waters. The present finding suggests a

cosmopolitan distribution for the species.

Previously, the nearest known occurrence of this

cestode to Australia was from the Java Sea under

the name Nybelinia narinari (MacCallum, 1917),

now a synonym of Kotorella pronosoma (see

Palm & Overstreet 2000).

Kotorelliella gen. nov.

Diagnosis: Tentacles elongate, slender; retractor

muscle originates at base of bulbs. Metabasal

tentacular armature homeoacanthous,

heteromorphous; hooks on bothridial surface

uncinate becoming slender towards antibothridial

surface. Basal hooks heteromorphous;

characteristic basal armature present, arranged in

a heteroacanthous atypica pattern with

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 73

FIGURES 55-61. Kotorelliella jonesi gen. et sp. nov. from Taeniura lymma. Holotype, QM G 218065. 55. Scolex.

56-57. Metabasal and apical armature, external view. 58. Basal armature, bothridial view. 59. Basal armature,

antibothridial view. 60-61. Basal towards metabasal armature, external view. Note the interpolated hooks as in Fig.

61. Scale bars: Figure 55, 150 um; Figures 56-61, 10 um.

74 H. W. PALM & I. BEVERIDGE

interspersed hooks on the antibothridial tentacle

surface. Hooks solid. Strobila unknown.

Type-species: Ko. jonesi sp. nov.

Kotorelliella jonesi sp. nov.

(Figs 55-61)

Types

Holotype, 1 postlarva from the spiral valve of

Taeniura lymma (Forsskal, 1775), Heron Island,

Queensland, coll. I. Beveridge & M. K. Jones,

11.v11.1998 (QM G 218065).

Material examined

Holotype.

Description

Scolex compact, 4 elongated bothridia, with

hook-like microtriches along bothridial borders.

Posterior margins of bothridia free, not fused with

peduncle. SL= 1910 (Fig. 55); SW = 390;

pbo = 800; pv = 1140; pb = 260; vel = 460;

BL = 259 (250-265); BW = 82.5 (80-85);

BR = 3.1:1; SP =3.1:4.4:1. Tentacles long and

slender; TW basal and metabasal = 21.2—23.8;

TW apical = 17.5—20; basal tentacular swelling

absent. Tentacle sheaths straight; TSW = 15.0—

17.7. Prebulbar organs and muscular rings around

basal part of tentacle sheaths absent; retractor

muscles originate in basal part of bulbs. Metabasal

(Fig. 56) and apical (Fig. 57) armature

homeoacanthous, heteromorphous. Rosethorn-

shaped hooks with anterior extension of base

cover bothridial tentacle surface (L = 13.5—15.0,

B = 7.5—10.0); antibothridial surface with slender

spiniform hooks, recurved at tip (L = 15.0-17.5,

B = 5.0-6.3); hooks decrease in size towards

apical armature; hook shape remains constant

(bothridial: L = 11.3-13.7, B = 10.0-11.2;

antibothridial: L = 12.5-13.7, B = 4.5-S.5). Basal

armature (Figs 58-61) with additional hooks

interspersed on antibothridial surface; hook

pattern heteroacanthous atypica, consisting of

about 5 rows of hooks on bothridial (Fig. 59) and

9-10 rows on antibothridial (Fig. 58) tentacle

surface; basal hooks on bothridial surface

L = 6.3-10.0, B = 6.3-8.8, on antibothridial

surface L = 2.0-10.0, B = 1.5-3.8, continuously

increasing in size from base of tentacle towards

metabasal armature (Figs 60-61).

Remarks

The holotype of Kotorelliella jonesi gen. et sp.

nov. has a unique form of armature within the

Tentaculariidae, changing from a

homeoacanthous, heteromorphous hook pattern

with rosethorn-shaped hooks in the metabasal

armature to a heteroacanthous atypica hook

pattern in the basal armature. The size of hooks in

the basal and apical regions is smaller than in the

metabasal region.

On the basis of the scolex morphology, Ko.

jonesi sp. nov. appears to be similar to Kotorella

pronosoma. Both species have an elongated

scolex, four elongated and slender bothridia with

free margins, and short oval bulbs. In both

species, the metabasal armature is

heteromorphous, with hooks changing from

uncinate on the bothridial surface to elongate on

the antibothridial surface. Both species have a

band of hook-like microtriches along bothridial

borders, which are clearly visible using light

microscopy. Such a microthrix pattern is

characteristic for tentaculariid trypanorhynch

cestodes.

The basal armature of K. pronosoma is

homeoacanthous, but that of Ko. jonesi sp. nov. is

unusual, having additional hooks interspersed

between the oblique spirals of hooks. Thus, the

basal tentacular armature is heteroacanthous

atypica. The basal region is homeoacanthous in

Nybelinia, Heteronybelinia and Mixonybelinia,

but in Tentacularia, the bothridial surfaces of the

tentacle have extra rows of small hooks (see

Beveridge & Campbell 1996). The unique feature

of the tentacular armature suggests that the

erection of a new genus is justified, even though

it is known only from the larval stage.

The new species was named after one of the

collectors, Dr M. K. Jones, from the Centre for

Microscopy and Microanalysis, University of

Queensland, Australia.

Unidentified material

The following additional material, some of

which represent new host records, was examined

but could not be assigned to a species:

Nybelinia sp. from the spiral valve of

Rhynchobatus djiddensis, Flat Top Island,

Queensland, coll. B. G. Robertson, 25.x.1985

(SAM AHC 28325)

Nybelinia sp. from the spiral valve of

Notorhynchus cepedianus (Péron, 1807), south

coast Kangaroo Island, South Australia, coll. B.

G. Robertson, 10.ii.1985 (SAM AHC 28327)

Nybelinia sp. from the stomach of

Notorhynchus cepedianus, Young Rocks,

Kangaroo Island, South Australia, coll. B. G.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 15

Robertson, 23.v.1985 (SAM AHC 21354, 28328)

Heteronybelinia sp. from the spiral valve of

Carcharhinus amblyrhynchoides, Tommy Cut

Mouth, Northern Territory, coll. B. G. Robertson,

19.ix.1986 (SAM AHC 18326).

DISCUSSION

The present study provides an overview for the

first time of the tentaculariid trypanorhynch fauna

from the Australian region. Three new species are

added to the genus WNybelinia, two to

Heteronybelinia and one to Mixonybelinia, with

the latter two genera, as well as Kotorella,

reported from the Australian region for the first

time. In addition, a new genus Kotorelliella gen.

nov., is erected, which is characterised by a

homeoacanthous, heteromorphous metabasal

armature and a heteroacanthous basal armature.

Other features such as the elongated, widely

62.

wy AN

: {TKN Ne

iS ‘ = iar

NIZA

I-ANICAA

TZN \ZZR

spaced bothridia and the short bulbs appear

similar to Kotorella.

The tentaculariid trypanorhynch fauna in

Australian waters is relatively species rich with 22

(48 %) of the total of 46 known species occurring

in these waters. Several of the new species may be

endemic. However, the distribution of many

species is still imperfectly known and a number of

species previously thought to be endemic to a

specific region have subsequently proven to be

cosmopolitan (Palm et al. 1997; Palm 1999; Palm

& Walter 1999, 2000). Reasons for broad

distributions within tentaculariid trypanorhynchs

may relate to life-cycle patterns, including widely

distributed fish intermediate and elasmobranch

definitive hosts. Another reason might be the low

host specificity of the parasites, particularly in the

intermediate host (Palm et al. 1997; Palm &

Walter 2000).

It is difficult to clearly distinguish between

FIGURES 62-63. Schematic drawing of scolex of Heteronybelinia pseudorobusta sp. nov. from Lepidotrigla

modesta, illustrating the arrangement of the different tentacle surfaces within tentaculariid trypanorhynchs. 62.

View from bothridial surface, with bothridial borders merging at the apex of the scolex. 63. View from external

surface, with bothridial borders widely spaced. Abbreviations, AB = antibothridial view, B = bothridial view,

Bo = bothridia, E = external view, I = internal view.

76 H. W. PALM & I. BEVERIDGE

some tentaculariid species solely on the basis of

the tentacular armature (eg N. strongyla), and

additional taxonomic characters are required. To

date, the strobilae of only a few tentaculariid

species have been described in detail. The

present study demonstrates that, apart from the

number and size of different genital structures,

the cirrus sac length:width ratio might be a

useful character as, for example, in H. australis

sp. nov. and N. hemipristis sp. nov. The

position of the anterior end of the cirrus sac and

the distance of the genital pore from the

anterior end of the segment, as well as the

number of layers of testis, whether the testes

are confluent posterior to the ovary, and the

number of testes anterior to the cirrus sac

appear to be useful taxonomic features. Further

species descriptions are needed to show whether

or not these characters are variable, as is

observed in a number of scolex characters, or

whether they can be used more widely in

tentaculariid systematics.

During this and previous studies, it became

evident that the descriptions of the different

tentacular surfaces of trypanorhynch cestodes

used by Dollfus (1942) and Campbell and

Beveridge (1994) may be difficult to apply to

larval tentaculariids. In many cases it was hard

to identify the various tentacle surfaces. For a

better understanding of the position of the four

tentacles within tentaculariid trypanorhynchs,

the following two schematic illustrations are

given (Figs 62, 63) as they may help to

standardise the orientation of the different

tentacle surfaces within tentaculariid

trypanorhynchs. The determination of tentacle

orientation is based upon whether the

bothridial borders merge at the apex of the

scolex (Fig. 62) or whether they are widely

spaced (Fig. 63). The former figure shows the

bothridial and the latter the external view of

the tentacle surfaces.

The description of Kotorelliella jonesi gen. et

sp. nov. is of particular interest since its armature

comprises both homeomorphous and

heteromorphous components (Campbell &

Beveridge 1994; Palm 1995, 1997). The species

seems to be closely related to Kotorella

Pronosoma, which has a similar scolex

morphology and metabasal armature. Both species

have characteristic microtriches along the

bothridial borders, which appear similar to those

of Tentacularia coryphaenae and species of

Nybelinia (see Palm & Overstreet 2000).

Although Beveridge et al. (1999) could not align

Kotorella with the other tentaculariid genera using

cladistic analyses, we consider that Kotorella

pronosoma and Kotorelliella jonesi belong to the

Tentaculariidae since they possess a

homeoacanthous metabasal armature and, more

importantly, bands of hook-like microtriches

along the bothridial borders. The latter character

has as yet only been described for tentaculariid

trypanorhynchs.

Following the classification of Dollfus (1942),

only the metabasal armature is considered in

classifying the tentacular armature. However, in

more recent classifications, trypanorhynchs with a

distinct basal armature such as Mixodigma

leptaleum Dailey & Vogelbein, 1974,

Paroncomegas araya (Woodland, 1934) and

Mixonybelinia beveridgei (Palm, Walter,

Schwerdtfeger & Reimer, 1997) have been placed

in distinctive genera based in part on the basal

armature (Campbell & Beveridge 1994; Campbell

et al. 1999; Palm 1999). Kotorelliella jonesi has

thus been placed in a new genus. Interestingly, K.

jonesi represents a _ transition from

homeoacanthous towards heteroacanthous

armature as suggested by Campbell and Beveridge

(1994, p. 57, lines 7-11).

In summary, tentaculariid trypanorhynchs show

considerable morphological variability, as can be

seen by differences in scolex morphology between

the genera Nybelinia, Heteronybelinia and

Mixonybelinia, and between Kotorella and

Kotorelliella. The tentacular armature also differs

between the species, varying from homeoacanthous

and homeomorphous (Nybelinia, Mixonybelinia) to

homeoacanthous and heteromorphous

(Heteronybelinia, Kotorella, Mixonybelinia)

patterns. Kotorelliella has a homeoacanthous

metabasal and a heteroacanthous basal hook

pattern. Additional methods, such as molecular

genetic studies, might help to resolve not only

species relationships but also the phylogenetic

relationships of the species and species groups

within this large trypanorhynch family.

ACKNOWLEDGMENTS

Collecting was supported financially by the

Australian Biological Resources Study and the

Australian Research Council. Financial support was

also provided by the German Research Council DFG

PA 664/3-1 and 3-2. We wish to thank R. A. Bray, S.

Butler, T. H. Cribb , M. K. Jones, R. Martin, M.

O’Callaghan, R. Norman, J. C. Pearson, S. Pichelin, J.

Reddich, B. G. Robertson , K. Sewell, P. Speare and J.

Stevens for their contributions to this study.

TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 77

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TWO NEW SPECIES OF THE GENUS LEPANUS BALTHASAR FROM

SOUTH AUSTRALIA (COLEOPTERA : SCARABAEIDAE)

E. G. MATTHEWS & T. A. WEIR

Summary

Ball-rolling dung beetles of the tribe Scarabaeini are recorded for the first time from South

Australia. The new species Lepanus penelopae and L. loftyensis are described from southern Eyre

Peninsula and the Mount Lofty Block respectively, and are compared with other species of the

genus. There are brief remarks comparing Lepanus Balthasar with the related genera Sauvagesinella

Paulian and Aptenocanthon Matthews.

TWO NEW SPECIES OF THE GENUS LEPANUS BALTHASAR FROM SOUTH AUSTRALIA

(COLEOPTERA: SCARABAEIDAE)

E. G. MATTHEWS & T. A. WEIR

MATTHEWS, E. G. & WEIR, T. A. 2002. Two new species of the genus Lepanus Balthasar

from South Australia (Coleoptera: Scarabaeidae). Records of the South Australian Museum.

35(1): 79-84.

Ball-rolling dung beetles of the tribe Scarabaeini are recorded for the first time from South

Australia. The new species Lepanus penelopae and L. loftyensis are described from southern

Eyre Peninsula and the Mount Lofty Block respectively, and are compared with other species

of the genus. There are brief remarks comparing Lepanus Balthasar with the related genera

Sauvagesinella Paulian and Aptenocanthon Matthews.

E. G. Matthews, South Australian Museum, North Terrace, Adelaide, South Australia S000;

and T. A. Weir, CSIRO Division of Entomology, P.O. Box 1700, Canberra, Australian Capital

Territory 2601. Manuscript received 8 August 2001.

At the time of the revision by Matthews (1974)

of the Australian Scarabaeini, generally known as

ball-rolling dung beetles, it was believed that this

group was absent from South Australia, although

it was known from Victoria and Western

Australia. However, in 1979 P. Greenslade

obtained specimens of an undescribed species of

the genus Lepanus Balthasar, which belongs to

this tribe, in the Marble Range of southern Eyre

Peninsula; and in 1982 another undescribed

species of the same genus was collected during

extensive pitfall trapping undertaken by J. and P.

Greenslade in Kuitpo Kyeema Forest, southern

Mount Lofty Ranges, in connection with a project

to determine the effects of fire on the forest-floor

fauna. Then in 1999 and 2000, more specimens of

the second species were picked up by members of

the Biological Survey of South Australia in the

area of Mount Remarkable at the extreme northern

end of the Mount Lofty Block environmental

province, as well as near the Barossa Valley, using

unbaited pitfall traps. Altogether, the two species

are now known from five collection localities

(Fig. 1), all situated on low mountain ranges at

altitudes from 300 to just over 500 m. Kangaroo

Island is part of the Mount Lofty Block but pitfall

traps baited with human faeces, set throughout the

island by one of the authors (EGM) in November

1990, failed to collect any Lepanus.

Lepanus is known from New Guinea, where it

was first described, and from densely vegetated

mesic and humid habitats along the northern,

eastern and southern coasts of Australia as far

west as the tingle forest near Walpole, Western

Australia (Matthews 1974). The habitat is leaf

litter, and food, where known, consists of

vertebrate faecal matter. The descriptions below

bring the total number of described Australian

species of Lepanus to 23, but several undescribed

species have been collected in recent years in

eastern montane forests (R. Storey, pers. comm.).

Specimens are deposited in the South

Australian Museum, Adelaide (SAMA); the

Australian National Insect Collection, Canberra

(ANIC); the Queensland Museum (QMBA); and

the Queensland Department of Primary Industries

collection at Mareeba (DPIM).

All figures except the map are by the senior

author.

DESCRIPTIONS OF NEw SPECIES

Lepanus penelopae sp. nov.

(Figs 1, 2, 3, 5, 7, 9, 12)

Holotype

‘S. Aust. Eyre Pen. Marble Rge. Dense

broombush. Pitfall. 4.10.79. P.J.M. Greenslade’,

male, SAMA.

Paratypes

Same data as holotype, 1 male and 5 females,

ANIC, SAMA.

Description

Body uniformly piceous and nitid, legs rufous.

Total length 5.0-6.0 mm. Maximum width across

80 E. G. MATTHEWS & T. A. WEIR

FIGURE 2. Lepanus penelopae male, habitus. Scale line

1 mm.

elytra 3.4—-3.6 mm. Head — Dorsal surface even,

very densely punctate with small shallow

punctures not running together, some bearing

short, very fine recumbent setae. Dorsal part of

eye small, its maximum width contained about 25

times in interocular distance. Prothorax — Anterior

angles subquadrate. Sides of pronotum rounded,

widest at broadly rounded posterior angles. Dorsal

surface very densely and uniformly finely

punctate, punctures separated by distances equal

to a little more than their diameter, glabrous.

Elytra — Striae shallowly impressed with regularly

spaced moderately deep punctures separated by 2-

4 diameters. Intervals nearly flat, smooth with

very slightly uneven surface, glabrous, with

minute punctures. Hind wings — Absent. Sterna —

Meso- and metasterna impunctate medially with

large shallow cicatricose punctures laterally,

glabrous. Legs — Protibia with three large teeth on

outer edge, which is serrate between teeth and

proximal to them. Claws (Fig. 7) a little expanded

basally but not distinctly angulate or dentate.

Abdomen — Pygidium (Fig. 9) with a very deep

TWO NEW SPECIES OF LEPANUS BALTHASAR 81

FIGURES 3-6. Tibiae of males in dorsal view. 3, L. penelopae, left protibia; 4, L. loftyensis, right protibia; 5, L.

penelopae, right metatibia; 6, L. loftyensis, right metatibia. b, brush of stiff setae; c, comb of flattened bristles.

Scale line 1 mm.

basal groove across whole width of disc, groove

narrower and shallower in middle, surface inside

it transversely striated. Disc very finely punctate,

glabrous. Aedeagus as in Fig. 12. Sexual

dimorphism — Male with rounded expansion of

inner apical end of protibia bearing apical comb

of short wide flattened bristles (Fig. 3, c) and on

lower surface a brush of long setae which is

directed outwardly parallel to surface, extending

beyond edge of comb (Fig. 3, b). Metatibia of

male (Fig. 5) a little expanded at inner apical edge

and bearing small apical tooth there.

Remarks

In the key to the species of Lepanus published

by Matthews (1974), L. penelopae will go to

couplet 16 because it has three protibial teeth, and

then to couplet 18 because it has a transverse

groove on the pygidium and glabrous dorsal

surfaces. However, it does not have dentate claws

as also required by this couplet. It can then be

forced through couplets 19 and 20 to end up with

L. illawarrensis Matthews of New South Wales,

which appears to be the most closely related

species, sharing many aspects of structure and

secondary sexual characters. L. penelopae differs

from L. illawarrensis in having simple claws, non-

geminate elytral striae with much deeper strial

punctures, the pygidial groove more strongly

narrowed in the middle, the basal ridge of the

pygidium straight in the middle, no trace of a

median tubercle on the metasternum of the male,

male hind tibia not twisted, and somewhat

differently shaped aedeagal parameres.

Etymology

This species is named after Penelope

Greenslade, who collected it as well as many of

the other specimens of Lepanus described in this

paper.

Lepanus loftyensis sp. nov.

(Figs 1, 4, 6, 8, 10, 11)

Holotype

‘S. AUST. Pitfalls Kuitpo Kyeema Forest 15th

March, 1982 P. Greenslade’, male, SAMA.

Paratypes

Same data as holotype, 12 males, 4 females,

ANIC, SAMA. ‘S. Aust. Deep Ck Boat Hbr Lane

82 E.G. MATTHEWS & T. A. WEIR

FIGURES 7-9. 7, L. penelopae, claw segment of

metatarsus; 8, ditto, L. loftyensis. Scale line 0.25 mm;

9, L. penelopae, pygidium of male. Scale line 1 mm.

turnoff pitfalls in heath 1-8 Dec 1983 P.

Greenslade’, 1 female, SAMA. ‘S. AUST. Mt

Remarkable NP 11 km SSW Wilmington

32° 44°56”S 138°04’03”E 20-25 Nov 99 pitfall

Flinders Ra. MAMO00401’, 5 females, ANIC,

SAMA. ‘S. AUS. 11 km SE Whispering Wall

(near Para Wirra) 34°34’54”S 138°55’12”E

27-10-00 SMLR Survey BAR 03201’, 17 males,

31 females, SAMA, QMBA, DPIM.

Description

Body uniformly fuscous to piceous, legs

rufous. Total length 3.4-5.5 mm, maximum width

across elytra 2.3-3.6 mm. Head — Surface even,

very densely punctate with shallow coarse

punctures running together in irregular transverse

rows, bearing small recumbent setae. Dorsal part

of eye small, slit-like, its maximum width

contained about 30 times in interocular distance.

Prothorax — Anterior angles obtuse. Sides of

pronotum widest about one-quarter of the distance

behind anterior angles, thereafter slightly

converging or subparallel to broadly rounded

posterior angles. Dorsal surface very densely and

uniformly punctate, punctures separated by

distances equal to less than their diameter, bearing

very fine, short, fully recumbent setae. Elytra —

Striae very superficial, geminate with slightly

crenulate edges, impunctate. Discal intervals flat,

in same plane as striae, with more or less undulate

surface, finely shagreened, with a row of

punctures bearing recumbent setae along edges of

each interval, an additional row usually present in

middle of intervals near base. Hind wings —

Atrophied, represented by short stubs. Sterna —

Mesosternum with shallow punctures on posterior

half. Metasternum entirely densely punctate with

shallow punctures laterally bearing short, fine

recumbent setae. Legs — Protibia (Fig. 4) with 3

large teeth on outer edge, which is serrate between

teeth and proximal to them. Claws (Fig. 8)

strongly dentate. Abdomen — Pygidial disc simple,

without groove, uniformly densely and shallowly

FIGURE 10. Lepanus loftyensis male, habitus. Scale

line 1 mm.

TWO NEW SPECIES OF LEPANUS BALTHASAR 83

punctate, punctures bearing very small recumbent

setae usually worn off. Aedeagus as in Fig. 11.

Sexual dimorphism — Male with rounded

expansion of inner apical end of protibia bearing

apical comb of short, wide, flattened bristles (Fig.

4, c) and a brush of long setae underneath which

is directed downward perpendicular to surface.

Metatibia of male (Fig. 6) with strongly recurved

apical end bearing acute inner tooth. Prothorax of

male broader anteriorly, subquadrate; that of

female slightly narrower anteriorly. Clypeal teeth

of male slightly smaller than those of female.

Remarks

In the key to species of Lepanus in Matthews

(1974) L. loftyensis will first go to couplet 16

because it has three teeth on the outer edge of the

protibia, then to couplet 17 because it has a simple

pygidium and setose dorsal surfaces, then to L.

villosus Matthews of north Queensland because

of its crenulate elytral striae and densely punctate

meso- and metasterna. It differs from L. villosus

in having superficial elytra striae, reduced hind

wings, very different parameres of the aedeagus,

and being of much larger size. Further, L.

loftyensis has the setae on the pronotum only

about half the length of those on the elytra, the

first elytral interval with a double row of seta-

bearing punctures, and the upper edges of the

epipleural carina not visible from directly above

due to the curvature of the elytra.

The designation MAM 00401 on labels of the

Mount Remarkable specimens refers to a locality

which is described in Brandle (2001) as

mountainous, on a hill crest, with loam/sand soil

and an overstorey of Allocasuarina verticillata at

a cover density of 25-50%, and at an altitude

above sea level of 430m. For the Parra Wirra

specimens the designation BAR 03201 refers to a

ridge top covered with Eucalyptus fasciculosa /

Acacia paradoxa woodland and an understorey of

Calytrix, Hakea and Xanthorrhoea, at an altitude

of 515 m (L. Queale, pers. comm.).

Etymology

The name refers to the Mount Lofty Block

environmental province in which all four

collection localities of this species are contained.

The block is Province number 3 as delineated in

the Biological Survey of South Australia, and

extends from Kangaroo Island northward to

Mount Remarkable. The biota of this province is

characterised as fully Bassian (Brandle 2001).

DISCUSSION

The existence of the species here described as

Lepanus loftyensis was briefly noted by Matthews

(1984: 6, footnote) erroneously as an undescribed

species of Sauvagesinella Paulian. Lepanus and

Sauvagesinella are closely related genera which

are not separable in the female sex, but in the

male there are several secondary sexual characters

which will distinguish them. Unlike Lepanus,

Sauvagesinella males have a prominent median

FIGURES 11 & 12. Aedeagi in right and left views. 11, L. loftyensis; 12, L. penelopae. Scale line 1 mm.

84 E. G. MATTHEWS & T. A. WEIR

tubercle on the metasternum (only a trace of a

tubercle in some Lepanus), a row of close-set

tubercles along the inner edge of the metatibia,

and a more or less compressed metafemur with a

longitudinal fold or ridge on its ventral surface.

Sauvagesinella as understood here does not occur

outside the extreme southern part of Western

Australia, where there are three species (Matthews

1974).

Storey and Monteith (2000) mention that males

of all species of Aptenocanthon Matthews, where

known, have a similar form of the fore tibial apex

with its inner angle expanded and bearing a short

brush of stiff setae bent downwards at right angles

to the upper tibial surface. This is a similar

situation to that found in the two species of

Lepanus described above (eg Fig. 3, b) and is also

known to occur in several other species of

Lepanus and all three species of Sauvagesinella.

As well as this brush, there is a comb of flattened

bristles at the protibial apex (Figs 3 and 4, c)

which occurs, to varying degrees, in all known

males of Aptenocanthon and Sauvagesinella as

well as some species of Lepanus. Clearly, these

male characters are not limited to Aptenocanthon

but their value in classification must await further

studies on the interrelationships of these genera of

Scarabaeini, as also pointed out by Storey and

Monteith (2000).

In South Australia the only other known

representatives of the dung beetle subfamily

Scarabaeinae belong to the tribes Onthophagini

(native and introduced Onthophagus Latreille),

Onitini (introduced Onitis Fabricius) and

Oniticellini (introduced Euoniticellus Janssens),

none of which make or roll food balls. All these

groups are differentiated from the tribe

Scarabaeini by their middle and hind tibiae,

which are strongly expanded apically. In

Scarabaeini these tibiae are slender and only a

little expanded, an adaptation for rolling the food

material.

REFERENCES

BRANDLE, K. 2001. A biological survey of the

Flinders Ranges, South Australia 1997-1999,

Biodiversity Survey and Monitoring, National Parks

and Wildlife, South Australia, Department for

Environment and Heritage, pp. i-xviii, 1-455.

MATTHEWS, E. G. 1974. A revision of the scarabaeine

dung beetles of Australia. II. Tribe Scarabaeini.

Australian Journal of Zoology, Supplementary Series

24: 1-211.

MATTHEWS, E. G. 1984. A guide to the genera of

beetles of South Australia. Part 3. South Australian

Museum, Special Educational Bulletin Series 6: 1-

60.

STOREY, R. I. & MONTEITH, G. B. 2000. Five new

species of Aptenocanthon Matthews (Coleoptera:

Scarabaeidae: Scarabaeinae) from tropical Australia,

with notes on distribution. Memoirs of the

Queensland Museum 46(1): 349-358.

A CHECKLIST OF CISSEIS (SENSU STRICTO) GORY & LAPORTE, 1839

(COLEOPTERA : BUPRESTIDAE : AGRILINAE)

SHELLEY BARKER

Summary

A checklist of Cisseis (s.s.) Gory & Laporte, 1839 is presented and discussed, summarising the

information available on the genus at the present time.

A CHECKLIST OF CISSEIS (SENSU STRICTO) GORY & LAPORTE, 1839

(COLEOPTERA: BUPRESTIDAE: AGRILINAE).

SHELLEY BARKER

BARKER, S. 2002. A checklist of Cisseis (sensu stricto) Gory & Laporte, 1839 (Coleoptera:

Buprestidae: Agrilinae). Records of the South Australian Museum 35(1): 85-90.

A checklist of Cisseis (s.s.) Gory & Laporte, 1839 is presented and discussed, summarising

the information available on the genus at the present time.

S. Barker, Department of Entomology, South Australian Museum, North Terrace, Adelaide,

South Australia 5000. Manuscript received 3 December 2001.

MATERIAL

Specimens examined came from the following

institutions:

AMSA - Australian Museum, Sydney

ANIC — Australian National Insect Collection,

CSIRO, Canberra

BMNH - The Natural History Museum, London

BPBM - B. P. Bishop Museum, Honolulu

HMOE - Hope Museum, Oxford

HUMB -— Humboldt University Museum, Berlin

MCSNG —- Museo Civico di Storia Naturale

Genoa, Italy

MNHN -— Museum National d’ Histoire Naturelle,

Paris

NMVA — National Museum of Victoria,

Melbourne

PMCE — National Museum of Czech Republic,

Prague

QMBA — Queensland Museum, Brisbane

RMBB - Royal Museum, Brussels

SAMA -— South Australian Museum, Adelaide.

Abbreviations:

WA — Western Australia

NWA - Kimberly area, Western Australia

Q — Queensland

NSW — New South Wales

V — Victoria

SA — South Australia

T — Tasmania

NT — Northern Territory.

INTRODUCTION

Many of the Australian buprestid genera have

not been reviewed since the death of H. J. Carter in

1941. Because of the large amount of subsequent

collecting and confusion with species

identification, the genera are in a parlous state of

order. One of worst has been Cisseis (Coleoptera:

Buprestidae: Agrilinae), last researched by Carter

(1923, 1929). With the intention of revising

Cisseis, the author commenced work on the group

in 1997. All of the readily available types, more

than half of which are housed in European

museums, have now been examined. Some of the

difficulties have been resolved and a number of

new species described (Barker 1998, 1999a, 1999b,

2001). Some species clearly did not belong in

Cisseis and these, together with nine new species,

were placed in a new genus Stanwatkinsius Barker

& Bellamy (2001). Carter (1923) did not examine

male genitalia, which the author has found to be

highly diagnostic. As a result, many of his

specimens located in various museum collections

are incorrectly identified. In order to summarise the

knowledge available on the genus, a checklist has

been constructed of valid species together with

their distributions by state or country. Types and

their locations are also listed. The following

abbreviations are used: HT = holotype; ST =

syntype; LT = lectotype; AT = allotype.

CHECKLIST OF CISSEIS (SENSU STRICTO) GORY &

LAPORTE, 1839.

aberrans Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 98. (6 HT, Woodridge, WA, H.

Demarz, SAMA I 21504). WA.

86 S. BARKER

acuducta (Kirby, 1836)(Trachys). Faun. Bor.

Amer.: 162. (6 HT, N. Scotia, BMNH).

Q, NSW, V, SA, T.

marmorea (Gory & Laporte, 1839)(Ethon).

Mon. Bupr. ii: 4. (HT, MNHN not seen)

cuprifrons Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 157. (6 HT, Australia,

Standing, BMNH)

laeta Kerremans, 1903. in Wytsman Genera

Ins. fas. 12: 227. (o HT, Australia,

Moffarts, BMNH)

adusta Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 101. (¢ HT, Lucindale, SA,

Feueheerdt, SAMA I 21505). SA.

albertisi Gestro, 1877. Ann. Mus. Civ. Stor. Nat.

Genova. 9: 357. (3 ST sex unknown,

Somerset, Australia, D’Albertis, 1/75

MCSNG; @ ST, Somerset, Australia,

RMBB). Q.

albosparsa Gory & Laporte, 1839. Monograph ii:

3. (HT, MNHN not seen). NT, Q.

cupriventris Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 161. (& HT, Australia,

Chevrolat, BMNH).

inflammata Carter, 1923. Proc. Linn. Soc.

N.S.W. 48: 167. (S HT, S. Johnstone R.,

Q, H. W. Brown AMSA K67240) (new

syn.).

aquilonia Bellamy, 1991. Tijdschr. v. Ent. 134:

171. (6 HT, BPBM not seen). Philippine

Arch.

armstrongi Barker, 2001. Trans. Roy. Soc. S.

Aust. 125: 101. (¢ HT, Bogan R., NSW,

J. Armstrong, NMVA). NSW.

augustgoerlingi Barker, 2001. Trans. Roy. Soc. S.

Aust. 125: 102. (6 HT, Marloo Stn.,

Wurarga, WA, A. Goerling, ANIC). WA.

aurocyanea Carter, 1934. Proc. Linn. Soc. N.S.W.

59: 258. (2 HT, Fletcher, Q, E. Sutton,

AMSA K67296). NSW, Q.

bedfordi Obenberger, 1923. Casopis Cs. Spol.

Entom. 32: 9. (2 HT, Cape Bedford, Q,

NMPC). Q.

brooksi Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 102. (6 HT, Mareeba, Q, J. G.

Brooks, ANIC). Q.

broomensis Barker, 2001. Trans. Roy. Soc. S.

Aust. 125: 103. (6 HT, Broome, WA, H.

W. Brown, SAMA I 21509). NWA.

browni Carter, 1934. Proc. Linn. Soc. N.S.W. 59.

258. (6 HT, Dedari, WA, H. W. Brown,

AMSA K67297). WA.

carterella Obenberger, 1935. Col. Catalogus 12:

846. (2 LT, Herberton, Q, C. J. Wild,

QMBA)(replac. name). Q.

elliptica var. frontalis Carter, 1923. Proc.

Linn. Soc. N.S.W. 48: 170. (homonym).

carteri Obenberger, 1924. Archiv. fiir Naturg. 90:

109. (3 LT, Yilgarn, WA, NMPC). WA.

chalcophora Barker, 2001. Trans. Roy. Soc. S.

Aust. 125: 103. (¢ HT, Kapunda, SA, S.

Barker, SAMA I 21507). SA.

clermonti Théry, 1945. Bull. Soc. Ent. Fr. 50: 46.

(HT, MNHN not seen). Bougainville.

corpulenta Barker, 2001. Trans. Roy. Soc. S.

Aust. 125: 104. (6 HT, Tallering stn.,

Pindar, WA, S. Barker, SAMA I 21511).

WA.

cupreicollis (Hope, 1846)(Ethon). Ann. Mag. Nat.

Hist. 17: 64. (6 HT, Moriatta (sic), SA,

HMOE no. 893). SA

aenicollis (Hope, 1846)(Ethon). Ann. Mag.

Nat. Hist. 17: 65. (2 HT, Adelaide,

HMOE no. 892 2/2).

cupreola Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 10S. (o HT, Mt Spec, Q, J. G. B.,

ANIC). SA, NSW, Q.

cupripennis (Guerin, 1836)(Buprestis). Voy.

Coquille: 65. (HT, MNHN not seen).

NSW, V.

semiscabrosa Thomson, 1879. Typi Bupr.

Mus. Thomson. App. 1A: 53. (3d HT,

Sydney)(new syn.).

nitidicollis Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 162. (6 HT, Australie, Bates,

BMNH)(new syn.).

cyanea Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 105. (¢ HT, Wialki, WA, S. Barker,

SAMA I 21512). WA.

cyaneopyga Carter, 1923. Proc. Linn. Soc. N.S.W.

48: 170. (2 LT, Lake Austin, WA, H.

W. Brown, AMSA K67292). WA.

goerlingi Carter, 1936. Proc. Linn. Soc.

N.S.W. 61: 104. (2 HT, Marloo stn.,

Wurarga, WA, A. Goerling, AMSA

K67466)(new syn.).

cyanura Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 163.(2 dd ST & 2 ST, BMNH). Q.

derbyensis Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 106. (¢ HT, Derby, H. W. Brown,

SAMA I 21520). NWA.

duodecimmaculata (Fabricius, 1801)(Buprestis).

Sys. Eleuth.: 191. (HT sex unknown,

BMNH). All mainland states.

duodecimguttata (Boisduval, 1835)

(Buprestis). Voy. Astrolabe, Entom. 2:

93. (HT, MNHN not seen).

quaturodecimnotata Hope, 1846. Ann. Mag.

Nat. Hist. 17: 64. (HT not seen, HMOE).

pustulata Thomson, 1879. Typi Bupr. Mus.

CHECKLIST OF CISSEIS (S.S.) GORY & LAPORTE, 1839 87

Thoms. App 1A: 51. (HT sex unknown,

MNHN).

biologia Froggatt, 1910. Australian Insects:

165. (HT not seen, AMSA?)(new syn.).

fallaciosula Obenberger, 1935. Acta. Soc.

Ent. Csl. 32: 36. (HT sex unknown,

PMCE)(new syn.).

elongatula Blackburn, 1888. Proc. Linn. Soc.

N.S.W. 3: 892. (6 HT, NT, SAMA). NT.

elliptica Carter, 1923. Proc. Linn. Soc. N.S.W. 48:

170. (2 LT, Cue, WA, H. W. Brown,

NMVA). WA.

excelsior Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 107. (d HT, Mt Carbine, Q, T. M. S.

Hanlon & M. Powell, SAMA I 21514). Q.

ernestadamsi Barker, 1999. Rec. S. Aust. Mus. 32:

46. (6 HT, Edungalba, Q, E. E. Adams,

ANIC). Q.

fascigera Obenberger, 1919. Ent. Mitteilungen 8:

20. (6 HT, New Guinea, PMCE no.

23772). New Guinea.

fraterna Kerremans, 1900. Mem. Soc. Ent. Belg.

7: 80. (2 HT, Germ. N. G., Webster,

BMNH). New Guinea.

frontalis Kerremans, 1898, Ann. Soc. Ent. Belg.

42: 157. (S HT, New Guinée, Standing,

BMNH). New Guinea.

fulgidicollis Macleay, 1888. Proc. Linn. Soc.

N.S.W. 3: 1231. (2 do ST, Derby,

NWA, ANIC). NWA, Q.

gibbera (Carter, 1937)(Neospades). Trans. Roy.

Soc. S. Aust. 61: 121. (2 HT, Mullaly,

NSW, H. J. Carter, AMSA K111950).

NSW.

gouldi (Hope, 1846)(Ethon). Ann. Mag. Nat. Hist.

17: 65. (2 HT, Port Essington, HMOE

no. 889). NT.

heroni Carter, 1934. Proc. Linn. Soc. N.S.W. 59:

259. (6 HT, Dorrigo, NSW, W. Herron,

AMSA K67298). NSW.

impressicollis Macleay, 1878. Ent. Soc. N.S.W. 2:

248. (2 HT, Gayndah, AMSA K32713).

viridiaurea Macleay, 1878. Ent. Soc. N.S.W.

2: 248. (2 2 2 ST, Gayndah, AMSA no.

K32712)(new syn.).

viridicuprea Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 160. (2 HT, Cocktown (sic),

Standing, BMNH)(new syn.).

nitida Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 164. (6 HT, Cocktown (sic),

BMNH)(new syn.).

inops Kerremans, 1898. Ann. Soc. Ent. Belg. 42:

168. (2 HT, Australie, Standing,

BMNH). NSW.

kohouti Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 107. (6 HT, Canberra, K. Pullen,

ANIC). NSW, Q.

laticollis Carter, 1923. Proc. Linn. Soc. N.S.W.

48: 169. (2 HT, BMNH). Q.

leucosticta (Kirby, 1818)(Buprestis). Trans. Ent.

Soc. Lond. 12: 382. (2 HT, BMNH). All

mainland states.

aurulenta Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 159. (@ HT, Australia,

Chevrolat, BMNH)(new syn.).

fulgidifrons Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 161. (2 HT, Australia,

Chevrolat, BMNH).

macmillani Barker, 2001. Trans. Roy. Soc. S.

Aust. 125: 108. (6 HT, Wanneroo, WA,

R. P. McMillan, SAMA I 21516). WA.

macqueeni Barker, 2001. Trans. Roy. Soc. S.

Aust. 125: 108. (6 HT, Milmerran, Q, J.

Macqueen, ANIC). Q.

maculata (Gory & Laporte, 1839)(Ethon). Mon.

Bupr. ii: 5. (HT, MNHN not seen). NSW.

marmorata Gory & Laporte, 1839. Mon. Bupr. ii:

4. (HT, MNHN not seen). NSW, Q.

viridicollis Thomson, 1879. Typi Bupr. Mus.

Thomson. App. 1A: 50. (6 HT, MNHN).

aena Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 158. (3 22 ST, Sydney, Bates,

BMNH).

minutissima Thomson, 1879. Typi Bupr. Mus.

Thomson. App. 1A: 54. (HT, MNHN not

seen). SA, V.

simplex Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 171. (2 dd ST & Q ST,

Australie, Standing, BMNH)(new syn.).

modesta Kerremans, 1898. Ann. Soc. Ent. Belg. 42:

170. (S HT, Australie, Stark, Chevrolat

collection, BMNH). WA, SA, V.

myallae Carter, 1934. Proc. Linn. Soc. N.S.W. 59:

260. (2 HT, Bogan R., NSW, J.

Armstrong, AMSA K67299). NSW.

nigrita Kerremans, 1898. Ann. Soc. Ent. Belg. 42:

168. (d HT, Australie, Fairmaire,

BMNBH).

nigripennis Macleay, 1888. Proc. Linn. Soc.

N.S.W. 3: 1231. (¢ HT, Barrior (sic)

Range, NWA, Froggett, ANIC). NT, WA.

ignicollis Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 164. (od ST, Australie,

Standing, BMNH; o ST, Australie,

Chevrolat, BMNH; 2 doo ST, Port

Denison BMNH)(new syn.).

nigromaculata Kerremans, 1895. Ann. Soc. Ent.

Belg. 39: 218. (HT not in BMNH).

Malacca.

88 S. BARKER

nitidiventris Carter, 1934. Proc. Linn. Soc.

N.S.W. 59: 260. (2 HT, Gosford, NSW,

N. MacGregor, AMSA K67300). NSW,

niveosparsa (Carter, 1927)(Neospades). Proc.

Linn. Soc. N.S.W. 52: 228. (6 HT, Bogan

R., NSW, J. Armstrong, AMSA

K67288). NSW, SA.

notulata (Germar, 1848)(Ethon). Linn. Ent.: 178.

(3d HT, Adelaide, HUMB). SA, NSW, T.

atroviolacea Thomson, 1879. Typi. Bupr.

Mus. Thoms. App. 1A: 52. (6 ST & @

ST, Austr, MNHN)(new syn.).

morosa Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 158. (2 d6¢ ST, Australie,

Stark, BMNH)(new syn.).

nubeculosa (Germar, 1848)(Ethon). Linn. Ent.:

176. (2 LT, Adelaide, HUMB). SA, V.

chalcoptera (Germar, 1848)(Ethon). Linn.

Ent.: 177. (2 36 3 ST, Adelaide, HUMB).

similis Saunders, 1871. Cat. Bupr.: 103. (2

HT, Adelaide, BMNH)(new syn.).

oblonga Kerremans, 1903. in Wytsman Genera

Ins. fas. 12: 229. (2 HT, Australie,

Fairm., BMNH). V, NSW, Q, T.

obscura Blackburn, 1887. Trans. R. Soc. S. Aust.

10: 252. (2 HT, T 320, type locality

unknown, BMNH)(unique).

opima Thomson, 1879. Typi Bupr. Mus.

Thomson. App. 1A: 50. (@ HT,

Champion Bay, MNHN). WA.

ovalis Carter, 1923. Proc, Linn. Soc. N.S.W. 48:

170. (2 HT, Nov. Holl. occid., Fry Coll.

1905.100, BMNH 37811)(unique). WA.

parva Blackburn, 1887. Trans. R. Soc. S. Aust.

10: 253. (6 HT, BMNH). WA, SA, V,

NSW.

pygmaea Blackburn, 1891. Trans. R. Soc. S.

Aust. 14: 299. (2 HT, Victoria, C.

French, NMVVA)(new syn.).

patricia Carter, 1935. Proc. Linn. Soc. N.S.W. 55:

180. (d6 HT, Bunbury, WA, F. L.

Whitlock, AMSA K67302). WA, V,

NSW.

pauperula Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 167. (do HT, Australie, A. Deyr.,

Chevrolat collection, BMNH). T.

prasina Carter, 1923. Proc. Linn. Soc. N.S.W. 48:

168. (6 LT, Australia, Blackburn’s

collection, 3267, SAMA). SA, Q.

puella Kerremans, 1898. Ann. Soc. Ent. Belg. 42:

170. (2 66 ST & @ ST, Australie,

Fairm., BMNH). Q.

curta Kerremans, 1903. in Wytsman Genera

Ins. fas. 12: 229. (6 HT, Peak Downs,

Fairmaire, Mus. Godeffroy no. 11142,

BMNB).

pulchella Carter, 1923. Proc. Linn Soc. N.S.W.

48: 171. (HT sex unknown, French

collection, NMVA). Q.

pulleni Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 109. (6 HT, Edungalba, Q, S.

Barker, SAMA I 21518). ACT, Q.

regalis Thomson, 1879. Typi Bupr. Mus.

Thomson. App. 1A: 50. (6 HT,

Queensland, MNHN). NT, Q.

robertfisheri Barker, 1999. Rec. S. Aust. Mus. 32:

47. (d HT, Melrose, SA, R. H. Fisher,

SAMA I 21407). SA.

roseocuprea (Hope, 1846)(Ethon). Ann. Mag.

Nat. Hist. 17: 64. (2 HT, Moriatta (sic),

SA, HMOE no. 889). WA, SA, V, NSW.

dispar Blackburn, 1891. Trans. R. Soc. S.

Aust. 14: 297. (2 HT, 3832, BMNH).

fairmairei Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 169. (2 HT, Australie,

Fairmaire, BMNH).

cuprea Kerremans, 1903. in Wytsman Genera

Ins. fas. 12: 228. (2 HT, NSW, Standing,

BMNB).

rubicunda Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 169. (¢ HT, Australie, Fairm.,

BMNH). WA, SA, V, NSW, Q.

undulata Kerremans, 1903. in Wytsman

Genera Ins. fas. 12: 227. (2 HT, NSW,

Standing, BMNH)(new syn.).

purpurea Kerremans, 1903. in Wytsman

Genera Ins. fas. 12: 228. (2 HT, Sidney

(sic), Fairm, Mus. Godeffroy no.

11108)(mew syn.).

scabiosa (Boisduval, 1835)(Buprestis). Voy.

Astrolobe Entom. 2: 96. (HT, MNHN not

seen).

scabrosula Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 167. (3 63d ST & @ ST, Australie,

Fairm., BMNH). NSW, Q.

semiobscura Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 170. (o ST, Australie, Fairm.,

BMNH; 2 ¢@ ST, Australie, Deyr.,

BMNH). SA, V.

septuosa Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 109. (6 HT, Charity Ck Bridge,

Manning R., NSW, S. Watkins, SAMA I

21521). SA, NSW, Q.

sexnotata Fauvel, 1891. Rev. d’Ent. 10: 181. (HT,

MNHN not seen). New Caledonia.

signaticollis (Hope, 1846)(Ethon). Ann. Mag. Nat.

Hist. 17: 64. (d6 HT, Port Essington,

HMOE no. 891). WA, NT, Q.

speciosa Barker, 2001. Trans. Roy. Soc. S. Aust.

CHECKLIST OF CISSEIS (S.S.) GORY & LAPORTE, 1839 89

125: 110. (6 HT, Midland Jctn, WA, R.

P. McMillan, SAMA I 21523). WA.

stellata Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 111. (6 HT, Cairns, SAMA I

21524). Q.

stigmata Gory & Laporte, 1839. Mon. Bupr. ii: 3.

(HT, MNHN not seen). WA.

subbifascialis Carter, 1927. Proc. Linn. Soc.

N.S.W. 52: 229. (2 HT, Bogan R., NSW.

J. A., AMSA K67291). NSW.

tasmanica Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 165. (2 ST, Tasmanie, Oberthur,

BMNH; ST, Tasmanie, Chevrolat,

BMNH). T, V, NSW.

trimentula Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 111. (6 HT, Pilliga East SF, NSW,

S. Watkins, SAMA I 21526). V, NSW, Q.

tyrrhena Carter, 1923. Proc. Linn. Soc. N.S.W.

48: 168. (2 HT, Kalamunda, WA, H. M.

Giles, AMSA K67293). WA

vicina Kerremans, 1898. Ann. Soc. Ent. Belg. 42:

164. (6 HT, Australie, Standing,

BMNH). V, NSW, Q.

collaris Kerremans, 1903. in Wytsman,

Genera Ins. fas. 12: 229. (2 36 ST,

Gayndah, Fairmaire, BMNH).

ornata Kerremans, 1903. in Wytsman Genera

Ins. fas. 12: 229. (d HT, NSW, Standing,

BMNH).

violacea Kerremans, 1903. in Wytsman Gen. Ins.

fas. 12: 228. (6 ST & 2 ST, Gayndah,

Fairmaire, BMNH). NSW, Q.

viridiceps Kerremans, 1898. Ann. Soc. Ent. Belg.

42: 168. (6 HT, Australie, Standing,

BMNH). NSW.

viridipurpurea Carter, 1924. Proc. Linn. Soc.

N.S.W. 49: 27. (2 36 ST, Geraldton,

WA, J. Clark, AMSA K67294). WA.

sapphira Carter, 1933. Proc. Linn. Soc.

N.S.W. 58: 163. (2 HT, Moore R., WA,

H. W. B., AMSA K67295)(new syn.).

watkinsi Barker, 2001. Trans. Roy. Soc. S. Aust.

125: 112. (¢ HT, Tinonee Rd, Manning

R., NSW, S. Watkins, SAMA I 21528).

NSW.

westwoodi (Gory & Laporte, 1839)(Coraebus).

Mon. Bupr. ii: 15. (6 ST & 2 2 ST,

MNHN ). SA, V, T.

verna Blackburn, 1891. Trans. R. Soc. S.

Aust. 14: 299. (HT damaged sex

unknown, Warburton District, Victoria,

French collection, NMVA).

viridana Kerremans, 1898. Ann. Soc. Ent.

Belg. 42: 165. (2 HT, Austral., Standing,

BMNH).

theryi Kerremans, 1903. in Wytsman Genera

Ins. fas. 12: 229. (2 6d ST & 2 22 ST,

Victoria, Théry, BMNH).

DISCUSSION

C. adspersa Kerremans (HT not in BMNH) hab.

India belongs in another genus.

C. albertisi Gestro is not a synonym of C.

acuducta (Carter 1923) which does not occur

at the tip of Cape York in the type locality.

C. cornuta Gestro (6 ST, MCSNG) = Hypocisseis

latipennis Macleay.

C. duodecimguttata Guérin-Méneville, 1830

(Buprestis). Type species by subsequent

designation (Duponchel, 1843).

C. fossicollis Kerremans (HT BMNH) = Alcinous

fossicollis (Kerremans).

C. fulgidicollis Macleay is very uncommon in

collections and has been confused with other

species, particularly with C. stellata Barker.

C. gestroi Kerremans 1892, Mem. Soc. Ent. Belg.

1: 225 is unknown to me; the holotype is not

in the BMNH and the species is not listed

under Cisseis in Kerremans (1903).

C. maculata Gory & Laporte was misidentified by

Carter (1929) who listed C. tasmanica

Kerremans and C. pauperula Kerremans as

its synonyms; both are good species.

C. modesta Kerremans was listed as a synonym of

C. rubicunda Kerremans Carter (1929). I find

they are separate species.

C. nigrita Kerremans was listed as a synonym of

C. notulata (Germar) by Carter (1929). Carter

(1923) followed Blackburn’s concept of C.

notulata, which was incorrect. C. nigrita is a

separate species.

C. nigripennis Macleay was listed by Carter

(1923) as belonging in Neospades. I find it

belongs in Cisseis (s.s.).

C. obscura Blackburn 1s a species that I have been

unable to find in any available collection.

C. ovalis Carter could be a synonym of C. opima

Thomson. The unique holotype is a

discoloured female specimen.

. rugiceps Thomson belongs in Neospades.

. scabiosa (Boisduval) is unknown to me.

. semiobscura Kerremans was listed as a

synonym of C. notulata by Carter (1929). It is

a separate species, much smaller than C.

notulata and common in South Australia.

C. spilota Hope, 1846 (Ann. Mag. Nat. Hist. 17:

64) (HT, HMOE not seen) is unknown to me.

Hope indicated that it belonged in another

AANA

90 S. BARKER

genus. Carter (1929) incorrectly listed it as a

synonym of C. signaticollis Hope, 1846;

spilota has precedence over signaticollis.

C. stigmata Gory & Laporte, 1839 selected as type

species by Bellamy (1998) (invalid).

C. violacea Kerremans was listed as a synonym of

C. notulata by Carter (1929). It is a separate,

smaller species occurring in the vicinity of

Brisbane, Queensland.

ACKNOWLEDGMENTS

My thanks for assistance to: C. L. Bellamy,

Sacramento; M. Moulds, AMSA; T. A. Weir, ANIC; M.

Kerley, BMNH; A. Samuelson, BPBM; I. Lansbury,

HMOE; M. Uhlig, HUMB; R. Poggi, MCSNG; M.

Hanlon, Sydney; J. Menier, MNHN; K. Walker & C.

McPhee, NMVA; S. Bily, PMCE; G. B. Monteith,

QMBA,; J. Cools, RMBB; E. G. Matthews, SAMA; J.

A. Gardner, Waite Agricultural Research Institute,

Adelaide.

REFERENCES

BARKER, S. 1998. Selection of lectotypes and

redescriptions of three Cisseis (Coleoptera:

Buprestidae) species. Records of the South

Australian Museum 31: 21-23.

BARKER, S. 1999a. Designation of a lectotype and

descriptions of four new species of Australian

Buprestidae (Coleoptera). Records of the South

Australian Museum 32: 45-49.

BARKER, S. 1999b. Designation of lectotypes of three

species of Cisseis (Coleoptera: Buprestidae).

Transactions of the Royal Society of South Australia

123: 155.

BARKER, S. 2001. Descriptions of twenty one new

species of Cisseis (sensu stricto) Gory & Laporte

1839 (Coleoptera: Buprestidae: Agrilinae).

Transactions of the Royal Society of South Australia

125: 97-113.

BARKER, S. & BELLAMY, C. L. 2001.

Stanwatkinsius, a new genus of Australian jewel

beetles (Coleoptera: Buprestidae: Agrilinae) with a

key to known species. Transactions of the Royal

Society of South Australia 125: 1-14.

BELLAMY, C. L. 1998. Type species designations in

the family Buprestidae (Coleoptera). Deutsche

Entomologische Zeitschrift 45: 9-15.

CARTER, H. J. 1923. Revision of the Genera Ethon,

Cisseis and their allies. (Buprestidae.). Proceedings

of the Linnean Society of N.S.W. 48: 159-176.

CARTER H. J. 1929. A check list of the Australian

Buprestidae. Australian Zoologist 5: 265-304.

DUPONCHEL, P. A. J. 1843 (1841-1848). In

‘Dictionaire universal d’histoire naturelle’. Vols 1-6.

Ed. C. D. D’Orbigny. Bureau Principal de l’Editeur

& Renard, Martinet & Cie: Paris.

GORY, H. L. & LAPORTE DE CASTELNAU, F. L.

1839. ‘Histoire naturelle et iconographie des insectes

Coléoptéres. Monographie des buprestides’. Volume

2, livraisons 25—35. P. Duménil: Paris.

KERREMANS C. 1903. Genera Insectorum 12: 1-338.

THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS

COMMITTEE 10 MARCH 1921 - 30 OCTOBER 2001

WOLFGANG ZEIDLER

Summary

The Flora and Fauna of South Australia Handbooks Committee has been operating for 80 years

producing a series of handbooks on the major elements of the South Australian natural

environment. The main role of the Committee was to encourage and persuade specialists to write

books on the flora and fauna for no financial reward. The completed manuscripts were then edited

by members of the Committee and, in the past, were printed by State Print or its equivalent and

published by the State Government Printer at the State Government’s expense. Several years ago

this part of the Government’s operations ceased, with no provision made for the further printing of

handbooks.

THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE

10 MARCH 1921 —- 30 OCTOBER 2001

WOLFGANG ZEIDLER

ZEIDLER, W. 2002. The Flora and Fauna of South Australia Handbooks Committee 10 March

1921 — 30 October 2001. Records of the South Australian Museum 35(1) 91-95.

Wolfgang Zeidler, South Australian Museum, North Terrace, Adelaide, South Australia 5000;

and Visiting Research Fellow, Adelaide University, Adelaide, South Australia 5005.

Manuscript received 13 December 2001.

The Flora and Fauna of South Australia

Handbooks Committee has been operating for 80

years producing a series of handbooks on the

major elements of the South Australian natural

environment. The main role of the Committee was

to encourage and persuade specialists to write

books on the flora and fauna for no financial

reward. The completed manuscripts were then

edited by members of the Committee and, in the

past, were printed by State Print or its equivalent

and published by the State Government Printer at

the State Government’s expense. Several years

ago this part of the Government’s operations

ceased, with no provision made for the further

printing of handbooks.

Since the privatisation of the Government Printer

the Committee has sought other means to publish

its books and the last two handbooks have been

printed with the help of the Board of the Botanic

Gardens (‘Fungi’) and the SA Research &

Development Institute (Aquatic Sciences) (‘Marine

Invertebrates LI’). However, it became very clear

to the Committee that the only way it could

continue was to establish a publications fund.

Initially, the State Government was approached

because of its traditional support for the

Committee, but this proved fruitless, although the

Department of Environment & Heritage and the

South Australian Museum did offer some support

if funds could also be found from other sources.

Various philanthropic trusts were approached and,

although some were sympathetic, no funding was

forthcoming. Similarly, all three of the State’s

universities, whose students and staff are amongst

the main users of the handbooks, declined to assist.

The formation of a partnership with a publisher

was also considered, but rejected as it would

potentially compromise the original intention of the

Committee because handbooks with commercial

potential would be favoured.

Left with no alternative, the Handbooks

Committee reluctantly concluded that it could not

continue to function as originally intended and

should be disbanded. The last meeting of the Flora

and Fauna of South Australia Handbooks

Committee was accordingly held on 30 October

2001.

With the demise of the Handbooks Committee,

which was once the envy of all other states, it is

important to record its history and major

achievements.

The Committee began in 1921 when Prof E.

Wood-Jones, Prof T. B. G. Osborn and Prof J. B.

Cleland presented the idea to the then Premier Mr

H. Barwell. The concept was for authoritative

handbooks to be written by experts in their field

without remuneration, to be published by the

Government Printer at its convenience, and to be

sold at a low price so as to be affordable to the

wider community. Here follows an extract

recording the foundation of the handbooks series,

prefacing Eichler’s (1965) ‘Supplement to J. M.

Black’s Flora of South Australia’.

After the first World War, the South Australian

Branch of the British Science Guild was very

active in devising ways for increasing scientific

knowledge and its applications. At the time an

up-to-date account of the Flora of our State was

a pressing need. With this view, Professor J. B.

Cleland suggested that the Science Guild might

undertake to arrange for the preparation of a

series of Handbooks dealing with the Fauna and

Flora of South Australia. If these were prepared

in an honorary capacity by leading authorities,

the Government might well be asked to publish

them, as it were, on a pound for pound basis.

The Branch accepted the suggestion and it was

arranged that a deputation of Professors F.

Wood Jones, T. G. B. Osborn and J. B. Cleland

should wait on the Hon. The Premier Mr.,

afterwards Sir Henry Barwell, with the proposal.

W. ZEIDLER

Fortunately Professor Cleland had prepared a

memorandum on the subject in the following

terms and this document was handed to the

Premier on 25" February, 1921.

PROPOSED SCHEME FoR THE PREPARATION OF

HANDBOOKS ON THE FAUNA AND FLORA

OF SOUTH AUSTRALIA

L There is an almost complete lack in South

Australia of any handbooks dealing with

the Fauna and Flora of the State.

Il. In consequence, many individuals with

scientific tastes are unable to develop

these. This is a great loss to the State, as

these persons, with suitable handbooks

available, might be led on to contribute

voluntarily to the State additions to

scientific knowledge of great value to our

pastoral, agricultural and other interests.

II. | A scheme under the auspices of the S.A.

Branch of the British Science Guild is

proposed for the preparation of a series

of such handbooks to be edited by a sub-

committee of experienced workers, the

individual parts to be prepared by

specialists (in an honorary capacity).

IX. The scheme would be spread over a

period of 10-15 years.

X. Parts would be issued as completed by

the authors. None could be available

during this financial year. One or two

may be so for 1921-1922.

XI. — The work involved will be highly skilled

and laborious. In the interests of science

the authors will be prepared, we believe,

to devote their time and knowledge to this

work without fee, if a means of

publication can be obtained.

XII. The value to the community of the works

of these authors, thus offered as a gift to

the State and world of science, can be put

at a very high figure. Would the

Government be prepared to accept this

offer of gratuitous service and as a return

— on the “pound for pound subsidy”

principle — arrange for the publication of

the Handbooks by the Government

Printer?

XII. As the work will be spread over many

years, this should not necessitate any

additions to the staff of the Government

Printer, the printing of the parts being

proceeded with by him as opportunity

offered. The actual out-of-pocket expense

to the State would consist, under these

circumstances, of the paper only. The

parts as completed might be submitted to

the Premier or to the Minister of

Education, and laid on the table of the

House and ordered to be printed and thus

be dealt with as Parliamentary Papers.

IX. Many of the parts will require to be

illustrated. In many cases, blocks are

already prepared which doubtless could

be used and so reduce expense.

X. The educational value of these handbooks

will be great. A copy of each might be

placed with advantage in every public

school in the State, when it could be made

available not only to teachers and

scholars but to the public of the district

as well. To meet this demand and to

enable distribution to be made to private

individuals (at a nominal price to induce

the wide use of these handbooks) about

1,500 copies of each should be printed.

XI. Probably from 20 to 30 handbooks would

eventually be published (over the period

of 10-15 years), the sizes varying from

about 20 to 150 pages according to

subject matter.

Professor CLELAND, Adelaide University.

Professor WOOD JONES, Adelaide University.

The Premier read through the memorandum and

said, “A very generous offer, gentlemen. I shall

lay it before Cabinet.” Professor Cleland

received from the secretary to the Premier, a

letter dated 10° March, 1921 in the following

terms accepting the offer.

“With reference to the proposed scheme under

the auspices of the South Australian Branch of

the British Science Guild for the preparation of

a series of handbooks on the Fauna and Flora of

South Australia which was submitted by

yourself and Professor Wood Jones to the

Premier on the 25" ultimo, I am directed by the

Premier to inform you that in consideration of

the contributors in the compilation of the

handbooks undertaking the work in an honorary

capacity the Government is prepared to

undertake the printing of the publication at the

Government Printing Office at the expense of

the State.”

In recent years the Chief Secretary of South

Australia decided that the Government Printer

should publish the handbooks on a commercial

basis, rather than that the Handbooks Committee

should apply each year for a government grant

for publication. Consequently there is now no

delay in printing when the authors offer their

completed manuscripts.

The British Science Guild has long since ceased

to exist but the work of the Handbooks Committee

survived until now. The main aim of the

THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 93

Committee has always been to produce books that

document, in a rigorously scientific way, the

biological heritage of the State, although in recent

years the Committee encouraged authors to

broaden their focus to southern Australia in

general. Perhaps the most outstanding work

published by the Committee is the ‘Flora of South

Australia’. The remarkable J. M. Black produced

the four-volume work single-handedly and revised

the first three volumes, working until he was 96.

Revision of the fourth volume was completed

after his death. For South Australia’s

sesquicentenary a completely revised and

rewritten new edition was produced, making this

State the only one at the time to have a

comprehensive up-to-date record of State flora.

There are also other monumental works for

which other states have no equivalent (see list of

handbooks) and, while some volumes are out of

date, many have been updated or reprinted to meet

the demand of scientists, naturalists and students.

It seems remarkable that Hale’s “The Crustaceans

of South Australia’ (1927, 1929), reprinted in

1976, is still used as a basic text, although very

much out of date and only providing a limited

coverage of the group.

It is clear that the handbooks provide invaluable

data for research workers, not only in South

Australia, or even nationally, but internationally

as well. Indeed, the handbooks series has been the

envy of scientists in other states. Their

contribution to secondary and tertiary education,

as significant textbooks, is well recognised. For

example, the four-part ‘Flora’, ‘Fishes’ and the

three-part ‘Marine Invertebrates of Southern

Australia’ series are essential texts for tertiary

students. Unfortunately, the later volumes

produced by the Government Printer were

managed on a commercial basis and were

remaindered within two years, so that they were

only available for a relatively short period.

Future plans for new editions of ‘Fishes’,

Amphibians’, ‘The Vegetation of South Australia’,

‘Crustacea’ (as part of the ‘Marine Invertebrates’

series) and ‘Flora’ have been abandoned, although

they may be taken up by others. In particular, the

State Herbarium is in the process of producing an

electronic version of the ‘Flora’, and the South

Australian Research & Development Institute

(Aquatic Sciences) intends to proceed with future

volumes of the ‘Marine Invertebrates’ series as well

as revising earlier volumes.

As part of the winding up process, the

Committee has transferred copyright of the

botanical texts to the Board of the Botanic Gardens

and State Herbarium, and of the zoological and

ecological texts to the Board of the South

Australian Museum. Copies of correspondence and

minutes of the Handbooks Committee have been

deposited with State Archives.

TitLes IssueD By THE FLorA AND FAUNA OF SOUTH

AUSTRALIA HANDBOOKS COMMITTEE

1922 ‘Flora of South Australia’. Part I.

Cyathaceae — Orchidaceae. J. M. Black.

Second edition 1943. Reprinted 1948.

Reprinted (facsimile) 1960, 1972. Third

edition, Lycopodiaceae — Orchidaceae

(revised by J. P. Jessop) 1978.

‘The Fishes of South Australia’. E. R.

Waite.

Reprinted 1962.

‘The Mammals of South Australia’. Part I.

The Monotremes and the Carnivorous

Marsupials. F. Wood-Jones.

Reprinted in one volume with parts II

(1924) and III (1925), with an Introduction

by P. Crowcroft and a Selected

Bibliography by J. H. Calaby 1968.

‘Flora of South Australia’. Part II.

Casuarinaceae — Euphorbiaceae. J. M.

Black.

Second edition 1948. Reprinted 1963.

Reprinted (facsimile) 1977.

‘The Mammals of South Australia’. Part

II. The Bandicoots and the Herbivorous

Marsupials. F. Wood-Jones.

Reprinted in one volume with parts I

(1923) and III (1925), as above, 1968.

‘The Building of Australia and the

Succession of Life’ with special reference

to South Australia. Part I. W. Howchin.

‘The Mammals of South Australia’. Part

Il. The Monodelphia. F. Wood-Jones.

Reprinted in one volume with parts I

(1923) and II (1924), as above, 1968.

‘Flora of South Australia’. Part III.

Meliaceae — Scrophulariaceae. J. M.

Black.

Second edition, Callitrichaceae —

Plumbaginaceae 1952. Reprinted 1964.

Reprinted (facsimile) 1977.

‘The Crustaceans of South Australia’. Part

J. H. M. Hale.

Reprinted in one volume with part II

(1929) 1976.

1923

1924

1925

1926

1927

1928

1929

1930

1934

1935

1936

1937

1938

1939

1940

1947

W. ZEIDLER

‘The Building of Australia and the

Succession of Life’ with special reference

to South Australia. Part II. Mesozoic and

Cainozoic. W. Howchin.

‘Flora of South Australia’. Part IV.

Bignoniaceae — Compositae. J. M. Black.

Second edition, Oleaceae — Compositae

(revised by E. L. Robertson) 1957.

Reprinted (facsimile) 1980.

‘The Crustaceans of South Australia’. Part

Il. H. M. Hale.

Reprinted in one volume with part I (1927)

1976.

‘The Reptiles and Amphibians of South

Australia’. E. R. Waite.

‘The Building of Australia and the

Succession of Life’ with special reference

to South Australia. Part III. Pleistocene.

W. Howchin.

“Toadstools and Mushrooms and other

Larger Fungi of South Australia.’ General

introduction; Toadstools and Mushrooms.

J. B. Cleland.

Reprinted in one volume with part II

(1935), including ‘Notes on some Edible

and Poisonous Fungi’ by P. H. B. Talbot,

1976.

‘Toadstools and Mushrooms and other

Larger Fungi of South Australia’.

Polypores, Coral Fungi and remaining

Hymenomycetes; Puff-balls, Jelly-like

fungi, the larger Ascomycetes and

Myxomycetes. J. B. Cleland.

Reprinted in one volume with part I

(1934), as above.

‘The Seaweeds of South Australia’. Part I.

Introduction and The Green and Brown

Seaweeds. A. H. S. Lucas.

‘The Vegetation of South Australia’. J. G.

Wood.

‘The Molluscs of South Australia’. Part I.

The Pelecypoda. B. C. Cotton & F. K.

Godfrey.

‘Primitive Insects of South Australia’.

Silverfish, Springtails and their allies. H.

Womersley.

‘The Molluscs of South Australia’. Part II.

Scaphopoda, Cephalopoda, Aplacophora

and Crepipoda. B. C. Cotton & F. K.

Godfrey.

‘The Seaweeds of South Australia’. Part

1959

1961

1962

1964

1965

1966

1972

1974

1975

1976

1978

1979

1980

1982

1984

1986

II. The Red Seaweeds. A. H. S. Lucas & F.

Perrin.

‘South Australian Mollusca’.

Archaeogastropoda. B. C. Cotton.

‘South Australian Mollusca’. Pelecypoda.

B. C. Cotton.

‘The Marine and Freshwater Fishes of

South Australia’. T. D. Scott.

‘South Australian Mollusca’. Chitons. B.

C. Cotton.

‘Supplement to J. M. Black’s Flora of

South Australia’ (second edition 1943-—

1957). H. J. Eichler.

‘Aboriginal Man in South and Central

Australia’. Part I. Edited by B. C. Cotton.

Only one part was published.

‘The Vegetation of South Australia’. R. L.

Specht. Second edition.

‘The Marine and Freshwater Fishes of

South Australia’. T. D. Scott, C. J. M.

Glover & R. V. Southcott. Second edition.

Reprinted (facsimile, with 16 pp of colour

plates) 1980.

‘Plant Feeding and other Bugs (Hemiptera)

of South Australia’. Heteroptera Part I. G.

F. Gross.

‘Plant Feeding and other Bugs (Hemiptera)

of South Australia’. Heteroptera Part II. G.

F. Gross.

‘Butterflies of South Australia’. R. H.

Fisher.

‘Amphibians of South Australia’. M. J.

Tyler.

‘Lichens of South Australia’. R. B. Filson

& R. W. Rogers.

‘Mosses of South Australia’. D. G.

Catcheside.

‘Acacias of South Australia’. D. J. E.

Whibley.

‘Marine Invertebrates of Southern

Australia’. Part I. Edited by S. A.

Shepherd & I. M. Thomas.

‘Psylloidea of South Australia’. F. D.

Morgan.

‘The Marine Benthic Flora of Southern

Australia’. Part I. H. B. S. Womersley.

‘The Dynamic Partnership: Birds and

Plants in Southern Australia’. Edited by H.

A. Ford & D. Paton.

1986

1987

1989

THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 95

‘Flora of South Australia. Volumes I-IV’.

Edited by J. P. Jessop & H. R. Toelken.

Fourth edition (third edition consisted of

part I only).

‘The Ecology of Forests and Woodlands

of South Australia’. Edited by H. R.

Wallace.

‘The Marine Benthic Flora of Southern

Australia’. Part II. H. B. S. Womersley.

‘Marine Invertebrates of Southern

Australia’. Part II. Edited by S. A.

Shepherd & I. M. Thomas.

1990

1992

1994

1997

‘Orchids of South Australia’. R. Bates &

J. Z. Weber.

‘Acacias of South Australia’. D. J. E.

Whibley & D. E. Symon. Second edition.

‘The Fishes of Australia’s South Coast’.

Edited by M. F. Gomon, J. C. M. Glover

& R. Kuiter.

‘Larger Fungi of South Australia’. C. A.

Grgurinovic.

‘Marine Invertebrates of Southern

Australia’. Part III. Edited by S. A.

Shepherd & M. Davies.

Dr Wolfgang Zeidler (Hon. Secretary), on behalf of the Flora and Fauna of South Australia Handbooks Committee

— Em Prof William D. Williams (Chairman), Dr Sue Barker (Botanical Editor), Dr Margaret Davies (Zoological

Editor), Dr Shelley Barker, Ms Robyn Barker, Dr John G. Conran, Mr Peter Copley and Dr Brian D. Morley.

OBITUARY GRAEME LLOYD PRETTY 1940 -— 2000

HELEN TOLCHER

Summary

Graeme Lloyd Pretty was born in Melbourne on 25 June 1940. His father was in the RAAF and the

family soon moved to Richmond NSW, where Graeme spent his childhood and went to school.

After graduating from Sydney University BA (Hons) in Classical Archaeology and Dip Ed, he

contemplated undertaking a PhD with Jack Golson at the Australian National University. However,

he began working at the South Australian Museum in January 1962 as Assistant Curator of

Anthropology under Norman Tindale, from whom he learned much and with whom he kept in touch

long after the latter’s retirement. His first interest was in Melanesian prehistory — he might well

have made this his major field of work.

OBITUARY

GRAEME LLOYD PRETTY

1940-2000

PCE

WES

Graeme Pretty and Henry Pollach (left) at Roonka, 8 August 1971.

Roonka photos R83: 2. South Australian Museum.

Graeme Lloyd Pretty was born in Melbourne on

25 June 1940. His father was in the RAAF and

the family soon moved to Richmond NSW, where

Graeme spent his childhood and went to school.

After graduating from Sydney University with a

BA (Hons) in Classical Archaeology and Dip Ed,

he contemplated undertaking a PhD with Jack

Golson at the Australian National University.

However, he began working at the South

Australian Museum in January 1962 as Assistant

Curator of Anthropology under Norman Tindale,

from whom he learned much and with whom he

kept in touch long after the latter’s retirement. His

first interest was in Melanesian prehistory—he

might well have made this his major field of work.

In 1964 he was appointed Assistant Curator

of Archaeology, and archaeological fieldwork

was added to his duties. He excavated or

assisted at a number of sites on the lower River

Murray, seeking the advice of John Mulvaney

who had recently excavated Fromm’s landing.

At the same time he continued research into

Melanesian culture, including six weeks

fieldwork in the Southern Highlands of Papua

New Guinea (PNG) in 1968-69. Subsequently,

he was invited to review the functions of the

PNG Museum and Art Gallery. Later, he also

advised the Australian National Gallery on its

Melanesian collections.

In 1968 Pretty began a project that was to

become the most important one of his working

life. A salvage operation to remove a surface

campsite and scattered human remains from a

sandhill beside the River Murray at Roonka, near

Blanchetown, developed into a major excavation

that would attract worldwide interest. At that time

he was unique in having consulted with the

relevant Aboriginal group and obtained approval

before beginning work—he continued this contact

throughout and after the conclusion of the dig.

This rapport with the Aboriginal people

undoubtedly contributed to making possible the

excavation on the scale and significance that was

achieved.

98 H. TOLCHER, J. PRESCOTT & J. MULVANEY

Despite the disadvantages and difficulties of

working on a surface fully exposed to gale-force

winds and summer heat rising to the 50°C range,

he retrieved a wealth of archaeological material

and information from a site which proved to have

been occupied for some 18 000 years. In all, 216

complete or fragmentary burials were uncovered.

The grave goods were unprecedented at that time.

In addition to almost every variant of human

mortuary practice, it also gave evidence for

hitherto undocumented customs, of which there

were well over 150 examples.

Roonka looms large for its deep significance.

Here is a statistical sample from one place, across

some thousands of years, permitting health, injury

and dietary studies of the population. Pretty

correctly saw that aspects of the rituals involved—

the mode of burial, including grave goods, and

the antiquity of the rituals—presented a major

window to past belief and behavioural systems.

No other burial ground has produced so much

evidence.

The almost exclusively amateur workforce of

volunteers became, over the eight years of active

fieldwork, a team of patient, dedicated, highly

skilled excavators. Under Pretty’s direction, the

site was strictly controlled, meticulously recorded

and kept secure from outside interference. The

standard and scope of the work attracted

substantial financial support as well as visitors

from abroad, including osteo-archaeologists and a

delegation from the People’s Republic of China.

One group will remember the visit not only for

the archaeology but because their bus became

bogged to the axles on the Roonka Flat.

Pretty was appointed to the position of Senior

Curator of Anthropology and Archaeology in

1975.

The section of Pretty’s work that has been best

documented is probably that dealing with

chronology. He had a critical appreciation of the

role of physical dating methods in archaeology

and sought to create a temporal framework for the

finds at Roonka. The chronology of the site was

determined by a larger number of different dating

methods than any other archaeological site in

Australia, including thermoluminescence,

palaeomagnetism, uranium fluorimetry and C-14

dating, both conventional and AMS.

It is not generally known that he was

responsible for planting the idea for developing

luminescence dating in the Physics Department at

the University of Adelaide in conversation with

Professor John Prescott. This was in 1972, at the

very infancy of the thermoluminescence

technique. The earliest Adelaide work on

thermoluminescence dating per se was based on

samples from Roonka.

Pretty extended his fieldwork research to Island

Melanesia in 1971 and 1973, finding additional

support from sources outside Australia. However,

the demands of attempting to elucidate the

intricacies of the Roonka information, together

with constraints on finance and staffing, resulted

in the Melanesian work being put aside—apart

from ongoing work, under his direction, on the

sorting of the Foreign Ethnology reserve

collections and their eventual rehousing at Kent

Town in 1986. As Senior Curator of Archaeology,

Pretty’s work was directed almost exclusively to

organising the mass of information from the

Roonka site in order to arrive at what he

considered an adequate assessment on which to

base his final report. During the 1980s a master

index of the Roonka data was prepared.

The Third Australasian Archaeometry

Conference was held in Adelaide in 1988. Pretty

devised the theme of the Symposium, ‘Early Man

in the Southern Hemisphere’, which was designed

to give a specifically archaeological balance to the

program. He arranged invitations and finance for

the international contributors: G. Delibrias, G. J.

Bartstraa and H. J. Deacon; and organised the

conference excursion and wrote the field notes.

In his own contribution to the symposium, a

paper analysing the cultural chronology of

Roonka, it is possible to see a foretaste of his

overall interpretation of Roonka in its wider

context. An early indication is also found in his

exciting and innovative 1977 paper, “The cultural

chronology of Roonka Flat’. In it he remarked

‘This present paper therefore celebrates the

author’s satisfaction that the archaeology is now

sufficiently clear to commit preliminary notice of

it to paper’. Yet, during the following 23 years, he

produced no paper with the promised detail. It is

difficult for those of us who knew him to avoid

reflection on his apparent reluctance to put his

ideas on paper where they could be exposed to the

critical appraisal of his peers. He did publish

sections of the work with his collaborators, but

the overall synthesis was yet to come. Perhaps he

was sensitive to possible criticism; perhaps the

very volume of his material was an obstacle. It

seems likely that he laboured over the precise

language with which to present a polished final

monograph.

In 1982 restructuring of the South Australian

Museum’s divisions and staffing made Pretty one

of a number of Curators of Anthropology. He

OBITUARY —- GRAEME LLOYD PRETTY 99

accepted a severance package in 1994, continuing

his work on the Roonka report until 1996, when

he was stricken with viral encephalitis, from

which he never fully recovered. He died on

6 November 2000.

In his work Pretty was patient, a seeker of

precision in language that was baffling to many,

and single-minded to a degree that ignored

workplace relationships and, at times, made it

difficult for people to work with him. His memory

was exceptional. The breadth of his reading was

remarkable and was reflected in his large personal

library. He left the Museum’s Foreign Ethnology

collection well organised and totally accessible,

and the Roonka material fully recorded and

stored. He gained the respect of his field workers,

instructing them in aspects of Aboriginal culture

and inspiring in them a dedication and loyalty to

the project that will ensure the publication of the

final report.

Graeme married Dana in 1973 and they had one

daughter. To those who knew him well he was a

kind and considerate friend. At home he was a

gracious and amusing host, an erudite and

entertaining conversationalist, a devoted husband

and a proud and loving father.

AUTHORED AND COLLABORATIVE WorK To 1995,

LisTED By FIELD

Archaeology

PRETTY, G. L. 1964. Stone objects excavated in New

Guinea. Man 64, 138: 17.

PRETTY, G. L. 1964. A stone figure of a crested

cockatoo from Melanesia. Man 64, 220: 183-184.

PRETTY, G. L. 1965. Two stone pestles from Western

Papua and their relationship to prehistoric pestles and

mortars from New Guinea. Records of the South

Australian Museum 15, 1: 120-130.

PRETTY, G. L. (with M. J. TYLER). 1966. A bizarre

refuge for some Australian leptodactylid frogs.

Records of the South Australian Museum 15, 2: 360-

361.

PRETTY, G. L. 1967. Excavations at the Anglo-Saxon

site of Wallingford, Berkshire, England, 1966.

Journal of the Anthropological Society of South

Australia 5, 2: 3-5.

PRETTY, G. L. 1967. Rescue excavation of an

aboriginal grave, Tailem Bend. South Australian

Naturalist 41, 4: 9-11.

PRETTY, G. L. (with S. A. GALLUS). 1967. The

anthropology and archaeology of the Nullarbor Plain,

South Australia. Pp. 47-49 in ‘Caves of the

Nullarbor’. Eds. J. R. DUNKLEY & T. M. L.

WIGLEY. Adelaide.

PRETTY, G. L. 1968. Excavation of Aboriginal graves

at Gidgealpa, South Australia. Records of the South

Australian Museum 15, 4: 671-677.

PRETTY, G. L. 1968. Two more prehistoric stone

artifacts from Western Papua. Records of the South

Australian Museum 15, 4: 693-697.

PRETTY, G. L. 1969. Prehistoric background to

Australia. Tradition: Journal of the History

Teachers’ Association (S.A.) May. Pp. 13-23.

PRETTY, G. L. 1969. Excavation of an Aboriginal

cemetery near Blanchetown, River Murray, South

Australia. Paper read to the 41st Congress of the

Australian and New Zealand Association for the

Advancement of Science, Adelaide, August 1969. 15

pp., 10 illus.

PRETTY, G. L. 1970. Excavation of an Aboriginal

cemetery on Roonka Station, River Murray. Kalori,

Journal of the Museums Association of Australia 38:

17.

PRETTY, G. L. 1971. Excavations at Roonka Station,

Lower River Murray, South Australia, 1968-1970.

Journal of the Anthropological Society of South

Australia 19, 9: 6-15.

PRETTY, G. L. 1971. Further investigations into

Melanesian culture-history: South Australian

Museum field research in Island Melanesia, 1971.

Kalori: Journal of the Museums Association of

Australia 42: 89-95.

PRETTY, G. L. 1972. ‘Report of an Inspection of

Archaeological Sites and Field Monuments in the

Territory of Papua New Guinea’. Papua New Guinea

Public Museum and Art Gallery: Port Moresby. vii +

SO pp., illus.

PRETTY, G. L. 1975. Archaeology in South Australia:

a report on recent work. Australian Archaeology:

Australian Archaeological Association Newsletter 3:

32-39.

PRETTY, G. L. 1975. ‘Social Change in Prehistoric

Australia; the Evidence from Roonka’. Paper read to

the 46th Congress of the Australian and New Zealand

Association for the Advancement of Science,

Canberra. 12 pp., 16 illus.

PRETTY, G. L. 1976. The excavations at Roonka Flat,

South Australia: an insight into ancient Australian

society. Actes et Proc. IX Congres, Union

Internationale des Sciences Prehistoriques et

Protohistoriques, Nice. Section 6, 2: 100-112.

PRETTY, G. L. 1977. Archaeology in South Australia.

Pp. 40-54 in ‘South Australian Yearbook’.

Australian Bureau of Statistics: Adelaide.

PRETTY, G. L. 1977. The cultural chronology of the

Roonka Flat: a preliminary consideration. Pp. 288-

331 in ‘Stone Tools as Cultural Markers: Change,

Evolution, Complexity’. Ed. R. V. S. WRIGHT.

Australian Institute of Aboriginal Studies:

Canberra.

PRETTY, G. L. 1978. Review of V. D. Watson & J. D.

100

Cole, ‘Prehistory of the Eastern Highlands of New

Guinea’. The Artefact 5: 98-102.

PRETTY, G. L. 1980. Prehistory. Pp. 9-17 in

‘Proceedings of the Fiftieth Anniversary Seminar of

the Anthropological Society of South Australia’.

Adelaide.

PRETTY, G. L. 1981. Trial excavation of an Aboriginal

camp site and site survey, Kinchega National Park.

Pp. 53-63 in ‘Darling Surveys, I’. Ed. J. M. HOPE.

Occasional Papers in Prehistory 3. Australian

National University: Canberra.

PRETTY, G. L. 1982. ‘The Archaeology of Roonka:

Examining the Human and Environmental History of

an Australian Place’. Paper read to the Nature

Conservation Society of South Australia, Adelaide.

23 pp., 20 illus.

PRETTY, G. L. 1982. Prehistory of the Australians: the

view from Roonka. University of Adelaide,

Department of Classics, Lecture Series-Archaeology

TH: 1-11.

PRETTY, G. L. (with R. C. PATON & R. D. J.

WEATHERSBEE). 1983. Tribal man. Pp. 115-126

in ‘Natural history of the South East of South

Australia’. Eds. M. J. TYLER, C. R. TWIDALE, J.

K. LING & J. W. HOLMES. Royal Society of South

Australia: Adelaide.

PRETTY, G. L (with T. J. GARA). 1983. Scientific

background. in ‘Roonka Conservation Park

Management Plan’. Ed. P. J. HUGHES. ANUTECH

Services: Canberra.

PRETTY, G. L. 1984. Review of John Aubrey (ed.),

‘Monumenta Britannica or a Miscellany of British

Antiquities (parts 1 & 2)’. Mankind 14, 3: 254-6.

PRETTY, G. L. 1986. The prehistory of South

Australia. Pp. 3-62 in ‘The Flinders History of South

Australia: Social History. 3’. Ed. E. RICHARDS.

Wakefield Press: Adelaide.

PRETTY, G. L. (with R. S. MERRILLEES & C. A.

HOPE). 1990. ‘Living with Egypt’s Past in

Australia’. Museum of Victoria: Melbourne. viii +

78 pp., illus.

PRETTY, G. L. 1990. The significance of the tombs of

evolved prehistoric hunters and gatherers; Southern

Australia and Southern France compared. (Invited

address, Royal Society of South Australia. May,

1990.) Newsletter, Royal Society of South Australia,

June. Pp. 4-7.

The following are in manuscript form and it is

intended that they will be published in due course.

PRETTY, G. L. ‘Archaeological Investigations at

Roonka, Part I - Description of the evidence gathered

from the Karmakalingbila Ngaiawang territory:

Geological and geochronological background;

cultural landscape and stratification; ecological,

demographic and cultural finds’. Text, tables, illus.

H. TOLCHER, J. PRESCOTT & J. MULVANEY

PRETTY, G. L. (with G. K. WARD) (eds).

‘Archaeological Investigations at Roonka, Part 2 -

Ethnoanalogical and comparative analysis:

Conjectures about territory, society and ideas in the

Karmakalingbila Ngaiawang territory in the light of

its Murray Darling Basin context’. (Accepted for

publication in British Archaeological Reports:

International Series.)

PRETTY, G. L. ‘Archaeological Investigations at

Roonka, Part 3 - Ethnodeductive and systemic

analysis: Conjectures about the mechanics and

dynamics of cultural change in_ the

Karmakalingbila Ngaiawang territory in the light

of its Central Southern Australian context’. Text,

tables, illus.

PRETTY, G. L. ‘Archaeological Investigations at

Roonka, Part 4 - Decoding the symbol inventory of

the Karmakalingbila Ngaiawang sequence: Restoring

immediate intelligibility and ultimate causal identity

to culturally unfamiliar modes of action through

appeals to phyletic behavioural repertoires’. Text,

tables, illus.

PRETTY, G. L. ‘Archaeological Investigations at

Roonka, Part 5 - The explanation of prehistoric

hunter-gatherer culture change in the

Karmakalingbila Ngaiawang territory during the

Holocene; tests of confirmation and concluding

synthesis’. Text, tables, illus.

Art

PRETTY, G. L. 1963. A turtle shell mask of Torres

Strait type in the Macleay Museum, University of

Sydney. Records of the South Australian Museum

14, 3: 421-425.

PRETTY, G. L. 1971. The National Collection of

Primitive Art, Canberra. Art and Australia 8, 4: 324-

329.

PRETTY, G. L. 1971. Comment on R. McGhee,

‘Differential artistic productivity in the Eskimo

cultural tradition’. Current Anthropology 17, 2: 216.

PRETTY, G. L. 1976. ‘Arts of Melanesia: Adelaide

Festival of Arts 1976’. (Exhibition Catalogue.) David

Jones Gallery: Adelaide.

Cultural heritage and public policy

PRETTY, G. L. 1970. ‘Report on the Commonwealth

Collection of Primitive Art from New Guinea and the

Pacific Islands’. Commonwealth Art Advisory

Board: Canberra. iv + 78 pp., illus.

PRETTY, G. L. 1971. Museums and controls on the

export of cultural property. Kalori: Journal of the

Museums Association of Australia 42: 89-95.

PRETTY, G. L. (General Editor). 1977. ‘National Folk

Province Feasibility Study Papers’. Commonwealth

Department for the Environment, Housing and

OBITUARY — GRAEME LLOYD PRETTY 101

Community Development and South Australian

Department for the Environment: Adelaide, as

follows:

No. 1 PRETTY, G. L. ‘The Ngaiawang Folk Province:

A Proposal’.

No. 2 PRETTY, G. L. ‘The National Folk Province

Concept - Specification for a feasibility study’.

No. 3 ROWNEY, B. G. ‘Kapunda - Historical

development of its town plan’.

No. 4 PRETTY, G. L. ‘The National Folk Province

Concept and the National Estate’.

No. 5 KALIBATAS, E. ‘Craigie’s Plains: Construction

and materials analysis of a historic vernacular sheep

station’.

No. 8 SPIERS, G. K. ‘Survey of Heritage Agencies in

South Australia’,

No 9 ROWNEY, B. G. ‘Surviving cottages in Mine

Square, Kapunda’.

No. 10 PRETTY, G. L. (with B. G. ROWNEY & P. F.

DONOVAN). ‘Ngaiawang Folk Province:

Preliminary inventory of the cultural landscape’.

No. 11 SPIERS, G. K. ‘The administration of heritage

properties through Local Government: I, The South

Australian Planning and Development Act 1996-

1975’.

No. 12 DONOVAN, P. F. & SCHEPPERS, K. H.

‘Report on the Kapunda Colonial Festival, 1976’.

No. 13 SPIERS, G. K ‘The administration of heritage

properties through Local Government: II, The

Municipal Heritage Trust’.

No. 15 DONOVAN, P. F. ‘Woakwine Folk Province:

Outline sketch of cultural landscape’.

No. 16 DONOVAN, P. F. & SCHEPPERS, K. H. ‘The

administration of heritage property through Local

Government: III, Report from the Heritage Officer,

District Council of Kapunda’.

No. 19 ROWNEY, B. G. & DONOVAN, P. F. ‘Old

Legislative Council Building, Adelaide : An

architectural assessment’.

No. 20 ROWNEY, B. G. & DONOVAN, P. F.

‘Establishing conservation priorities for the built

environment: The example of Kapunda’.

Ethnography

PRETTY, G. L. 1969. The Macleay Museum mummy

from Torres Straits; a postscript to Elliot Smith and

the diffusion controversy. Man: the Journal of the

Royal Anthropological Institute NS 4, 1: 24-43.

PRETTY, G. L. 1969. ‘Salvage Ethnography in New

Guinea: the South Australian Museum Expedition to

the Southern Highlands District, Papua, 1968-1969’.

Report to the Wenner-Gren Foundation for

Anthropological Research, New York. iv + 85 pp.,

illus.

PRETTY, G. L. 1970. Aboriginal Studies. (Review of

‘Man, Land and Myth in North Australia: the

Gunwinggu people’ by R. M. & C. H. Berndt ;

‘Yiwara-Foragers of the Australian Desert’ by

Richard A Gould.) Australian Book Reviews 9, 12:

332-334.

PRETTY, G. L. 1974. Comment on Francis L. Utley,

‘The migration of folk tales: four channels to the

Americas’. Current Anthropology 15, 1: 19-20.

PRETTY, G. L. 1980. The Aboriginal cultural

landscape of the Lower Murray Valley. Pp. 43-52 in

‘Preserving Indigenous Cultures; a New Role for

Museums’. Eds. R. Edwards & J. Stewart. UNESCO

and Australia Council: Canberra.

PRETTY, G. L. (with A. CALDER). 1980.

Mummification in Australian and Melanesia.

Pp. 194-210 in ‘Mummies, Disease and Ancient

Cultures’. Eds. A. & E. Cockburn. Cambridge

University Press: New York.

PRETTY, G. L. and course participants, Visual Arts

Discipline, Flinders University for the Gerard

Aboriginal Community. 1983. ‘Artifacts of the

Ngaiawang Tribe, Lower Murray Valley, South

Australia, obtained by E. J. Eyre (1841-1844)’.

(Document basis for the renewal of the Gerard

Community artifact industry.)

PRETTY, G. L. 1993. The Aborigines of the South East

of South Australia and the Western District of

Victoria before white contact. Community History

(Adelaide) 3, 4: 2-5.

Evolutionary biology

PRETTY, G. L. (with P. SMITH & M. PROKOPEC).

1988. Dentition of a prehistoric population from

Roonka Flat, South Australia. Archaeology in

Oceania 23: 31-36.

PRETTY, G. L. (with M. PROKOPEC & P. SMITH).

1994. Prehistoric South Australian Aboriginals: New

evolutionary perspectives. Variability and Evolution

(Adam Mickiewicz University, Poznan) 4: 21-56.

Forensic anthropology

PRETTY, G. L. 1975. The recovery of human remains

for forensic purposes. Proceedings Australian

Forensic Science Society 1, 3: 68-74.

PRETTY, G. L. (with D. J. POUNDER & M.

PROKOPEC). 1983. A probable case of euthanasia

amongst prehistoric Aborigines at Roonka, South

Australia. Forensic Science International 23: 99-

108.

PRETTY, G. L. (with M. PROKOPEC, D. SIMPSON

& L. MORRIS). 1984. Craniosynostosis in a

prehistoric Aboriginal skull: A case report. Ossa.

International Journal of Skeletal Research 9-11:

111-118.

102

Heritage preservation

PRETTY, G. L. 1967. ‘Preliminary catalogue of

archaeological monuments in the Territory of Papua

New Guinea’. Papua New Guinea Public Museum

and Art Gallery: Port Moresby. iv + 95 pp., map.

PRETTY, G. L. 1968. Surveying archaeological

monuments in Papua New Guinea. Journal of the

Anthropological Society of South Australia 6, 2:

5-8.

PRETTY, G. L. 1970. Survey of prehistoric monuments

in South Australia. Pp. 39-50 in ‘Aboriginal

antiquities in Australia: their nature and

preservation’. Ed. F. D. McCarthy. Australian

Institute of Aboriginal Studies 22.

PRETTY, G. L. 1976. The Ngaiawang Folk Province.

Journal of the Anthropological Society of South

Australia 14, 9: 13-22.

PRETTY, G. L. 1978. The Ngaiawang Folk Province.

Pp. 28-45 in ‘The Tide of Australian Settlement —

Conservation of the Physical Evidence’. Australia

ICOMOS Proceedings, Beechworth, 14-16 April

1978. Australia/ICOMOS: Hawthorn, Vic.

History

PRETTY, G. L. 1967. Edward Gibbon Wakefield.

Pp. 559-562 in ‘Australian Dictionary of

Biography’. Ed. D. Pike. Melbourne.

PRETTY, G. L. 1975. The historical collections

in the South Australian Museum. Pp. 173-176

in ‘Proceedings of the National Seminar on the

Conservation of Cultural Material, Perth,

August 1973’. Eds C. Pearson & G. L. Pretty).

Institute for the Conservation of Cultural

Material: Perth.

PRETTY, G. L. 1986. Australian history at Roonka.

Journal of the Historical Society of South Australia,

14: 107-122.

PRETTY, G. L. 1986. Foreword to ‘People, Places and

Buildings’. J. Faull & G. Young. South Australian

Centre for Settlement Studies: Adelaide.

Materials conservation

PRETTY, G. L. (with C. PEARSON). (Eds). 1976.

‘Proceedings of the National Seminar on the

Conservation of Cultural Material, Perth, August

1973’. Institute for the Conservation of Cultural

Material: Perth. 297 pp., illus.

Museology

PRETTY, G. L. 1967. The British Museum. Friends of

the South Australian Museum Newsletter 4: 9-11.

PRETTY, G. L. 1968. ‘Report on the State of the Papua

New Guinea Museum: the Collections of

Archaeology and Ethnology’. Trustees, Papua New

H. TOLCHER, J. PRESCOTT & J. MULVANEY

Guinea Public Museum and Art Gallery: Port

Moresby. iv + 95 pp., illus.

PRETTY, G. L. 1971. Oceanic museum resources in

Australian museums. Pp. 179-186 in ‘Meeting on

Studies of Oceanic Cultures’. Australian National

Advisory Committee for UNESCO: Canberra.

PRETTY, G. L. 1976. The Ethnographic Collections in

the South Australian Museum. Pp. 107-119 in

‘Proceedings, National Seminar on Conservation of

Cultural Material’. Eds. C. Pearson & G. L. Pretty.

Institute for the Conservation of Cultural Material:

Perth.

Palaeoecology

PRETTY, G. L. (with W. E. BOYD). 1989. Some

prospects for archaeological palaeobotany in

Australia; an example from South Australia.

Australian Archaeology 28: 40-52.

PRETTY, G. L. (with W. E. BOYD). (in press). From

the sensible to the spiritual — Cultural signals from

palaeobotanic evidence at Roonka, South Australia.

Archaeological Dialogues.

Palaeopathology

PRETTY, G. L. (with M. E. KRICUN). 1989. Health

status of the prehistoric Roonka population. World

Archaeology 21, 2: 198-224.

PRETTY, G. L. (with M. PROKOPEC). 1990.

Skeletal aging. Rate of tooth attrition in hunter-

gatherer populations, prehistoric Roonka, South

Australia. Colloquiae Anthropologicae 14, 2: 331-

334.

PRETTY, G. L. (with M. PROKOPEC). 1991.

Observations on health, genetics and culture from

analysis of prehistoric population data from Roonka,

South Australia. Pp. 151-158 in ‘Human

Palaeopathology: Current Syntheses and Future

Options’. Eds. D. J. Ortner & A. C. Aufderheide.

Smithsonian Press: Washington, DC.

PRETTY, G. L. (with T. BROWN & M. E. KRICUN).

1992. Extensive compensatory remodeling of

craniofacial structures; a case from prehistoric South

Australia. Pp. 139-143 in ‘Craniofacial Variation in

Pacific Populations. Papers presented at the XVII

Pacific Science Congress Symposium, Honolulu,

Hawaii, May 30, 1991’. University of Adelaide:

Adelaide.

PROKOPEC, M. and PRETTY, G. L. 1994.

Praehistorische Bewohner der Roonka-Siedlung

(Sudaustralien). Ethnographisch-Archaeologische

Zeitschrift 35, 1: 136-148.

SIMPSON, D., PROKOPEC, M., MORRIS, L., &

PRETTY, G. L. 1984. Prehistoric craniosynostosis.

A case report. Records of the Adelaide Children’s

Hospital 3, 2: 163-168.

OBITUARY —- GRAEME LLOYD PRETTY 103

Radiometric dating

PRETTY, G. L. (with J. R. PRESCOTT, H. POLACH

& B. W. SMITH). 1983. Some comparisons of C-14

and thermoluminescent dates from Australia.

(Proceedings of the 1981 Groningen C-14

Conference). Council of Europe PACT Journal 8:

205-211.

PRETTY, G. L. 1988. Radiometric chronology and

significance of the fossil hominid sequence from

Roonka, South Australia. Pp. 32-52 in ‘Early Man in

the Southern Hemisphere’. Ed. J. R. Prescott.

Supplement to Archaeometry: Australasian Studies.

University of Adelaide Department of Physics and

Mathematical Physics: Adelaide.

Technology

PRETTY, G. L. 1970. Review of ‘Excavations of the

prehistoric iron industry in West Borneo’ by T.

Harrisson & S. J. O'Connor. Man: The Journal of

the Royal Anthropological Institute, London. NS. 5:

710-711.

Trace element chemistry

PRETTY, G. L. 1991. (with F. D. PATE, J. T.

HUTTON & R. A. GOULD). Alterations of in vivo

elementary dietary signatures in archaeological bone.

Evidence from the Roonka Flat Dune, South

Australia. Archaeology in Oceania 26: 58-69.

Helen Tolcher, c/- South Australian Museum, North Terrace, Adelaide, South Australia 5000; Professor Emeritus

John Prescott, Physics Department, Adelaide University, South Australia 5005; and Professor Emeritus John

Mulvaney, Centre for Archaeological Research, Australian National University, ACT 0200. Records of the South

Australian Museum 35(1): 97-103.

IRNIECOIRIDS

OJF

plpteiils

SOUTH

AUSTRALIAN

MUSEUM

VOLUME 35 PART 1.

MAY 2002

ISSN 0376-2750

CONTENTS:

oF.

ARTICLES

I. BEVERIDGE

New species and new records of Cloacina von Linstow, 1898 (Nematoda: Strongyloidea)

parasitic in macropodid marsupials from Papua New Guinea.

B. P. KEAR

Darwin Formation (Early Cretaceous, Northern Territory) marine reptile remains in the South

Australian Museum.

H. W. PALM & I. BEVERIDGE

Tentaculariid cestodes of the order Trypanorhyncha (Platyhelminthes) from the

Australian region.

E.G. MATTHEWS & T. A. WEIR

Two new species of the genus Lepanus Balthasar from South Australia (Coleoptera:

Scarabaeidae).

S. BARKER

A checklist of Cisseis (sensu stricto) Gory & Laporte, 1839 (Coleoptera: Buprestidae:

Agrilinae).

W. ZEIDLER

The Flora and Fauna of South Australia Handbooks Committee 10 March 1921 —

30 October 2001.

NOTE

H. TOLCHER, J. PRESCOTT & J. MULVANEY

Obituary — Graeme Lloyd Pretty

Published by the South Australian Museum,

North Terrace, Adelaide, South Australia 5000.

INE CORDS

Ole

Ie0e,

SOUTH

~ AUSTRALIAN

MUSEUM

VOLUME 35 PART 2

OCTOBER 2002

THE LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE

(COLEOPTRA : INSECTA)

C. H. S. WATTS

Summary

The larvae of the aquatic Hydrophilid genera Allocotocerus Kraatz, Hybrogralius Orchymont and

Regimbartia Zaitzev are described and figured for the first time.The larvae of the following

Australian species are also described and figured, most for the first time: Allocotocerus punctatus

(Blackburn); Amphiops aueenslandicus Balfour-Browne; Berosus australiae Mulsant; Enochrus

eyrensis (Blackburn), E. mastersi (W. MacLeay); Helochares tristis (W. MacLeay), H. clypeatus

(Blackburn), H. tenuistriatus Regimbart, H. foveicollis (Montrouzier); Hydrobiomorpha sp. ;

Hydrophilus bilineatus (MacLeay); Hybrogralius hartmeyeri (Regimbart); Laccobius decipiens

Gentili; Limnoxenus zealandicus (Broun); Regimbartia attenuata (Fabricius); and Sternolophus

margincollis (Hope).

THE LARVAE OF SOME AUSTRALIAN AQUATIC

HYDROPHILIDAE (COLEOPTERA: INSECTA)

CHS WATTS

WATTS, CHS. 2002. The larvae of some Australian aquatic Hydrophilidae (Coleoptera:

Insecta). Records of the South Australian Museum 35(2): 105-138.

The larvae of the aquatic Hydrophilid genera Allocotocerus Kraatz, Hybogralius Orchymont

and Regimbartia Zaitzev are described and figured for the first time. The larvae of the

following Australian species are also described and figured, most for the first time:

Allocotocerus punctatus (Blackburn); Amphiops queenslandicus Balfour-Browne; Berosus

australiae Mulsant; Enochrus eyrensis (Blackburn), E. mastersi (W. MacLeay); Helochares

tristis (W. MacLeay), H. luridus (W. MacLeay), H. clypeatus (Blackburn), H. tenuistriatus

Regimbart, H. foveicollis (Montrouzier); Hydrobiomorpha sp.; Hydrophilus bilineatus

(MacLeay); Hybogralius hartmeyeri (Regimbart); Laccobius decipiens Gentili; Limnoxenus

zealandicus (Broun); Regimbartia attenuata (Fabricius); and Sternolophus marginicollis

(Hope).

The larvae were identified by rearing larvae collected in the field, or occasionally by

association and elimination, or by using biochemical methods.

The pupae of Amphiops queenslandicus and Hybogralius hartmeyeri are described and

figured. Unlike most other Hydrophilids the larvae of Amphiops are thought to pupate exposed

on the stems of emergent vegetation.

A key is provided to the larvae of Australian genera of aquatic Hydrophilids other than

Notohydrus Balfour-Browne, Paranacaena Blackburn, Phelea Hanson and Agraphydrus

Regimbart, whose larvae remain unknown.

CHS Watts, South Australian Museum, North Terrace, Adelaide, South Australia 5000.

Manuscript received 23 April 2001.

The Hydrophilidae (sensu Hansen 1991)

comprise a major part of the Australian aquatic

beetle fauna, occurring in all types of fresh water

and most commonly among vegetation at the edge

of standing water. Adults and larvae occur

together. The taxonomy of the adults is now

relatively well known thanks to the work of

Hansen (1991) at the generic level and above; and

that of Gentili (1980, 1992, 1993, 2000) and

Watts (1987, 1988, 1989, 1990, 1995, 1998a,

1998b) on the aquatic fauna at the species level.

No recent work has been done on the species level

taxonomy of the extensive portion of the family in

which both adults and larvae are terrestrial.

In contrast to the situation with adults, the only

work specifically on Australian larvae is that of

Anderson (1976) who described the larvae of

Helochares tristis (W. MacLeay), Enochrus

maculiceps (W. MacLeay) and Chasmogenus

nitescens (Fauvel) together with details of their

life histories.

One feature of the Australian aquatic

Hydrophilid fauna is its low endemicity at the

generic level. Consequently, descriptions of a

number of Australian genera are available in

scattered Northern Hemisphere publications. The

work on the New World genera was summarised

and added to by Archangelsky (1997). This work

included a number of genera which also occur in

Australia.

For a number of years I have been rearing field-

collected larvae of Australian aquatic

Hydrophilids with the aim of discovering and

describing the larvae of all genera and, for those

already known from non-Australian species, to

extend the descriptions to incorporate Australian

species. The basic motivation was to produce a

key to enable the larvae of all Australian aquatic

Hydrophilidae to be identified at least to genus.

As a result I have reared 12 species in 9 genera.

Two additional genera, Allocotocerus and

Regimbartia, were identified by associating adults

and larvae by the biochemical technique of

alloenzyme electrophoresis. Others were identified

using the descriptions in Archangelsky (1997), eg

Hydrobiomorpha;, or by association and

elimination, eg Enochrus mastersi.

I have not attempted any phylogenetic analysis

106

since, with such a high proportion of genera also

occurring widely outside Australia, the only

sensible approach would have been to include an

examination of all known genera, which was

beyond the scope of this study. Some comments

on relationships are given under most generic

descriptions.

In preparing this paper I have drawn heavily on

the work of Archangelsky (1997) on the New

World fauna. The descriptions follow his format

and a number of the figures used to illustrate the

key come from his important publication.

Although their larvae are known from other

places, I have not seen Australian examples of the

following genera: Chaetarthria Stephens,

Chasmogenus Sharp, Crenitis Bedel, Coelostoma

Brulle or Paracymus Thomson, and since I have

nothing to add have not included descriptions.

Detailed descriptions can be found in

Archangelsky (1997). In addition, the larvae of

Paranacaena Blackburn, Notohydrus Balfour-

Browne, Phelea Hanson and Agraphydrus

Regimbart are unknown.

MATERIALS AND METHODS

Larvae were collected in the field and reared in

small aquaria formed by placing the base of a

small petrie-dish inside a larger petrie-dish and

filling the space between them with damp sand

(Watts 1963). Mosquito and chironomid larvae

were the main food items provided. These were

accepted by all larvae other than those of

Hydrophilus (see under Hydrophilus). All species

successfully reared pupated in cells constructed in

the wet sand with the exception of those of

Amphiops which pupated on the surface or

attached to the sides of the container.

Larvae were preserved in 75% ethanol and

examined under a stereomicroscope. Permanent

microscope slides were prepared for detailed

examination. Drawings were made with the aid

of a camera lucida. Habitus drawings were only

prepared when no previous illustration was

available or when Australian species differed

from those previously illustrated. Unless

otherwise mentioned the descriptions are based

on third instar larvae. Earlier instars usually

differ somewhat, most noticeably in the stouter

cephalic appendages and in fewer spines on the

mentum. The key is constructed to accommodate

all instars. Measurements of the head capsule

were made under a stereoscopic microscope with

a graduated eyepiece. Measurements of total

CHS WATTS

length were made with dial callipers on both

mounted and alcohol preserved material, but in

this case the measurements can only be

indicative as factors such as larval age and

preservation effects can alter the length

considerably. Measurements of the head capsule,

particularly the width, are a more reliable

indicator of size. Unless stated otherwise, the

specimens were collected by myself and are in

the collection of the South Australian Museum.

SYSTEMATICS

KEY TO THE GENERA OF AUSTRALIAN

AQUATIC HYDROPHILIDAE BASED ON

LARVAL CHARACTERS

1) Not including the genera Notohydrus,

Paranacaena, Phelea and Agraphydrus,

whose larvae are not known.

2) Including Spercheus which is in the family

Spercheidae but which has larvae that can be

confused with Hydrophilidae.

1 — _ Tip of mandible bifid (Fig. 2g) .............

yderkeyedaapeigegi ok Spercheidae (Spercheus)

— _ Tip of mandible not bifid .....................

ceuetesieeeten Ae gas Hydrophilidae. ..2

Hypopharyngeal lobe well developed, like

a pubescent tongue, originating at the base

of the labium on the left side (Figs 2a, 2b)

Subfamily Sphaeridiinae* .......0..00.....

bupsisatosdyestosissssparsorsee Coelostoma Brulle

— Hypopharyngeal lobe reduced or absent

paths tinahabet Subfamily Hydrophilinae...3

Abdominal segments with long

filamentous gills (Fig. Sa), or with multiple

setose lateral projections on abdominal

and thoracic segments (Fig. 15a) ..........

pesdboodEodnsedaacds sthabharaee Tribe Berosini...4

— Lacking long filamentous abdominal gills

or complex lateral projections, at most

with simple short fleshy projections (Figs

4a, 14a, 16a) oo. eecccseeeesesseeseessenes 6

Gills long, without setae (Fig. 5a);

labroclypeus asymmetric (Fig. 5b);

mandibles asymmetric (Fig. Sf);

prementum small and squat, without ligula

(Fig. 5c), basal segment of antenna without

lateral projection (Fig. Se) 0... eee eee

SinacaSeaap evleny HHitepavier abun 23 Berosus Leach

— Abdominal and thoracic segments each

with two or more setose, lateral projections

3 =

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 107

(Fig. 15a); mandibles nearly symmetrical

(Fig. 15h); prementum long and thin, with

ligula (Fig. 15d); basal segment of antenna

with lateral projection (Fig. 15g) ........ 5

Body spines predominantly blunt, weakly

bifid at tips (Fig. 3b); lateral projection on

basal segment of antenna near apex, sharp,

spine-like (Fig. 3f); apex of second segment

of antenna with lateral extension (Fig. 3d);

maxillary stipe without spines (Fig. 3e);

ligula relatively short, prementum stouter

(Fig) 36) <....0-1.. Allocotocerus Kraatz

Body spines predominantly pointed (Fig.

15c); lateral projection on basal segment

of antenna thumb-like, further from apex

(Fig. 15g), second segment of antenna

without lateral extension (Fig. 15e);

maxillary stipe with spines on inside (Fig.

15f); ligula relatively long, prementum

more elongate (Fig. 15d)... ee

shes ebdgebibetnadsatentcbedeas Regimbartia Zaitzev

Apical segment of antenna about same

length as penultimate (Figs 12d, 16e);

femora with fringe of swimming-hairs (Fig.

2h); with prostyles (two retractable, fleshy

appendages on last abdominal segment)

se Dred aL Subtribe Hydrophilina. ..7

Apical segment of antenna much shorter

than penultimate (Figs 9e, 14e); femora

lacking, or virtually lacking, swimming-

hairs; without prostyles ...........-2eee 9

Labroclypeus without teeth (Fig. 12a);

mandibles asymmetrical, left one very

robust, right one more slender (Fig. 12e);

ligula shorter than first segment of labial

palpus (Fig. 12b); up to 40 mm long ....

RENE are cet ae Hydrophilus Muller

Labroclypeus with weak to moderate teeth

(old specimens may lack teeth due to wear)

(Figs lle, 16f); mandibles symmetrical or

nearly so; ligula longer than first segment

of labial palpus (Figs 11b, 16c); up to

20am long. nag oe 8

Prementum longer than mentum (Fig. 16c);

first segment of antennae with few spines

(Fig. 16f); mandibles each with two large

distal teeth and one small proximal denticle

(Fig. 16f) 0... Sternolophus Solier

Prementum slightly shorter than mentum

(Fig. 11b); first segment of antenna with

numerous spines on inside (Fig. 11c);

mandibles each with one large distal tooth

which is pick-shaped at the tip and one or

two small proximal denticles (Fig. 1 le)

wacedeaeseetatins Hydrobiomorpha Blackburn

Left expansion of epistoma much more

prominent than the right and with a row of

stout setae on front edge (Fig. 13b); left

mandible with group of stout setae at base

of middle tooth which is lacking on the

right mandible (Fig. 13f) .......... 10

Lateral expansions of epistoma similar,

without a row of stout setae on front edge

(Fig. 14b); mandibles without sucha group

C0) Bo] Fo 11

Frontal sulci parallel and not uniting to

form a coronal sulcus (Fig. 13a); nasale

with three teeth (Fig. 13b); prementum

wider than long (Fig 13c); ligula absent or

Virtually so (Fig. 13C) oo. sceeseeeeeeeeeeeee

pabbadeanetpragaresiitealaryeas Laccobius Erichson

Frontal sulci meet just before occipital

foramen to form a coronal sulcus (Fig.

10a); nasale with five teeth (Fig. 10b);

prementum longer than wide (Fig. 10c);

ligula present though small (Fig. 10c) ..

ebagagasneaterpetast ete Hybogralius Orchymont

Legs very short, without claws, prementum

with a large round ligula (Fig. 2e); length

up to 6 mm......... Chaetarthria Stephens

Legs usually longer, with claws.

Prementum with or without ligula, but if

present never large and round; length up to

USS TO peisk easy oa sca egeeesecbectveeebabeate 12

Mentum large, wider than long, prementum

small, lacking ligula (Fig. 4b) ..............

Be ceadpomedd st tshe heen dteg’ Amphiops Erichson

Mentum square or longer than wide,

prementum well developed, ligula present

(Figs 6b, 8b, 14c) .... eee ecceeeeeeeeee 13

All abdominal segments with dorsal

sclerites, although often small; without

coronal sulcus (Fig. 2c); lengthupto6 mm;

antennal appendage as long as apical

segment; mandibles symmetrical with three

teeth. stone Tribe Anacaenini....14

Abdominal segments, other than 1 and 8,

lacking dorsal sclerites; with short coronal

sulcus (Fig. 1a); antennal appendage half

length of apical segment (Fig. 8d);

mandibles often asymmetrical and with

fewer than three teeth; length up to 15 mm

wigan Fes 5 aaa sean pate sev pstegesttupedinateeg’ 15

With lateral abdominal flaps (Fig. 2c);

without cervical sclerites Crenitis Bedel

108 CHS WATTS

FIGURE 1. a, dorsal side of head capsule of Hybogralius hartmeyeri. 1, maxillary palpus; 2, stipes of maxillary

palpus; 3, mandible; 4, mandibular teeth; 5, labium; 6, antenna; 7, right lobe of epistome; 8, teeth of nasale; 9,

nasale (7 & 9 make up the labroclypeus); 10, stemmata; 11, frontal sulci; 12, coronal sulcus; 13, cervical sclerites;

14, occipital foramen; b, details of dorsal side of labium of Hybogralius hartmeyeri. 1, labial palpus; 2, ligula; 3,

prementum; 4, mentum.

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 109

S <f\ MM Ang

FRA

omens oy,

we Ate.

le ee

h i

FIGURE. 2. a, ventral view of labium and hypopharyngeal lobe (arrowed) of Sphaeridium scarabaeoides

(Linnaeus) (subfamily Sphaeridiinae); b, dorsal view of labium and hypopharyngeal lobe (arrowed) of

Dactylosternum sp. (subfamily Sphaeridiinae); c, habitus of Crenitis morata (Horn); d, ditto, dorsal view of

labium; e, ditto of Chaetarthria sp.; f, ditto, Paracymus subcupreus (Say); g, mandible of Spercheus platycephalus

MacLeay; h, hindleg of Hydrobiomorpha sp., Northern Territory; i, labroclypeus and mandible of Chasmogenus

nitescens (Fauvel). a-f, North American taxa from Archangelsky 1997; i, from Anderson 1976.

110

— Lacking abdominal flaps; with small

cervical sclerites . Paracymus Thomson

15 — Apical segment of antenna about half

length of penultimate (Fig. 14e); mandibles

symmetrical with three teeth, without

serrations (Fig. 14f) ........ceeeeeeeeeeeeee

achessegng tadingsdared Limnoxenus Motschulsky

— Apicalsegment of antenna about a quarter

the length of the penultimate segment

(Fig. 8d); mandibles weakly to strongly

asymmetric, with one or two teeth, often

with serrations (Figs 6e, 8e) ............0.

Revvonspaperetact Subtribe Acidocerenia....16

Nasale moderately developed, with right

side more prominent (Fig. 6a) .......... 17

— Nasale poorly developed, all areas equally

prominent (Fig. 9b) oo... eects 19

Mandibles asymmetric, the right one

with one tooth, left with two (Fig. 6e)

asasadganbzegteabaesupeaesaseareosassast atte: Enochrus

Thomson (other than E. mastersi)

— Mandibles almost symmetrical, both with

two teeth (Fig. 8€) .....ceeceeeesseeeeeereee 18

Inside of maxillary stipe with group of

spines near base (Fig. 8c); ligula longer

than first segment of labial palpus (Fig.

8b); prementum approximately the same

length as mentum (Fig. 8b)....Helochares

Mulsant (other than H. foveicollis)

— Inside of maxillary stipe lacking such

spines (Fig. 7c); ligula shorter than first

segment of labial palpus (Fig. 7b);

prementum approximately twice length of

mentum (Fig. 7b) ......ccecseeesseeeteeeseeeees

divasgersegsevecdtt E. mastersi (W. MacLeay)

First segment of antenna with bulge on

inside near apex (Fig. 9e); mandibles each

with two teeth (Fig. 8f) ..... cesses

oben ans H. foveicollis (Montrouzier)

— First segment of antenna without bulge;

right mandible with one tooth, left one

with two (Fig. 21) oo... eeeeeeeseeeeeeeneeeeee

16 —

17 —

18 —

* Members of the subfamily Sphaeridiinae are

predominantly terrestrial, occurring in moist

situations such as dung and rotting vegetation.

Members of only one genus, Coelostoma, are

commonly found in aquatic situations in

Australia.

CHS WATTS

DESCRIPTIONS

The following descriptions are arranged in

alphabetic order of the genera.

Allocotocerus Kraatz

Allocotocerus punctatus (Blackburn) Fig. 3

Size of third instar. Length 8.5—12.0 mm; head

capsule, 0.70—-1.00 mm long, 0.85—1.0 mm wide.

Head capsule. Subquadrangular, bulging

outwards at anterolateral angles. Labroclypeus

symmetrical; nasale very short, with numerous

very short teeth on anterior border of epistome;

lateral lobes of epistome rounded, very short.

Frontal sulci straight, reaching from antennal

bases to rear of head. Coronal sulcus absent. Gular

sclerite absent. Cervical sclerites small,

subrectangular, very weakly sclerotised.

Antennae. Three-segmented. First segment

slender, longer than other two combined, with a

spine-like inner process close to apex; third

antennal segment short, connected to edge of the

second segment before apex. Sensory appendage

on second segment, slim, shorter than third

segment.

Mandibles. Symmetrical, long and slender, with

two inner teeth on basal half; distal tooth large,

basal one small.

Maxillae. Five-segmented. Stipes slender, much

longer than remaining segments combined, with

two or three fine setae on inner margin. Palpus

four- segmented; first and third segments subequal

in length, first segment bearing a short inner

process; second segment very short; fourth

segment shorter than third.

Labium. Mentum large, subglobular, dorsal

surface with some cuticular spines towards sides,

anterolateral angles rounded with a few spines.

Prementum much longer and narrower than

mentum, subrectangular, longer than wide. Palpus

two-segmented, basal segment short. Ligula about

two times as long as basal segment of palpus, tip

broadened.

Thorax and legs. Prothorax with broad dorsal

sclerite, with sagittal line, surface covered by

minute spines, fringe of long, slender setae around

the margins and three short setae-bearing

projections at sides; ventral surface with large,

subrectangular sclerite, without sagittal line.

Mesothorax with two dorsal pairs of subtriangular

sclerites. Metathorax with one dorsal pair of

irregular shaped sclerites. Both mesothorax and

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 111

FIGURE. 3. Allocotocerus punctatus. a, labroclypeus; b, detail of abdominal stella; c, labium; d, tip of antenna; e,

maxilla; f, antenna; g, mandibles.

112

metathorax with three lateral pairs of setiferous

projections and a number of short setiferous

papillae on dorsal surface. Legs five-segmented,

long, visible from above.

Abdomen. Segments | to 7 with four pairs of

setiferous projections similar to those on thorax,

three on the side and a smaller dorsal one

towards the midline. In addition, the abdomen is

covered with minute, simple spines and the

dorsal surface with small projections, each with

six to eight, relatively long setae, truncated or

weakly bifid at tips (in most specimens these

structures are obscured by an accumulation of

sand grains and detritus). Segment 8 without

setiferous projections; with dorsal semispherical

sclerite covering the spiracular atrium; with a

pair of small finger-like ‘procerci’. Segment 9

trilobed, with a pair of small, unsegmented

urogomphi.

Spiracles. Mesothorax and abdominal segment

2 with pair of small non-functional spiracles on

short papillae. (I have been unable to detect

spiracles on other abdominal segments.) Spiracles

on segment 8 annular, large and functional, within

the spiracular atrium.

Interspecific variation

There is little variation among the few

specimens seen — including one from the

Northern Territory which is almost certainly A.

tibialis (Balfour-Browne).

Identification

By association of adult and larva by isoenzyme

electrophoresis.

Remarks

The larvae closely resemble those of the New

World genus Derallus (Archangelsky 1997;

Spangler 1966) in the setiferous body structures,

form of the labroclypeus, mandibles, labium,

maxillary palpi and antennae. The differences are

small: blunt rather than sharp-pointed body spines,

shape of the distal part of the antennae, number of

spines on the maxillary palpi and details of the

labium. The larva of Allocotocerus is also closely

similar to those of Regimbartia, differing from

this genus in the details of the same suite of

characters. On balance it more closely resembles

Derallus. Although placed by Hansen (1991) with

Berosus in the tribe Berosini, on larval characters

the three genera form a very distinctive and

cohesive group well separated from Berosus, a

conclusion reached by others (Bertrand 1972;

Oliva 1992; Spangler 1966).

CHS WATTS

Specimens examined

Queensland: 2km S Mt Molloy, 30/3/96.

Allocotocerus sp. Northern Territory: Manton

Dam, 23/3/98.

Amphiops Erichson

Amphiops queenslandicus Balfour-Browne. Figs

4, 17a

Size of third instar. Length 6.5—9.0 mm; head

capsule 0.65—0.80 mm long, 0.78-0.82 mm wide.

Head capsule. Subquadrate. Labroclypeus

symmetrical; nasale very short, with five small

teeth; lateral lobes of epistome rounded,

projecting further than nasale. Frontal sulci

inversely bell-shaped, meeting before reaching

occipital foramen. Coronal sulci very short. Gular

sclerite absent. Cervical sclerites small,

rectangular.

Antennae. Three-segmented. First segment as

long as second; third half the length of second.

Sensory appendage on second segment short, a

quarter the length of the third segment.

Mandibles. Symmetrical with three inner teeth,

the two distal teeth large and with slightly

crotchet-shaped tips; third tooth much smaller.

Maxillae. Five-segmented. Stipes narrow,

longer than the remaining segments combined,

with a row of five to six small setae on inner

margin. Palpus four-segmented; first segment

elongate, rectangular, with inner process as long

as very short second segment; third segment

longer and narrower than first; fourth segment a

little longer than second.

Labium. Mentum large, subrectangular, anterior

edge with five to six large spines/protuberances,

and numerous cuticular spines at sides towards

base. Prementum ovoid, much smaller than

submentum. Palpus short, two-segmented, basal

very short, distal segment two to three times

longer than basal one. Ligula absent.

Thorax and legs. Pro-, meso- and metasterna,

almost completely covered by dorsal sclerites,

each with a sagittal line. Ventral surface of

prothorax with a subrectangular sclerite,

subdivided by a sagittal line. Sclerites with very

small spines arranged in short lines towards rear

(only visible under high magnification). Bases of

setae on sclerites pigmented. Legs five-segmented,

moderately developed, visible in dorsal view.

Dorsal surface with sparse covering of small dark

patches.

Abdomen. First segment with dorsal pair of

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 113

e f

FIGURE 4. Amphiops queenslandicus. a, habitus; b, labroclypeus; ¢, labium; d, maxilla; e, antenna; f, mandible.

moderate sized subtriangular sclerites, segments 2

to 5 with smaller dorsal sclerites, sclerites with

scattered small darker patches. Segments 1 to 7

similar in size and shape, subdivided by a

transverse fold; pleural areas with four pairs of

moderate bulges/lobes each with several long

setae at apex, ventral one largest; with a pair of

small dorsal papillae, each with long terminal

setae, abdominal surface strongly folded. Segment

8 with pair of small ‘procerci’, with a large dorsal,

suboval sclerite; segment 9 trilobed, with a pair of

small one-segmented urogomphi. Integument with

only a few small spines.

Spiracles. Eight abdominal pairs; pairs 1 to 7 non-

functional, on tips of small papillae and each with a

long seta immediately inwards on a small cylinder-

shaped base. Spiracles on segment 8 annular, large

and functional, within the spiracular atrium.

Pupa. (Fig. 17a)

Colour. Light to dark brown.

Head. With one pair of supraorbital styli; styli

without apical setae. Appendages adpressed

tightly to body, weakly differentiated externally.

Thorax. Pronotum with 16 styli, all on margins

of pronotum. Mesothorax lacking obvious styli;

metanotum with two pairs of styli, lateral ones

near the base of wingpads very small, central pair

prominent. Styli without setae. Wing cases

grooved/ridged. Legs held tightly against body

almost totally covered by pterothecae.

114

Abdomen. Segments 1 to 7 with a transverse

row of four to five very small tergal styli on a

sharp ridge that is stronger laterally. Segments 1

to 7 with well-developed rigid flap-like lateral

extensions, those on segments | to 3 upright and

slightly bent over inwards, those on segments 4 to

7 horizontal and each with a small stylus on

posterior edge, each stylus with a small seta.

Anterior edge of first tergite with very prominent,

thin, rigid ridge. Segment 8 without obvious styli;

segment 9 bearing a pair of medium-sized cerci.

Interspecific variation

There is little variation among the specimens of

Amphiops available, although it is almost certain

that several species are included. There are some

differences, namely: the extent of the dorsal small

dark patches/spots varies from a few to moderate

numbers; the area between the abdominal sclerites

and spiracles sometimes is a bit darker than rest;

and there is some variation in the width of the

nasale.

Identification

By rearing larvae collected in the field.

Remarks

Bertrand (1972) keyed out Amphiops larvae and

briefly illustrated a presumed larva. Australian

species appear to have less well-developed lateral

abdominal protuberances than the specimen from

the Indonesian Archipelago (Insulinde) illustrated

by Bertrand (1972). Berge Henegouwen (1982)

described an African species as having short

conical outgrowths.

The noticeably lumpy sides to the abdomen, the

very small prementum and labial palpi, and the

lack of a ligula most readily identify specimens of

Amphiops.

The form of the pupae is unusual, in particular

the virtual absence of setae, the compact ventral

surface and hard shield-like dorsal surface.

In the laboratory A. queenslandicus (and also A.

duplopunctatus, of which a single larva was

reared through to adult but the larval exuvium was

lost) pupated above ground in the aquaria. This,

together with the unusual shape and dark colour

of the pupae, suggest that in the wild they pupate

attached to the stems of emergent vegetation. Just

how they physically attach themselves remains to

be discovered.

Specimens examined

Queensland: 1 km W Mingela, 4/5/98.

Amphiops sp. Northern Territory: Jabiru,

CHS WATTS

21/3/98; 11 km E Jabiru, 19/3/98; Manton Dam,

23/3/98; Ormiston Gorge, 27/1/99. Queensland:

10 km S Cardwell, 7/2/97; 40 km S Cardwell,

7/2/97; 5 km NE Mt Molloy, 30/3/96; 2km S Mt

Molloy 30/3/96; Nardello’s Lagoon near Mareeba,

29/3/96; same locality, 6/2/97; 10km W Petrie,

23/11/95; 12km NW Townsville, 8/2/97; 25 km

S Townsville, 25/3/96.

Berosus Leach

Berosus australiae Mulsant. Fig. 5

Size of third instar. Length 6.5-1.05 mm

(exclusive of gills); head capsule 0.75—-0.95 mm

long, 0.85—1.02 mm wide.

Head capsule. Subquadrate. Labroclypeus

strongly asymmetrical; nasale narrow, strongly

protruding, left of centre, with five to seven short

teeth; lateral lobes of epistome asymmetrical, right

one weakly projecting, left one strongly projected

anteriorly, covering basal third of mandible, with

several short, strong, curved spines. Frontal sulci

lyriform, not coming together; absent on third

instar larvae. Coronal sulcus absent, Gular

sclerites absent, a pair of small, weakly

sclerotised, cervical sclerites. With inverted U-

shaped darker markings on dorsal surface.

Antennae. Three-segmented, relatively close

together, arising just inwards of mandible bases.

First segment a little longer than other two

combined, with a stout subapical seta on inner

margin. Sensory appendage on second segment,

about half length of short third segment.

Mandibles. Slender, strongly asymmetrical;

right mandible with large distal tooth and a very

small central one; left mandible with group of

seven small teeth or strong spines, some of which

are bifid at tips.

Maxillae. Five-segmented. Stipes much longer

than remaining segments combined, with four or

five setae on inner margin. Palpus four-

segmented; first segment short, subquadrate, with

an inner process; second segment very short; third

segment the longest; fourth a little longer than

first.

Labium. Mentum short, poorly sclerotised.

Prementum small, subquadrate. Palpus long, two-

segmented, basal segment shorter than distal.

Ligula absent or reduced to small bulge.

Thorax and legs. Prothorax with large dorsal

sclerite, with sagittal line; ventral surface with a

large subrectangular sclerite, hind edge strongly

sinuate, with sagittal line. Mesothorax with two

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 115

FIGURE 5. Berosus australiae. a, habitus; b, labroclypeus; ¢, labium; d, maxilla; e, antenna; f, mandibles.

dorsal pairs of sclerites, inner pair small, outer

pair much larger, subtriangular. Metathorax

lacking sclerites. Mesothorax and metathorax with

a pair of small lateral tubercles bearing a long

apical seta. Legs five-segmented, long, visible in

dorsal view.

Abdomen. Abdominal segments 1 to 7

subdivided by a transverse fold; each with a pair

of long, lateral, tracheal gills, and two pairs of

small cylindrical structures on dorsal surface in a

line inward from spiracles. Segment 8 small, with

a dorsal sclerite; spiracular atrium absent.

Urogomphi reduced. Integument densely covered

with small setae. Bases of gills with covering of

small setae similar to those on adjacent abdominal

segments. Each gill with a relatively long seta,

arising from a small cylindrical base, towards tip.

Dorsal surface often with darker patterning.

Spiracles. Nine pairs of non-functional

spiracles, one mesothoracic and eight abdominal.

Intraspecific variation

There is some variation in the specimens I have

identified as this species in the number and shape

of the nasale teeth and in the presence/absence/

strength of a dorsal colour pattern.

116

Interspecific variation

The description of B. australiae fits most of

the specimens of Berosus that I have seen,

many of which are unlikely to be B. australiae.

I suspect that the description would fit the

larvae of most species in the B. australiae, B.

majusculus Blackburn, B. veronicae Watts

group of Australian Berosus. There is some

variation in colour and in the strength of setae

on the integument (weak in B. australiae).

Another group of specimens show clear

differences from B australiae: the nasale is

central rather than to the left; the integument

is covered with small dark-pigmented patches

in various patterns; the abdominal segments

have variably sized, but often marked, colour

pattern; there are finger-like projections on the

sides of the prothorax and a differently shaped

eighth abdominal segment. These larvae are

associated with species such as B.

queenslandicus Blackburn and B. involutus

(W. MacLeay).

Identification

By association and elimination.

Remarks

The long filamentous lateral gills and the

absence of functional spiracles readily identify

larvae of Berosus. The asymmetry of the

mandibles and labroclypeus, stout maxillae

and lack of a ligula are also distinctive, but are

characters shared variously by a number of

other genera. Within Australian species there

is no sign of the variation in the number of

pairs of lateral gills seen in African (Bertrand

1972) and New World (Archangelsky 1997)

species.

Specimens examined

Queensland: Bohle

Townsville, 23/3/96.

Berosus sp. New South Wales: 8 km N

River, 10km N

Bombala, 28/11/98; ditto, 3/11/97;

Collector, 20/1/97; 5 km NE Dartmoor,

11/10/96. Northern Territory: Mt

Borradaile Station, 26/5/99; Nawurlandja,

Kakadu National Park, 29/1/99.

Queensland: Burdekin River, 2/11/95; 10 K

S Cardwell, 7/2/97; Jourama Falls, 31/10/95;

5 km NE Mt Molloy, 30/3/96. South

Australia: 10 km N Coonawarra; 19/10/99.

Victoria: Simpson Creek, 12 km SW Orbost,

16/1/97. Western Australia: Gin Gin,

15/10/96; 4 km S New Norcia, 15/10/96.

CHS WATTS

Enochrus Thomson

Enochrus eyrensis (Blackburn). Fig. 6

Size of third instar. Length 9.1 mm; head

capsule 0.70—0.75 mm long; 0.75—0.80 mm wide.

Head capsule. Subquadrate. Labroclypeus

asymmetrical; nasale obliquely truncate, left side

shorter than right, with seven to nine short teeth,

first two on right side and last one on left side

largest; left lobe of epistome rounded, shorter than

nasale; right lobe triangular, sharply pointed,

about same length as nasale. Frontal sulci as an

inverted bell, fusing just before reaching occipital

foramen. Coronal sulcus very short. Gular sclerite

absent. Cervical sclerites moderate,

subrectangular.

Antennae. Three-segmented. In third instar first

segment wider than second segment and about

same length; second segment with small setae-

bearing appendage on inside near middle; third

segment about a third the length of the second.

Sensory appendage on second segment much

shorter than third segment.

Mandibles. Strongly asymmetrical; right

mandible with two strong inner teeth on basal

half; left mandible with one strong inner tooth.

Inner margin of right distal tooth and distal parts

of both mandibles serrated.

Maxillae. Five-segmented. Stipes wide, much

longer than remaining segments combined, with

four strong setae on inner margin. Palpus four-

segmented; first segment subrectangular, a little

wider that long, with an inner process a little

shorter than second palpal segment; second

segment short; third and fourth segments about

twice as long as second, subequal in length.

Labium. Mentum subtrapezoidal, dorsal surface

with two short stout spines in middle towards

front, three strong spines laterally, anteriolateral

angles each with a strong spine. Prementum squat,

a little wider at base, shorter than mentum. Palpus

two-segmented, first segment short, second

segment approximately twice as long as first.

Ligula slender, longer than first palpal segment,

with partial ring of small spines around bases of

palpal segments.

Thorax and legs. Prothorax completely covered

by a dorsal sclerite, with sagittal line; two small

very narrow sclerites on rear edge; front half of

ventral surface with large sclerite with sagittal

line. Mesothorax with large rectangular dorsal

sclerite with sagittal line, with numerous strong

setae; anterior half of metathorax with dorsal

sclerite with sagittal line, posterior half with two

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 117

c d e

FIGURE 6. Enochrus eyrensis. a, labroclypeus; b, labium; c, maxilla; d, antenna e, mandibles.

small foot-shaped sclerites. Legs five-segmented,

well developed, visible in dorsal view.

Abdomen. Segments 1 to 7 similar in shape and

size, subdivided into three or four transverse

folds; segment 1 with two dorsal pairs of small

suboval sclerites, the anterior pair much the

smaller. Segment 8 with a large, suboval, dorsal

sclerite. Segment 9 trilobed, with a pair of short

one-segmented urogomphi. Segments 2 to 7 with

ventral pair of small protuberances covered with

relatively strong, hooked setae (prolegs).

Integument rather sparsely covered with small

setae.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles non-functional. Spiracles on

segment 8 annular, large and functional, within

the spiracular atrium.

Intraspecific variation

No significant variation within the few

specimens seen.

Interspecific variation

Within the few specimens available, which

must include at least two additional species, there

is some variation in the distribution of setae on

the thoracic sclerites, in the number and relative

size of the nasale teeth, and in the length of the

ligula.

Specimens of E. mastersi differ more

substantially in: pattern of setae on integument,

118

the presence of abdominal sclerites, lack of

prolegs, the number of mandibular teeth, and the

smaller and more distal lateral protuberance on

the second antennal segment. The species is

separately described below.

Identification

By rearing larvae collected in the field.

Remarks

The larvae of the Australian E. (Methydrus)

maculiceps (MacLeay) (Anderson 1976) and E.

(M.) eyrensis differ little from those of the North

American E. (Lumetus) ochraceus (Melsheimer)

(Archangelsky 1997) and E. (L.) fimbriatus

(Richmond 1920, as E. perplexus (le Conte))

except possibly in a squatter mentum. Enochrus

(Hydatotrephis) mastersi differs from all of the

above by the nearly symmetrical mandibles and the

lack of obvious setiferous prolegs, and from E. (M.)

eyrensis, by the more elongate and spinose

mentum. In these characters, other than the absence

of prolegs, it more closely resembles larvae of the

subgenus Lumetus than those of Methydrus.

All known larvae of Enochrus share with most

Helochares the angled nasale. Apart from E.

mastersi, they are readily separated from

Helochares by the asymmetrical mandibles.

Enochrus mastersi can be separated from

Helochares by having the ligula shorter rather

than longer than the basal segment of the labial

palpus, in the lack of a basal patch of strong setae

on the maxillary stipe, and in the presence of

small dorsal sclerites on the abdominal segments.

Specimens examined

South Australia: 2 km S Penola, 10/99.

Enochrus sp. Queensland: Bohle River, 10 km

N Townsville, 23/3/96; 30 km SE Townsville

4/5/98. South Australia: Mandina Lakes, 15/10/00;

Tea Tree Gully, 7/5/97.

Enochrus mastersi (W. MacLeay). Fig. 7

Size of third instar. Length 12.0 mm; head

capsule 1.30 mm long, 1.13 mm wide.

Head capsule. Subquadrate. Labroclypeus

weakly asymmetrical; nasale obliquely truncate,

left side shorter than right, with seven to eight

short teeth, first two on right side and last one on

left side largest; left lobe of epistome more

rounded than right, shorter than nasale. Frontal

sulci as an inverted bell, fusing before reaching

occipital foramen. Coronal sulcus very short.

CHS WATTS

Gular sclerite absent. Cervical sclerites small,

subrectangular.

Antennae. Three-segmented. First segment

wider than second segment, shorter than second in

first instar larvae, about 1.5 times as long as

second in third instar larvae, slightly bulging on

inner apical corner; third segment about a third

the length of the second. Sensory appendage on

second segment much shorter than third segment.

Mandibles. Weakly asymmetrical; right

mandible with two strong inner teeth on basal

half; left mandible with one strong inner tooth

and one much weaker tooth. Inner margins of

distal teeth and distal parts of mandibles serrated.

Maxillae. Five-segmented. Stipes wide, much

longer than remaining segments combined, with

four strong setae on inner margin. Palpus four-

segmented; first segment subrectangular, as wide

as long, with an inner process a little shorter than

second palpal segment; second segment short;

third and four segments slightly longer, subequal

in length.

Labium. Mentum rectangular, dorsal surface

with scattered cuticular spines and several strong

spines at front and side edges. Prementum

rounded, a little wider at base, shorter than

mentum, Palpus two-segmented, first segment

short, second segment 1.0-1.5 times longer than

first, some small spines at the bases of both

segments. Ligula slender, as long as first palpal

segment.

Thorax and legs. Prothorax completely covered

by a dorsal sclerite, with sagittal line, with

numerous relatively long setae; ventral surface

with two subrectangular sclerites. Mesothorax

with a pair of large dorsal sclerites with irregular

posterior margins, covered with relatively long

setae. Metathorax with narrow pair of dorsal

sclerites each with a small foot-shaped backward

extension, covered with relatively long setae. Legs

five-segmented, well developed, visible in dorsal

view.

Abdomen. Segments | to 7 similar in shape and

size, subdivided into three or four transverse

folds; segments 1 to 6 with a dorsal pair of small

suboval sclerites. Segment 8 with a large, suboval,

dorsal sclerite. Segment 9 trilobed, with a pair of

short one-segmented urogomphi. Integument

covered with dense, small setae, in many places

organised into tight bundles.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles non-functional. Spiracles on

segment 8 annular, large and functional, within

the spiracular atrium.

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE

119

(

FIGURE 7. Enochrus mastersi. a, labroclypeus; b, labium; c, maxilla; d, antenna; e, mandibles.

Intraspecific variation

Only three specimens are known. They vary

slightly in size of the abdominal sclerites but

otherwise are very similar.

Interspecific variation

See under E. eyrensis.

Identification

By association and elimination.

Specimens examined

New South Wales: Salisbury, 26/11/95.

Northern Territory: Ormiston Gorge, 27/1/99.

Victoria: King Parrot Creek, 2/12/98.

Helochares Mulsant

Helochares tristis W. MacLeay. Fig. 8

Size of third instar. Length 6.8-10.0 mm; head

capsule, 0.53-0.56 mm long, 0.66—0.69 mm wide.

Head capsule. Subquadrate. Labroclypeus

asymmetrical; nasale obliquely truncate, shorter

on left side, with six small teeth; lateral lobes of

epistome project nearly as far as nasale. Frontal

sulci inversely bell-shaped, meeting before

reaching occipital foramen. Coronal sulcus very

short. Gular sclerite absent. Cervical sclerites

narrow, subrectangular.

Antennae. Three-segmented. First segment as

120

long as second (first instar larvae) or longer

(second and third instar larvae); second segment

with a slim apical sensory appendage on inside

which is half the length of third segment; third

segment much smaller than second.

Mandibles. Slightly asymmetrical; two inner

teeth in middle, distal one larger on left mandible,

only slightly larger on right mandible. Distal inner

margin and inner margins of teeth slightly

serrated. Distal part of teeth darker.

Maxillae. Five-segmented. Stipes wide, longer

than remaining segments combined, with a row of

four stout setae on inner margin, base with a tuft

of strong spines. Palpus four-segmented; first

segment subrectangular, with an inner process a

little longer than the second palpal segment;

second segment the shortest; third segment

longest; last segment slightly shorter than third.

Labium. Mentum large, subquadrate, with

dorsal surface covered by strong cuticular spines.

Prementum subrectangular, wider than long.

Palpus two-segmented, basal segment very short,

distal segment three to four times longer than

basal; a few small spines at base of apical

segment. Ligula about twice as long as first palpal

segment.

Thorax and legs. Prothorax almost completely

covered by a dorsal sclerite, with sagittal line;

ventral surface with a subrectangular plate, with a

sagittal line. Mesothorax with a pair of large

rectangular dorsal sclerites; metathorax with a pair

of small dorsal sclerites, irregular in shape,

composed of a wide and narrow basal portion and

an L-shaped central portion arising from the

middle of the basal piece. Legs five-segmented,

well developed, visible in dorsal view.

Abdomen. Segment 1 with two dorsal pairs of

small, narrow, irregularly shaped sclerites; anterior

pair much smaller. Segments 1 to 7 similar in size

and shape, subdivided by three or four transverse

folds; pleura of segments 1 to 7 with three

longitudinal slight bulges, the most ventral one in

three lobes, each segment with a dorsal pair of

setae towards the middle arising from a small

circular sclerite or dark-pigmented area, and a

small seta a little inwards and backwards from

each spiracle. Segment 8 with a large, suboval,

dorsal sclerite, serrate on posterior edge and pair

of short apical flaps. Segment 9 trilobed; with a

pair of small one-segmented urogomphi; central

lobe largest. Integument densely covered in short

fine setae.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal, raised slightly above the surface.

Mesothoracic and first seven abdominal spiracles

CHS WATTS

non-functional. Spiracles on segment 8 annular,

large and functional, within the spiracular atrium.

Intraspecific variation

There is some variation in the shape of the

thoracic sclerites and in the shape and

configuration of the nasale teeth.

Interspecific variation

H. luridus (W. MacLeay) and H. clypeatus

Watts. As for H. tristis.

H. tenuistriatus Regimbart. The integument is

predominantly covered by long thin setae rather

than the short and very curved setae found in H.

tristis, H. luridus and H. clypeatus. This gives a

distinct furry look to the larvae.

H. foveicollis (Montrouzier). Differs in lacking

nasale, having a slight bulge on the inside of

antenna and a number of other characters (see

below). These differences are sufficient to warrant

a separate description (see below).

Other Helochares species. Within specimens

unidentified to species there is variation in: form

of the small setae on the integument; the shape of

the metasternal sclerites; and the presence/

absence, position and shape of the abdominal

sclerites.

Identification

H. tristis, H. luridus and H. clypeatus by rearing

larvae collected in the field; H. tenuistriatus by

association and elimination.

Remarks

Within the known Australian Helochares larvae

there are clearly two groups: H. foveicollis; and

H. tristis, H. luridus, H. tenuistriatus and H.

clypeatus. These correspond to the subgenera

Helochares and Hydrobaticus Blackburn,

respectively, and strongly reinforce the

distinctiveness of these groups as reflected in the

classification. Within Australian 4d.

(Hydrobaticus) there is very little difference

between the known species or among the large

number of unidentified larvae. In addition, there

is minimal difference between these Australian H.

(Hydrobaticus) and the North American H.

(Hydrobaticus) maculicollis Mulsant

(Archangelsky 1997; Richmond 1920) or the

South American H. (Sindolus) talarum Fernandez)

and H. (Helochares) pallipes (Brulle) (Fernandez

1983) or the European H. (Helochares) obscurus

(Muller) (as griseus Heer) (Boving & Henriksen

1938).

The larvae of H. (Helochares) foveicollis differ

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 121

FIGURE 8. Helochares tristis. a, labroclypeus; b, labium; ¢, maxilla; d, antenna e, mandibles.

considerably from all of these species in a number

of characters, most noticeably the symmetrical

labroclypeus and the more elongate antennae and

maxillary palpi (see later).

The larva of the related Australian genus,

Chasmogenus, is only known from the work of

Anderson (1976). It also has a symmetric

labroclypeus but differs from both JH.

(Helochares) foveicollis and H. (Hydrobaticus)

tristis in the asymmetric mandibles, one with one

tooth and the other with two, and in the complete

rather than partial dorsal sclerites on the

metathorax.

Species of H. (Hydrobaticus) appear to breed

more or less continuously and females are

frequently caught with egg masses attached to

their abdomens, although Anderson (1976) found

that there was a distinct spring breeding season in

H. tristis near Sydney. Larvae are among the most

commonly collected Hydrophilid larvae, often

found together with those of Limnoxenus in the

south and both Limnoxenus and Sternolophus in

the north. They can be readily separated from both

of these by their angled nasale and mandibles with

two rather than three teeth.

Although adults of Enochrus are often more

common than those of Helochares, their larvae

are, for unknown reasons, rarely found. They

share with HA. (Hydatotrephis) the angled nasale

but, other than E. mastersi, have strongly

asymmetrical mandibles. Larvae of E. mastersi

resemble those of H. (Hydatotrephis) quite closely

122 CHS WATTS

but differ in lacking a cluster of strong setae at the

base of the maxillary palpus, and in having the

ligula shorter than the basal segment of the labial

palpus.

Specimens examined

New South Wales: 20 km W Nelligan,

3/11/97. South Australia: 10 km N Coonawarra,

26/9/98; Cudlee Creek, 10/11/96; Kuitpo, 5/10/95;

13 km W Meadows, 26/9/96; Mt Crawford State

Forest, 10/11/96; Tea Tree Gully, 7/5/97.

Victoria: 10km W Cowwarr, 30/11/98;

Healsville, 12/68.

H. tenuistriatus. Western Australia: 30 km N

Perth, 14/10/96

H. clypeatus. Northern Territory: Nourlangie

Creek, 20 km SSW Jabiru, 11/10/98.

H. luridus. Queensland: Burdekin River E of

Charters Towers, 4/5/98.

Helocharus sp. New South Wales: 12km E

Tamworth, 26/11/95. Northern Territory:

Nawurlandja, Kakadu National Park, 22/3/98.

Queensland: Alligator River 20 km S Townsville,

25/3/96; Bluewater, 22/3/96; Bowling Green Bay

National Park, 6/11/95; 10 km N Cairns, 4/1/97;

Eubenangee Swamp, 4/2/97; 8km S Greenvale,

27/3/96; Jourama Falls near Townsville, 31/10/95;

1km W Mingela, 4/5/98; Star River, 1/11/95;

30 km SE Townsville, 4/5/98.

Helochares foveicollis (Montrouzier). Fig. 9

Size of third instar. Length 13.5 mm; head

capsule 0.78 mm long, 0.94 mm wide.

Head capsule. Subquadrate. Labroclypeus

symmetrical, without nasale; with eight to nine

small teeth along front edge in middle; lateral

lobes of epistome slight. Frontal sulci inversely

bell-shaped, meeting before reaching occipital

foramen. Coronal sulcus short. Gular sclerite

absent. Cervical sclerites small, subrectangular.

Antennae. Three-segmented. First segment as

long as second (first instar larvae) or longer

(second and third instar larvae), with distinct

protruberance on inside near apex; second

segment with a slim apical sensory appendage on

inside, half the length of third segment; third

segment much thinner than second.

Mandibles. Relatively slim, symmetrical; two

inner teeth in middle, distal one much larger.

Outer margin of apical tooth weakly serrated.

Maxillae. Five-segmented. Stipes relatively

narrow, much longer than remaining segments

combined, with a row of five stout setae on inner

margin, base with a few short, strong spines.

Palpus four-segmented; first segment

subrectangular with an inner process as long as

the second palpal segment; second segment the

shortest; third segment the longest; fourth segment

shorter than third.

Labium. Mentum large, subquadrate, with

central portion of dorsal surface covered by weak

cuticular spines and a semicircle of six spines,

anterolateral angles sharp, each with a small spine.

Prementum elongate-—rectangular, longer than

wide. Palpus two-segmented, basal very short,

distal segment four to five times longer than basal

one. Ligula short, thick, about twice the length of

first palpal segment.

Thorax and legs Prothorax almost completely

covered by a dorsal sclerite, with sagittal line;

ventral surface with a subrectangular sclerite,

subdivided by a sagittal line. Mesothorax with a

pair of large dorsal sclerites; metathorax with a

pair of small dorsal sclerites composed of a wide

and narrow basal portion and an L-shaped portion

arising from the middle of the basal piece, ‘heel’

reaching posterior margin. Legs five-segmented,

well developed, visible in dorsal view.

Abdomen. Segments 1 to 7 similar in size and

shape, subdivided by three or four transverse

folds; segment 1 with two dorsal pairs of small

narrow basal sclerites, the anterior pair smallest;

pleura of segments 1 to 7 each with three to four

weak longitudinal bulges, a well separated pair of

strong dorsal setae arising from a very small

circular darkly pigmented area, a moderately

strong seta just inwards and behind each spiracle.

Segment 8 with a large, suboval, dorsal sclerite

with a ragged hind edge, and a pair of prominent

apical flaps; segment 9 trilobed, central lobe

largest and sclerotised, with a pair of prominent

one-segmented urogomphi. Integument covered

with moderately dense hair-like setae.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles very small, non-functional.

Spiracles on segment 8 annular, large and

functional, within the spiracular atrium.

Intraspecific variation

Among the few specimens known there is some

variation in the number of larger spines on the

mentum and in wear on the nasale teeth.

Interspecific variation

Helochares foveicollis differs from other known

Australian Helochares larvae by the lack of a

nasale, presence of a prominent bulge on the

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 123

a d e

FIGURE 9. Helochares foveicollis. a, habitus; b, labroclypeus; ¢, labium; d, maxilla; e, antenna; f, mandibles.

inside of the antennae, proportionally longer and

narrower prementum and ligula, more elongate

maxillary palpi and the shape of the metathoracic

sclerites (Fig. 9).

Identification

By rearing field-caught larvae and rearing from

egg mass attached to female.

Remarks

On adult characters H. foveicollis is presently

placed in the subgenus Helochares together with

H. obscurus and H. pallipes. However larval

characters do not support this placement, nor its

placement in either of the subgenera

Hydatotrephis or Sindolus.

124 CHS WATTS

Specimens examined

Northern Territory: Manton Dam, 2/2/99;

2km S Adelaide River, 1/2/99. Queensland:

40 km S Townsville, 2/2/97.

Hybogralius Orchymont

Hybogralius hartmeyeri (Regimbart). Figs 10,

17b

Size of third instar. Length 10.0 mm; head

capsule 1.60—1.79 mm long, 1.62—2.00 mm wide

Head capsule. Rectangular. Labroclypeus

asymmetrical; nasale narrow, projecting well

forward, with five teeth, the two lateral ones more

distant than rest; lateral lobes of epistome

rounded, projecting farther than nasale, left lobe

often larger, with a strong comb of stout spines on

front edge, front edge of right lobe with a few

scattered spines. Frontal sulci U to V-shaped,

fusing just before reaching occipital foramen.

Coronal sulcus short. Gular sclerite absent.

Cervical sclerites relatively small, subrectangular.

Antennae. Three-segmented. First segment

longer than remaining segments combined; second

segment about 1.5 times the length of third

segment which is very thin. Sensory appendage

on second segment minute.

Mandibles. Asymmetrical; right mandible with

three inner teeth, the anterior one largest; left

mandible with three teeth of approximately the

same size, central one with patch of strong setae

at base.

Maxillae. Five-segmented. Stipes stout, longer

than remaining segments combined, with four

long stout setae on inner margin. Palpus four-

segmented; first segment widest, with a short

inner process; second segment short; third

segment longest; fourth segment about half length

of third.

Labium. Mentum small, subrectangular, sides

slightly convex, dorsal surface with small spines,

anterolateral angles rounded with several stout

spines. Prementum elongate, rectangular, as long

as mentum. Palpus two-segmented, first segment

the shortest. Ligula present, about as long as the

first palpal segment, tip weakly bifid.

Thorax and legs. Prothorax with large dorsal

sclerite, with sagittal line; ventral surface with a

large subrectangular sclerite, with sagittal line.

Mesothorax with a pair of large triangular dorsal

sclerites. Metathorax with a pair of semicircular

dorsal sclerites and a more posterior pair of much

smaller circular ones. Sclerites, particularly the

anterior ones, covered with very short spines,

occasionally arranged in short rows; rest of thorax

densely covered with short fine setae with a few

much longer setae laterally. Legs five-segmented,

relatively short, just visible in dorsal view.

Abdomen. Segments 1 to 8 similar in shape,

tapering towards the caudal end; Segments 1 to 7

subdivided into three or four transverse folds;

sides of each segment with some slight, fleshy

bulges, with one to two long setae. Segment 8

with two transverse folds, without dorsal sclerite.

Segment 9 with hind margin weakly concave.

Below segment 9, and projecting just beyond, are

two prominent broad lobes. Integument densely

covered with small fine setae.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles on small papillae. Spiracles

on segment 8 (if present at all) obsolete and

certainly non-functional.

Pupa (Fig. 17b). Head without styli. Pronotum

with 22 styli, 16 around margins and 6 on disc.

Mesonotum and metanotum each with one pair of

styli near the midline. Abdominal segments 1 to 7

with a row of four styli on the terga, segments 2

to 7 with one stylus on each pleuron; segment 8

without styli; segment 9 with a pair of long

urogomphi.

Intraspecific variation

One specimen has a slight mandibular tooth

anterior to the normal ones (Fig. 1); otherwise,

there is little variation other than in tooth wear

between the few known specimens.

Identification

By rearing from larvae collected in the field.

Specimens examined

Western Australia: Gooseberry Hill, 14/9/00.

Remarks

Hybogralius is a monospecific Australian genus

known only from a couple of localities in the

escarpment region of the Darling Ranges east of

Perth, Western Australia where it lives in small

temporary winter/spring streams. Although placed

with Limnoxenus in the subtribe Hydrobiina of

the tribe Hydrophilini (Hansen 1991), the larvae

of the two genera have little in common. Several

features of the larvae of Hybogralius stand out:

the lack of a spiracular atrium, the strongly

asymmetric labroclypeus, the narrow prominent

nasale, the strongly asymmetrical mandibles, and

the rings of small spines at the base of the

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE

d e

125

Carer tN aM

FIGURE 10. Hybogralius hartmeyeri a, habitus; b, labroclypeus; c, labium; d, maxilla; e, antenna; f, mandibles.

segments of the labial palpi. In most of these

characters it most closely resembles Berosus.

Apart from the lack of a spiracular atrium and the

presence of a coronal sulcus, it also resembles the

larvae of Oocyclus Sharp and Laccobius in the

tribe Laccobiini. To what degree these similarities

reflect phylogenetic relationships remains to be

seen.

As well as the lack of a functional apical

spiracle, the main tracheal trunks are narrower

than most other genera and the spiracles are not

much larger, if at all, than those of other genera

where they are considered non-functional. How

the larvae manage to breathe is a puzzle.

Hydrobiomorpha Blackburn

Hydrobiomorpha sp. Fig. 11

Size of third instar. Length 15.0 mm; head

capsule 1.65 mm long, 1.70 mm wide.

126

CHS WATTS

FIGURE 11. Hydrobiomorpha sp. a, labroclypeus; b, labium; c, maxilla; d, antenna; e, mandibles.

Head capsule. Subquadrate. Labroclypeus

slightly asymmetrical; nasale truncate, edge rough

rather than toothed; lateral lobes of epistome

rounded, projecting further than nasale. Frontal

sulci V-shaped, meeting before reaching occipital

foramen. Coronal sulcus short. Gular sclerite

absent. Cervical sclerites small, subquadrate.

Numerous small darker patches on posterolateral

angles and within frontal sulci.

Antennae. Three-segmented. First segment

slender, three times longer than remaining

segments combined (third instar larvae), inner

margins with numerous short spines; second

segment as long as third. Sensory appendage of

second segment very small.

Mandibles. Symmetrical, sharply pointed; right

mandible with three teeth on basal half, distal one

large, bifid apically, central tooth smaller, basal

one much smaller; left mandible similar to right

except for basal tooth placed slightly more

ventrally.

Maxillae. Five-segmented. Stipes slender,

longer than remaining segments combined, with a

group of short spines on inner margin at base, and

several setae along inner and outer margins.

Palpus four-segmented; first segment longest, with

a short inner process; second segment the shortest;

third and fourth segments subequal in length, each

twice as long as second segment.

Labium. Mentum approximately square,

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE

anterolateral angles projected, each with a pair of

very small spines; dorsal surface with eight short,

stout setae on distal two-thirds, and short cuticular

spines on basal third. Prementum longer than

wide, half the width of mentum. Palpus two-

segmented, distal segment three to four times

longer than basal segment. Ligula a little longer

than first palpal segment, weakly bifid.

Thorax and legs. Prothorax covered by a large

dorsal sclerite, posterior end rounded, sagittal line

present; ventral sclerite large, subrectangular, with

sagittal line. Mesothorax with pair of irregularly

shaped sclerites, wider anteriorly, with sagittal

line. Metathorax with pair of small irregular

shaped sclerites. Legs five-segmented, long,

visible in dorsal view. Prothorax and mesothorax

covered with small, scattered, dark patches.

Abdomen. First segment subdivided into two

transverse folds, with a dorsal pair of narrow

sclerites near anterior edge; segments 2 to 7

subdivided into four or five transverse folds,

without sclerites. Segments 1 to 7 each with

eight setose tubercles, four dorsal and two on

each lateral margin; pleura weakly lobed.

Segment 8 with a small, suboval, dorsal sclerite,

posterior apex subdivided into four lobes and a

pair of small procerci. Segment 9 round, with

three small dorsal sclerites, bearing a pair of

short, one-segmented urogomphi, and a pair of

long paracerci; a pair of gill-like appendages

(prostyli) originate on ventral side. Integument

sparsely covered with relatively long setae.

Single, narrow, darker line along midline of

dorsal surface.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles non-functional. Spiracles on

segment 8 annular, large and functional, within

the spiracular atrium.

Identification

Agreement with generic description by

Archangelsky (1997). Specimens were taken in

association with H. bovilli Blackburn and are

almost certainly that species.

Interspecific variation

There is little variation between the two known

specimens.

Remarks

The larvae described above differ little from

those of the New World H. casta (Archangelsky

1997; Spangler 1973) or the unidentified African

species described by Berge Henegouwen (1982).

127

The peculiar shape of the anterior mandibular

tooth is diagnostic within the Australian fauna.

Specimens examined

Northern Territory: Holmes Jungle, 28/11/99.

Hydrophilus Geoffroy

Hydrophilus bilineatus (MacLeay). Fig. 12

Size of third instar. Length 35 mm; head

capsule 3.25 mm long, 4.25 mm wide.

Head capsule. Suboval. Labroclypeus virtually

symmetrical; nasale undeveloped; lateral lobes of

epistome rounded, projecting farther than nasale.

Frontal sulci broadly U-shaped, fusing just before

reaching occipital foramen. Coronal sulcus

present, very short. Gular sclerite absent. Cervical

sclerites small, suboval.

Antennae. Four-segmented. First segment

slender, slightly constricted near base, longer than

remaining segments combined, with some slender

setae on distal two-thirds of inner margin; second

segment short, constricted near base in first instar

larvae, third and fourth segments subequal in

length. Sensory appendage on third segment

reduced to a small papilla.

Mandibles. Asymmetrical; right mandible

longer, more slender, with a large tooth on basal

half which is divided by a longitudinal groove

into ventral and dorsal sections; left mandible

shorter, with one small inner tooth on basal half.

Maxillae. Five-segmented. Stipes narrow,

elongate, longer than remaining segments

combined, with three stout setae on inner margin.

Palpus four-segmented; first segment the longest,

with a short inner process; second and third

segments subequal in length; fourth segment

slightly shorter.

Labium. Mentum subrectangular, sides convex,

anterolateral angles lobed, central third with some

small spines (without spines in first and second

instars). Prementum subtrapezoidal, anterior end

wider with a pair of small protuberances towards

front. Palpus two-segmented, first segment

shortest. Ligula present, much shorter than first

palpal segment.

Thorax and legs. Prothorax with two large

sclerites together covering most of dorsal surface,

the triangular area anterior to these sclerites

usually sclerotised; ventral surface with a large

subrectangular sclerite, with sagittal line.

Mesothorax with a pair of subtriangular dorsal

sclerites. Metathorax with a pair of narrow, basal

128 CHS WATTS

c d

FIGURE 12. Hydrophilus bilineatus. a, labroclypeus; b, labium; c, maxilla; d, antenna; e, mandibles.

sclerites and a more central pair of small

irregularly-shaped ones. Legs five-segmented,

relatively short, visible in dorsal view, with rows

of long swimming setae on both dorsal and ventral

edges of femur and tibiotarsus.

Abdomen. Segments 1 to 6 similar in shape,

tapering towards the caudal end; segments 7 to 8

more elongate and slender. Segments 1 to 7

subdivided into three transverse folds; third fold

with two lateral and two dorsal small tubercles,

more obvious on posterior segments. Pleura | to 8

each with a small lateral tubercle. Segment 8 with

two small subtriangular sclerites at posterior end;

segment 9 slightly trilobed, with a pair of short

one-segmented urogomphi, ventromedially with a

pair of long gill-like appendages. Integument in

first instar very densely covered with small thin

setae; in second and third instars moderately

covered with thin, relatively long setae and

numerous small ‘bumps’.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles very small, non-functional.

Spiracles on segment 8 annular, large and

functional, within the spiracular atrium.

Identification

By association and elimination.

Intraspecific variation

I have not noticed any significant variation

within the few available specimens I have

identified as H. bilineatus.

Interspecific variation

Within the larvae available there is considerable

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE

variation in a number of characters, most notably:

number of antennal segments vary from three to

four; the presence/absence/form of the lateral

projections on the abdominal segments; form of

covering of the integument; dorsal colour pattern.

Remarks

The larvae of two European species, H. piceus

Linnaeus and H. aterrimus Eschscholtz, and the

North American H. triangularis (Say) are well

known (Archangelsky 1997; Boving &

Henriksen 1938; Richmond 1920). In addition,

the larvae of H. senegalensis (Percheron) (Berge

Henegouwen 1982; Africa) and H. acuminatus

Motschulsky (Morioka 1955; Japan) have been

described. Within the known larvae there is

considerable variation in the number of antennal

segments in different instars, in the development

of lateral abdominal flaps and, at least within the

Australian species, in the form of the integument

coverage. Somewhat unusually for Hydrophilidae

larvae, it appears that specific differences are

sufficient to enable the larvae of many species to

be identified.

In the lack of lateral abdominal flaps and its

four segmented antennae, H. bilineatus most

closely resembles the North American H.

triangularis but differs in a number of details,

most obviously in the more robust mandibles with

a well-developed molar region. The other

Australian species appear more distant,

particularly in the well-developed abdominal flaps

and number of antennal segments. With the more

robust mandibles and abdominal flaps these

Australian species appear to be closer to the

subgenus Diblocelus (sensu Hansen 1991) than

subgenus Hydrophilus (Archangelsky 1997). On

adult characters they fit clearly into the subgenus

Hydrophilus.

Egg cases of several species were collected in

the field and each produced between 15 and 30

larvae. Those thought to be of H. bilineatus

refused all food offered but the other species

accepted, often with some excitement, small snails

and some accepted freshly killed mosquito larvae.

None accepted chironomid larvae. Unfortunately,

despite seemingly accepting snails as food, none

was reared to the second instar. Species varied in

the degree of cannibalism. Some broods were

quite strongly cannibalistic but in others, apart

from the occasional unfortunate individual,

siblings lived well together.

Specimens of Aydrophilus bilineatus

(MacLeay) have previously been identified as H.

picicornis Chevrolat ( Hansen 1999).

129

Specimens examined

Northern Territory: Manton Dam, 23/3/97;

2km S Adelaide River 1/2/99,

Hydrophilus sp. Northern Territory: Newry

Station, 2/86 col. M. J. Tyler. Queensland: 25 km

S Townsville, 3/5/98, 2/2/97; 8km S Gieenvale,

27/3/96; Skm NE Mt Molloy, 30/3/96. South

Australia: 10 km N Coonawarra, 10/11/97,

16/10/97; 1 km S Nangwarry, 9/10/97. Tasmania:

12 km N Hobart, 2/12/00; 2km W Fingle,

23/1/00; 3 km SW Clifton Beach, 4/12/00.

Victoria: 5 km NE Dartmoor, 11/10/97. Western

Australia: Murchison River, 13/3/95, col. S.

Halse; 6 km S Pinjarrah, 23/10/96.

Laccobius Erichson

Laccobius decipiens Gentili. Fig. 13

Size of third instar. Length 5.0-6.5 mm; head

capsule 0.43-0.50 mm long, 0.45—0.46 mm wide.

Head capsule. Subrectangular. Labroclypeus

strongly asymmetrical; nasale prominent, with

three teeth; lateral lobes of epistome

asymmetrical, both projecting further than nasale;

left lobe larger, covering basal third of mandible,

with a row of about 12 strong curved setae on

inner margin, larger ones dentate; right lobe

smaller, covering a quarter of mandible, without

setae. Frontal sulci parallel, reaching occipital

foramen without coming together, difficult to see

in third instar larvae. Coronal sulcus absent. Gular

sclerite absent; cervical sclerites small and oval,

difficult to see in first instar larva.

Antenna. Three-segmented; second segment the

longest, with a small outer sensory appendage

about a quarter the length of third segment; first

segment half the length (first instar larvae) to

slightly shorter (third instar larvae) than second

segment.

Mandibles. Strongly asymmetrical. Left

mandible with three inner teeth, central one

largest, with five strong spines, region behind

third tooth with a number of very short spines;

right mandible with two inner teeth, larger than

those of left mandible, front one much larger,

occasionally also with a very small third tooth, or

spine, behind the second.

Maxillae. Five-segmented. Stipes wide, longer

than remaining segments combined, inner margin

with four setae. Palpus four-segmented; first segment

subquadrate, incompletely sclerotised, with a small

inner process; second segment shortest; third and

fourth segments subequal in length.

130 CHS WATTS

FIGURE 13. Laccobius decipiens. a, habitus; b, labroclypeus; c, labium; d, maxilla; e, antenna; f, mandibles.

Labium. Mentum small and_ narrow,

subrectangular. Prementum slightly larger,

trapezoidal. Palpus two-segmented, first segment

shorter, second segment two to three times longer

than first. Ligula obsolete, at most reduced to a

slight bulge.

Thorax and legs. Prothorax completely covered

by a large dorsal sclerite, with sagittal line;

ventrally with a pair of large subrectangular

sclerites, fused in anterior half. Mesothorax with

two pairs of dorsal sclerites, anterior one small,

subrectangular; posterior pair larger,

subtriangular. Metathorax with one pair of small,

irregularly shaped sclerites intermediate in size to

the two mesothoracic ones. Legs five-segmented,

visible in dorsal view.

Abdomen. Segments 1 to 7 similar in shape,

tapering towards posterior end, ‘segments

subdivided by transverse folds; sides with several

slight bulges, each segment with a pair of long

dorsal setae arising from small, darkly pigmented

areas, one just behind spiracle and the other near

midline. Segment 8 with a subtriangular dorsal

sclerite. Segment 9 small, trilobed, bearing a pair

of small one-segmented urogomphi. Integument

with quite dense covering of short fine setae.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles non-functional, on short dark-

pigmented papillae; spiracles on segment 8

annular, large and functional, within the spiracular

atrium.

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 131

Intraspecific variation

Within specimens that I have identified as this

species there is some variation as follows: some

specimens have an additional small third tooth on

the right mandible; in newly moulted specimens

there are slight serrations on the upper mandibular

teeth which appear to wear away quickly; the first

abdominal segment occasionally has a pair of very

small dorsal sclerites.

Interspecific variation

Within specimens unidentified to species there

is variation in the shape of the dorsal sclerites on

abdominal segment 8; the width of the nasale and

the shape of the nasale teeth; the presence, size

and position of abdominal sclerites. Overall these

differences are slight.

Identification

By rearing larvae collected in the field.

Specimens examined

New South Wales: Bombala, 28/11/98; ditto,

4/11/97; ditto, 18/1/97. Victoria: Stratford,

7/11/97.

Laccobius spp. Queensland: Kauri Creek near

Tinarro Dam, 24/10/93, col. D. Larson.

Remarks

There is little difference between the larvae of

the Australian species and those of the North

American L. minutoides Orchymont and L. agilis

(Randall) described and illustrated by Richmond

(1920) and Archangelsky (1997).

Limnoxenus Motschulsky

Limnoxenus zealandicus (Broun). Fig. 14

Size of third instar. Length 11.5—15.0 mm; head

capsule 1.60-1.79 mm long, 1.62—2.00 mm wide.

Head capsule. Square. Labroclypeus slightly

asymmetrical; nasale moderately developed,

angled slightly with right side more forward, with

five teeth, left tooth a little distant from others;

lateral lobes of epistome rounded, projecting not

quite as far as nasale, right lobe often a little more

rounded and a little more projected than left lobe.

Frontal sulci U to V-shaped, fusing just before

reaching occipital foramen. Coronal sulcus very

short. Gular sclerite absent. Cervical sclerites

relatively small, subrectangular.

Antennae. Three-segmented. First segment

longer than remaining segments combined, with

some slender setae on distal two-thirds of inner

margin; second segment about twice the length of

third segment which is very thin. Sensory

appendage on second segment minute.

Mandibles. Symmetrical with three inner teeth,

progressively smaller towards base.

Maxillae. Five-segmented. Stipes stout, longer

than remaining segments combined, with four

long stout setae on inner margin. Palpus four-

segmented; first segment the widest, with a short

inner process; second and fourth segments short,

subequal in length; third segment longest.

Labium. Mentum subrectangular, sides slightly

concave, dorsal surface with small spines,

anterolateral angles rounded, with several stout

spines. Prementum square, not much smaller than

mentum. Palpus two-segmented, first segment the

shortest. Ligula present, slightly longer than first

palpal segment, tip bifid.

Thorax and legs. Prothorax with large dorsal

sclerite, with sagittal line; ventral surface with a

large subrectangular sclerite, with sagittal line.

Mesothorax with a pair of large triangular dorsal

sclerites. Metathorax with a pair of wineglass-

shaped dorsal sclerites sometimes with stem of

wineglass absent. Rest of surface of thorax

covered with very short spines, occasionally

arranged in short rows. Legs five-segmented,

relatively short, barely visible in dorsal view.

Abdomen. First segment with a dorsal pair of

small subovoid sclerites close to anterior margin.

Segments 1 to 6 similar in shape, tapering towards

the caudal end; segments 7 and 8 more elongate

and slender. Segments 1 to 7 subdivided into three

transverse folds; dorsal surface of third fold with

four small, dark, narrowly cylindrical tubercles;

sides of each segment with three slight, fleshy

bulges. Segment 8 with a large suboval dorsal

sclerite, with three to four quite deep indentations

on hind edge, lighter coloured towards middle.

Segment 9 trilobed, with a pair of short one-

segmented urogomphi. Integument densely

covered with small setae and a sparse covering of

longer setae which get progressively smaller

posteriorly. Dorsal surface often with two darker

Stripes through sclerites and inner pair of

tubercles.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles non-functional, on small

papillae. Spiracles on segment 8 annular, large

and functional, within the spiracular atrium.

Intraspecific variation

The number of nasale teeth is variable with up

132 CHS WATTS

a d f

FIGURE 14. Limnoxenus zealandicus. a, habitus; b, labroclypeus; ¢, labium; d, maxilla; e, antenna; f, mandible.

to ten smaller teeth in some specimens; in some _/dentification

specimens these are worn down to the stage where By rearing from larvae collected in the field.

the front of the nasale is almost smooth. The

central nasale teeth are not well developed in first Remarks

instar larvae. In one specimen the antennae are The larva of the European L. niger (Gimelin)

four segmented with the normal third segments has already been described by Berge Henegouwen

clearly divided into two. (1975). The larva of the sole Australian and New

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 133

Zealand species differs little from those of this

species,

One of the most commonly collected

hydrophilid larvae in Australia, it is quickly

recognised by the parallel darker stripes on the

dorsal surface and combination of relatively short

basal segment to the antenna and symmetrical

three-toothed mandibles. The dense covering of

small setae gives the abdomen a fur-like look.

Specimens examined

New South Wales: 10 km E Braidwood, 30/11/95;

Braidwood, 19/1/97.

South Australia: 10 km N Coonawarra, 10/11/97;

10km E Mt Compass, 13/9/97; Mt Crawford

Forest, 10/11/96; 6km N Forreston, 3/10/95;

19 km N Forreston, 3/9/99; 1 km S Nangwarry,

9/10/97; ditto, 29/8/99; 2 km S Penola, 20/11/99;

Warburton River, 2-8/10/99, 27°52'23"S

137°54'40"E; Watervalley, 7.2 km NNE Mt

Rough, 15/10/00. Tasmania: Lake St Clair, 4 km

N Derwent Bridge, 25/1/00. Victoria: 5 km NE

Dartmoor, 11/10/97; 2 km W Brimpaen, 23/9/98;

Healsville, 12/68; 6km N Noojee, 16/1/97.

Western Australia: Gin Gin, 15/10/96; 10km E

Kalamunda, 16/10/96; 2 km W Nannup, 20/10/96;

6km §S Pinjarrah, 23/10/96; 1 km S Serpentine,

24/10/96; 10 km S Yallingup, 22/10/96.

Regimbartia Zaitzev

Regimbartia attenuata (Fabricius). Fig. 15

Size of third instar. Length 5.0-7.5 mm; head

capsule 1.00—1.01 mm wide, 0.75—0.80 mm long.

Head capsule. Subquadrangular. Labroclypeus

symmetrical; nasale very short, with numerous

short teeth on anterior border of epistome; lateral

lobes of epistome rounded, very short, not

projecting as far as nasale, each with two long

setae. Frontal sulci straight, short, meeting at

about the level of the antennal bases. Coronal

sulcus long, about half the length of the head

capsule. Gular sclerite absent. Cervical sclerites

small, subrectangular.

Antennae. Three-segmented. First segment

slender, longer than other two combined, with a

prominent subapical inner process some distance

from apex. Sensory appendage on second segment

slim, slightly shorter than third antennal segment.

Mandibles. Virtually symmetrical, long and

slender, with two inner teeth on basal half; distal

tooth large, basal one small.

Maxillae. Five-segmented. Stipes slender, much

longer than remaining segments combined, with

four or five setae on inner margin. Palpus four-

segmented; third segment longest; second segment

shortest; first and fourth segments subequal in

length, first segment with a short inner process.

Labium. Mentum large, wider than long, sides

convex, dorsal surface with cuticular spines.

Prementum elongate, much narrower than

mentum. Palpus two-segmented, basal segment

short. Ligula annular, two to three times as long

as basal segment of palpus.

Thorax and legs. Prothorax with a large dorsal

sclerite, with sagittal line; surface covered by

minute spines; fringe of long, slender setae around

the margins, four small spines, one on each

anterolateral angle and two towards the middle;

ventral surface with large, subrectangular sclerite,

without sagittal line. Mesothorax with two dorsal

pairs of subtriangular sclerites; metathorax with

one dorsal pair of irregular shaped sclerites. Both

mesothorax and metathorax with five pairs of

setiferous projections, about half of them with a

narrow cylindrical projection with a long seta

attached at the end. Legs five-segmented, long,

visible from above.

Abdomen. Segments 1 to 7 with five pairs of

setiferous projections similar to those on thorax,

four lateral ones with one more posterior than

others and one closer to midline. In addition, the

abdomen is covered with small stellate setae and

the dorsal surface with small projections each

with six to eight long, sharply pointed setae (in

most specimens these structures are obscured by

an accumulation of sand grains and detritus).

Segment 8 without setiferous projections, with a

pair of small finger-like ‘procerci’, with dorsal

semispherical sclerite which covers the spiracular

atrium. Segment 9 trilobed, with a pair of small,

unsegmented urogomphi.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles non-functional. Spiracles on

segment 8 annular, large and functional, within

the spiracular atrium.

Interspecific variation

There is little variation between the few

specimens seen.

Identification

By association of adults and larvae by

isoenzyme electrophoresis.

Remarks

The larvae closely resemble both Allocotocerus

134

me a

EROS

CHS WATTS

FIGURE 15. Regimbartia attenuata. a, habitus; b, labroclypeus; ¢, labium; d, detail of abdominal stella; e, tip of

antenna; f, maxilla; g, antenna; h, mandibles.

and the New World Derallus (Archangelsky 1997;

Spangler 1966), all three genera showing a suite

of unusual characters that clearly separate them

from other Hydrophilid larvae. The differences

between the genera are relatively slight (see key

and under Allocotocerus).

Specimens examined

Queensland: 6 km N Bluewater, 3/2/97; 15 km

W Mareeba, 6/12/90, col. D. Larson; 5 km NW

Mareeba, 22/9/90, col. D. Larson; 2km N Mt

Molloy, 1/4/96.

Sternolophus Solier

Sternolophus marginicollis (Hope). Fig. 16

Size of third instar. Length 11.5—15.0 mm; head

capsule 1.80—1.90 mm long, 1.55—1.70 mm wide.

Head capsule. Subrectangular. Labroclypeus

weakly asymmetrical; nasale short, weakly

projecting with five short teeth, left one a little

distant from rest; lateral lobes of epistome

symmetrical not projecting beyond nasale,

undulating. Frontal sulci V-shaped, fusing before

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 135

a d

e f

FIGURE 16. Sternolophus sp. a, habitus; b, labroclypeus; ¢, labium; d, maxilla; e, antenna; f, mandible.

reaching occipital foramen. Coronal sulcus short.

Gular sclerite absent. Cervical sclerites small,

suboval, longitudinally oriented. Frontal sulcus

outlined in a darker colour.

Antennae. Three-segmented. First segment

much longer than other two combined, with an

annular ring of long setae near apex in second and

third instars; second segment and third segment

subequal. External apical appendage on second

segment lacking; a small flat button-like structure

on outside towards apex.

Mandibles. Virtually symmetrical, with three

inner teeth, distal two large, basal one much

smaller.

Maxillae. Five-segmented. Stipes as long (first

instars) or longer (second and third instars) than

remaining segments combined, with four or five

setae on inner margin and patch of small spines

on inside near base. Palpus four-segmented; first

segment with an inner apical process; second

segment short; third and fourth segments a little

shorter than first.

Labium. Mentum with sides convex, slightly

wider than long, anterolateral corners pointed,

dorsal surface with numerous small cuticular

spines and row of longer spines towards apex;

small spines mostly absent in first instar.

Prementum long and thin. Palpus two-segmented,

basal segment much shorter than distal. Ligula a

little shorter than distal segment of palpus, slightly

bifid at tip.

Thorax and legs. Prothorax with large dorsal

136 CHS WATTS

sclerite, with sagittal line, with pattern of darker of prothorax. Legs five-segmented, long, visible

lines and dots; ventral surface with a large in dorsal view, with rows of long swimming-setae

subrectangular sclerite, with sagittal line. on femur and tibiotarsus.

Mesothorax and metathorax each with a dorsal Abdomen. Segments 1 to 6 similar in shape,

pair of subtriangular sclerites, smaller than those _ tapering towards the caudal end; segments 7 and 8

FIGURE 17. a, Amphiops queenslandicus: ventral, lateral and dorsal views of pupa. b, Hybogralius hartmeyeri:

ventral and dorsal views of pupa.

LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE

more elongate and slender. Segments 1 to 7

subdivided into three transverse folds, each

segment with transverse row of six pairs of long

setae arising from short dark-pigmented cylindrical

bases on dorsal surface of posterior fold. Pleura 1

to 8 each with a small, lateral, setiferous tubercle.

Segment 8 with two small, dorsal, subtriangular

sclerites at posterior end. Segment 9 slightly

trilobed, with a pair of short one-segmented

urogomphi. Integument densely covered with both

very short and moderately long setae.

Spiracles. Nine pairs; one mesothoracic and

eight abdominal. Mesothoracic and first seven

abdominal spiracles non-functional. Spiracles on

segment 8 annular, large and functional, within

the spiracular atrium.

Intraspecific variation

There is some suggestion of two longitudinal

dark lines on the dorsal surface in some

specimens, and in an occasional specimen the

small third mandibular tooth is virtually absent.

Interspecific variation

In some specimens of Sternolophus there is a

greater development of the lateral abdominal

tubercles into quite prominent finger-like

projections, especially towards the rear; a row of

setiferous papillae across the abdominal segments

instead of small cylinder-like structures; a more

symmetrical nasale; a stronger third mandibular

tooth; and some difference in the shape of the

meso and metathoracic sclerites (the habitus

drawing (Fig. 16) is of this form). These larvae

are S. centralis Watts and/or S. immarginatus

Orchymont. The distribution of specimens would

suggest the latter but not certainly so.

Identification

By rearing from field-collected larvae.

Remarks

The larva of S. marginicollis is one of the

137

few Australian larvae already known (Bertrand

1935) as S. tenebricosus Blackburn, a junior

synonym (Watts 1989), although the

identification was by association, which is

unsafe in this genus because it often has two

or more species common at the same locality.

Other species have been described from Africa

and Japan (see Bertrand 1972). The present

larvae, certainly of S. marginicollis and

probably also including S. immarginatus

Orchymont, do not differ significantly from

the published descriptions and illustrations.

Larvae of Sternolophus are commonly

encountered in Northern Australia and can be

readily separated from other Hydrophilid larvae

by the combination of long thin prementum, long

ligula and mandibles with three teeth.

Specimens examined

Northern Territory: Jabiru, 19/3/98; Manton

Dam, 2/2/99; Ormiston Gorge, 27/1/99.

Queensland: Killymoon Creek, 25 km S

Townsville, 2/2/97.

Sternolophus sp. Northern Territory: Holmes

Jungle near Darwin, 28/1/99. Queensland: 8 km

N Bluewater, 31/10/95; Bohle River, 10 km N

Townsville, 23/3/96; Eubenangee Swamp, 4/2/97;

2km S Mt Molloy, 30/3/96.

ACKNOWLEDGMENTS

I wish to thank Mr Rod Gutteridge, Ms Eloese

O’Grady and Mr Harold Hamer for so ably preparing

the illustrations; Dr David Larson for donating larvae he

collected whilst working in North Queensland; the

South Australian Library staff, Ms Marianne Anthony

and Ms Jill Evans, for procuring references that we did

not already have in the library; and Ms Debbie Churches

for preparing the final manuscript copy. I would

particularly like to thank Dr Steve Donnellan of the

Evolutionary Biology Unit of the South Australian

Museum for undertaking the biochemical matching of

adult and larval Allocotocerus and Regimbartia.

REFERENCES

Anderson, JME. 1976. Aquatic Hydrophilidae

(Coleoptera). The Biology of some Australian

species with descriptions of immature stages in the

laboratory. Journal of the Australian Entomological

Society 15: 219-228.

Archangelsky, M. 1997. Studies on the biology, ecology

and systematics of the immature stages of new world

Hydrophiloidea (Coleoptera: Staphyliniformia). Ohio

Biological Survey Bulletin New Series 12: 1-207.

Berge Henegouwen, AL van. 1975. Description of the

egg-case and larva of Limnoxenus niger (Zschach)

(Coleoptera, Hydrophilidae). Entomologische

Berichten Amsterdam 35: 27-30.

Berge Henegouwen, AL van. 1982. Notes on the larval

stages of some East African Hydrophilinae

(Coleoptera: Hydrophilidae). Entomologische

Berichten Amsterdam 42: 11-16.

138 CHS WATTS

Bertrand, H. 1935. Larves de Coleopteres aquatic de

l’Expedition Limnologique Allemande en Insulinde.

Archiv fur Hydrobiologie 6: 193-215.

Bertrand, H. 1972. Larves et nymphes des Coleopteres

aquatic du globe. F. Paillart: Abbeville, France.

Boving, AC & Henriksen, KL. 1938. The developmental

stages of the Danish Hydrophilidae (Ins. Coleoptera).

Videnskabelige Meddelelser fra Dansk Naturhistorik

Forening i Khobenhaven 102: 27-162.

Fernandez, LA. 1983. Helochares (Sindolus) talarium

sp. nov., redescripcion de Helochares (Helochares)

pallipes (Brulle) y descripcion de sus estados

preimaginales. Coleoptera, Hydrophilidae. Limnobios

2: 439-449,

Gentili, E. 1980. The genera Laccobius and Notohydrus

(Coleoptera, Hydrophilidae) in Australia and New

Zealand. Records of the South Australian Museum

18: 143-154.

Gentili, E. 1992. The Notohydrus of Australia

(Coleoptera, Hydrophilidae). Bolettino Society

Entomologia Italiano, Genova 124: 21-26.

Gentili, E. 1993. Paranacaena Blackburn, 1889: a valid

genus (Coleoptera, Hydrophilidae). Gioruale italiano

di entomologia 6: 285-296.

Gentili, E. 2000. The Paracymus of Australia

(Coleoptera: Hydrophilidae). Records of the South

Australian Museum 33: 101-122.

Hansen, M. 1991. The Hydrophiloid beetles. Phylogeny,

classification and a revision of the genera

(Coleoptera, Hydrophiloidea). Biologiske Skrifter 40:

1-367.

Hansen, M. 1999. Hydrophiloidea (s. str.) (Coleoptera).

In ‘World Catalogue of Insects’, vol. 2: 1-416.

Apollo Books: Stenstrup, Denmark.

Morioka, A. 1955. Description of four Hydrophilid

larvae from Japan (in Japanese). Shin-Konchu 8: 15-

18.

Oliva, A. 1992. Cuticular microstructure in some genera

of Hydrophilidae (Coleoptera) and their phylogenetic

significance. Bulletin de l’Institut Royal des Sciences

Naturelles de Belgique, Entomologie 62: 33-56.

Richmond, EA. 1920. Studies on the biology of aquatic

Hydrophilidae. Bulletin of the American Museum of

Natural History 42: 1-94.

Spangler, PJ. 1966. A description of the larva of

Derallus rudis Sharp, (Coleoptera, Hydrophilidae).

The Coleopterists Bulletin 20: 97-100.

Spangler, PJ. 1973. A description of the larva of

Hydrobiomorpha casta (Coleoptera: Hydrophilidae).

Journal of the Washington Academy of Science 63:

160-164.

Watts, CHS. 1963. The larvae of Australian Dytiscidae

(Coleoptera). Transactions of the Royal Society of

South Australia 87: 23-40.

Watts, CHS. 1987. Revision of Australian Berosus

Leach (Coleoptera: Hydrophylidae). Records of the

South Australian Museum 21: 1-28.

Watts, CHS. 1988. Revision of Australian Hydrophilus

Muller, 1764 (Coleoptera: Hydrophilidae). Records

of the South Australian Museum 22: 117-130.

Watts, CHS. 1989. Revision of Australian Sternolophus

Solier (Coleoptera: Hydrophilidae). Records of the

South Australian Museum 23: 89-95.

Watts, CHS. 1990. Revision of the Australian

Hydrobiomorpha Blackburn (Coleoptera:

Hydrophilidae). Records of the South Australian

Museum 24: 35-42.

Watts, CHS. 1995. Revision of the Australasian genera

Agraphydrus Regimbart, Chasmogenus Sharp and

Helochares Mulsant (Coleoptera: Hydrophilidae).

Records of the South Australian Museum 28: 113-

130.

Watts, CHS. 1998a. Revision the Australian Amphiops

Erichson, Allocotocerus Kraatz and Regimbartia

Zaitzev (Coleoptera: Hydrophilidae). Records of the

South Australian Museum 30: 93-106.

Watts, CHS. 1998b. Revision of Australian Enochrus

(Coleoptera: Hydrophilidae). Records of the South

Australian Museum 30: 137-156.

EMBOLOCEPHALUS YAMAGUCHI (BRINKHURST, 1971) (CLITELLATA:

TUBIFICIDAE) FROM SOUTH AUSTRALIAN STREAMS

A. M. PINDER & P. K. MCEvoy

Summary

Papillate tubificids from streams in the Mount Lofty Ranges are identified as Embolocephalus

yamaguchii (Brinkhurst, 1971) and represent the first records of this largely Holarctic genus from

Australia. Collection details are provided for records of a second papillate species, identified as

belonging to the genus Spirosperma.

EMBOLOCEPHALUS YAMAGUCHI (BRINKHURST, 1971) (CLITELLATA: TUBIFICIDAE)

FROM SOUTH AUSTRALIAN STREAMS

AM PINDER & PK McEVOY

PINDER, AM & McEVOY, PK. 2002 Embolocephalus yamaguchii (Brinkhurst, 1971)

(Clitellata: Tubificidae) from South Australian streams. Records of the South Australian

Museum 35(2): 139-145.

Papillate tubificids from streams in the Mount Lofty Ranges are identified as

Embolocephalus yamaguchii (Brinkhurst, 1971) and represent the first records of this largely

Holarctic genus from Australia. Collection details are provided for records of a second papillate

species, identified as belonging to the genus Spirosperma.

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

Western Australia 6946. PK McEvoy, Australian Water Quality Centre, Private Mail Bag 3,

Salisbury, South Australia 5108. Manuscript received October 2001.

Most tubificids with papillate body walls are

contained within a complex of genera

(including Embolocephalus Randolph and

Spirosperma Eisen) established, or re-

established, following revision of the genus

Peloscolex Leidy (Brinkhurst 1991; Brinkhurst

& Wetzel 1984; Holmquist 1978, 1979). These

genera have a largely Holarctic distribution,

with few records from southern continents. The

latter consist of a record of the otherwise

European Embolocephalus velutinus (Grube,

1879) from Venezuela (Dumnicka, 1983), an

identification questioned by Brinkhurst and

Marchese (1989), and papillate tubificids of

uncertain identity from Peru (Brinkhurst &

Marchese 1989). The North American species

Quistadrilus multisetosus has been recorded as

far south as Mexico. In Australia, Timms (1978)

recorded Peloscoiex (sic) from a lake in

Tasmania and Pinder & Brinkhurst (1994, 2000)

noted papillate tubificids from Victoria and

New South Wales. The latter were all immature

so descriptions were not possible, but they were

tentatively assigned to Spirosperma, based on

the presence of papillae and the form of the

chaetae. Collection details for this species,

including the first records from South Australia,

are presented below for the first time. Other

specimens, collected from streams in the Mount

Lofty Ranges in South Australia, are herein

described and identified as Embolocephalus

yamaguchii (Brinkhurst, 1971), a species

otherwise known only from Lake Biwa in

Japan.

METHODS

Serially sectioned specimens were cut at 6 um

and stained in haematoxylin and eosin.

Measurements were taken using an eyepiece

graticule on a Zeiss Jenamed 2 compound

microscope calibrated with an Olympus stage

micrometer. Abbreviations used in the line-

drawings are a: atrium, cp: cuticular pad, e: large

epidermal papillae, ed: ejaculatory duct, ff: female

funnel, mf: male funnel, p; prostate, pe: penis, sa:

spermathecal ampulla, sc: spermathecal chaeta,

scg: spermathecal chaetal gland, v: vas deferens.

Specimens are either in the senior author’s

collection (AP) or the Australian Water Quality

Centre collection (AWQC); returned to WSL

Consultants (WSL) or Australian Water

Technologies (AWT); or deposited with the South

Australian Museum (SAM) or the Museum of

Victoria (NMV).

SYSTEMATICS

Embolocephalus Randolph, 1892

Type species

Embolocephalus velutinus (Grube, 1879)

Diagnosis

From Holmquist (1978) and Brinkhurst (1981).

Prostomium and sometimes first segments

retractable within the rest of body. Body wall with

epidermal papillae, generally concealed by a

140

secreted layer with embedded foreign particles.

Hair chaetae present dorsally, usually broad and

sabre-like, with pectinate crotchet chaetae. Bifid

and/or simple pointed chaetae present ventrally,

usually both in anterior bundles. Spermathecal

chaetae modified on X, lying in a well-developed

chaetal sac. Male ducts each with a long winding

vas deferens, usually narrower entally and broader

(up to twice the width) ectally, entering the atria

apically. Atria long and tubular to crescentic,

broadest near the prostate union, tapering to a

short ejaculatory duct. One large prostate gland

joining each atrium towards the middle of the

latter. Penes well developed without thickened

cuticular sheaths. Spermathecae with ovoid to

oblong ampullae, well set off from the ducts, with

pores usually more or less in front of and lateral

to the spermathecal chaetal sacs.

Distribution

Europe, North America, Japan, Kamchatka,

?7Venezuela and now Australia.

Embolocephalus yamaguchii (Brinkhurst,

1971)

Peloscolex yamaguchii Brinkhurst, 1971: 505

Peloscolex sp. Yamaguchi, 1953: 295

Embolocephalus yamaguchii (Brinkhurst, 1971)

Ohtaka, 1994: 52; 1995: 174

Material examined

South Australian specimens. SAM E3095—

3101: 3 mature specimens serially sectioned, 3

mature dissected and 1 immature whole-mounted,

Aldgate Creek at Strathalbyn Road, Mylor,

tributary of the Onkaparinga River (AWQC site

3217), 35°02’S 138°45’E, samples E33, 3 June

1997 and F32, 27 Oct 1997. AP: 1 mature serially

sectioned, collected as above. SAM E3102-3103:

1 mature in alcohol, Onkaparinga River at

Houlgraves Weir (AWQC site 3205), 35°05’S

138°43’E, sample E37, 3 June 1997, and 1 mature

whole-mounted from same site, sample F37,

28 Oct 1997. SAM E3104: 1 mature in alcohol,

Aldgate Creek at Aldgate Valley Road (AWQC

site 13022), 35°02’S 138°45’E, sample H41, 2

Dec 1998. AWQC: 5 immature in alcohol,

Onkaparinga River at Hack Bridge (AWQC site

3212), 35°03’S_ 138°45’E, sample H43, 1 Dec

1998. AWQC: 1 immature in alcohol, Lenswood

Creek at gauging station (AWQC site 3208),

AM PINDER & PK McEVOY

34°56’S 138°50’E, sample OCB148, 17 Jul 2000:

AWQC: 2 mature in alcohol, Onkaparinga River

at Silver Lake Road (AWQC site 13023), 35°04’S

138°45’E, sample OCB181, 17 Oct 2000.

Collections by V. Tsymbal, P. McEvoy, A. Lang

and S. Wade (Australian Water Quality Centre,

South Australia).

Japanese specimens. From collection of A.

Ohtaka (Hirosaki University, Japan): 1 mature

specimen serially sectioned, 1 mature whole-

mounted and 2 mature in alcohol (now dissected),

Lake Biwa, Japan.

Habitat

This species has been collected only from a

sixth order reach of the Onkaparinga River

upstream of Mount Bold Reservoir and from two

fourth order tributaries, between 245 and 295

metres above sea level. Specimens were collected

from both riffle and edge habitats, with substrates

ranging from those composed mainly of bedrock,

boulder or cobble to those dominated by finer

sediments and/or detritus. Willow root mats are a

common feature on the stream bed of most sites.

Riparian zones had an overstorey of either

indigenous Eucalyptus species and Acacia

melanoxylon and/or introduced willows (Salix

spp.). Land uses include grazing, horticulture and

urban settlement. Apparently similar habitats in

adjacent catchments were sampled with the same

sampling intensity without finding E. yamaguchii.

Description of Australian specimens

Length of preserved specimens 15.6—25.5 mm,

width at clitellum 0.7—1.1 mm. Each pre-clitellar

segment with a band of tall, broad epidermal cells

forming a raised transverse ridge in the middle of

the segment, pronounced dorsally and laterally but

virtually absent ventrally. Body surface with

foreign material adhered to elongate ovoid

papillae covering the entire body except for the

clitellum (Fig. 1). Papillae mostly 15-22 um long

on pre-clitellar segments (measured on the longest

axis, including foreign particles) and 20-34 pm

long on post-clitellar segments, and mostly 17—

25 um high (with most of the height constituted

by the foreign material). Ridges of anterior

segments with fewer or no papillae. Each segment

also with 2 rings of papillae that are taller and

broader than normal (up to 40 um wide and

40 um high including foreign material), which

give the appearance of darker (or sometimes

lighter) bands around each segment, 1 ring about

at the level of the chaetae and 1 anteriorly on each

segment, most pronounced on posterior segments.

EMBOLOCEPHALUS YAMAGUCHII FROM SOUTH AUSTRALIAN STREAMS 141

Dic

FIGURE 1. E. yamaguchii A, Enlarged view of body

wall of segments towards the posterior end of SAM

E3098 (prior to slide mounting) showing normally sized

and enlarged (e) papillae; B, Portion of SAM E3104

showing lateral view of anterior segments (top),

clitellum (centre) and post-clitellar segments (bottom).

Scale bar for A 100 jim, B 500 um.

Other (possibly sensory) papillae present with the

foreign material adhered only to the sides, with

the top of the papillae exposed, arranged in ill-

defined rings around each segment (about 10 of

these papillae per ring), usually 1 ring at the level

of the chaetae and 1 or more rings between the

chaetae and intersegmental furrow. These partially

naked papillae are about the same size as the

shorter type of fully encased papillae and are

sometimes visible as darker or paler spots. A pad

of tall, thin epidermal cells, with cuticle 10 times

thicker (up to 17 wm) than elsewhere, present

medially on the spermathecal segment, adjacent to

the anterior spermathecal chaeta. Clitellum

covering posterior half of X and all of XI, paler

than other segments due to a smooth even coating

of soft white granular material (Fig. 1).

Prostomium partially retracted into first

segment in fixed animals. Pharynx with dorsal

muscular pad in II to Il, with pharyngeal gland

cells on pharynx and on septa 3/4 to 4/5.

Oesophagus from IV to VI, widening into broader

thicker-walled intestine in VII.

Ventral chaetae 130-211 um long and 7-14 pm

wide at nodulus, generally smallest in pre-clitellar

bundles. Ventral bundles on II to VII usually with

1 bifid and 1 simple pointed chaeta, rarely 1 extra

bifid chaeta and/or 1 or 2 extra simple pointed

chaetae. Bifid chaetae with teeth equal in width

but upper tooth up to 1.5 times longer than lower

(Fig. 2B). Posterior ventral bundles with 1 (rarely

2) bifid chaetae, with upper teeth becoming

shorter than lower and lower tooth greatly

expanding in width and becoming notably

recurved (Fig. 2C). Dorsal chaetal bundles

normally with 2 to 4 slightly curved hair chaetae,

220-420 pm long and 4—7 um wide, with finely

plumose shafts and blunt, slightly swollen, tips,

with an equivalent number of short (32-75 um

long and 1—2 um wide) finely pectinate crotchets

with thin parallel outer teeth (Figs 2D,E).

Posterior dorsal bundles with fewer of both types

of chaetae. Chaetae of spermathecal segment

modified (Figs 2A, 3A,B), ventral chaetae absent

in XI.

Genitalia paired (Figs 2, 3B). Testes

anteroventral in X, ovaries anteroventral in XI.

Male funnels large, ciliated vasa deferentia broad

near funnels (35 um), narrowing shortly thereafter

(17-20 wm) and widening slightly ectally

(25-33 um), highly coiled, joining ental end of

atria. Atria each with a narrow lumen and layer of

tall glandular lining cells, crescentic, broadest

where a single prostate gland joins (slightly entad

of medial), tapering at either end, the ectal end

forming a short ejaculatory duct which enters the

penis apically. Atrial muscle layer thin, wider

where prostate joins. Atrium, including

ejaculatory duct, measuring 800 x 110 pm to 1000

x 185 um. Prostate glands voluminous, sometimes

ballooning into following segment. Penes broad

entally, narrowing at about one-third of the

distance from the tip to form a narrower ectal end,

142 AM PINDER & PK McEVOY

D E

FIGURE 2. E. yamaguchii. A, Reconstruction of genitalia (with organs drawn in the same plane for simplicity)

from serial sections of SAM E3095; B, tips of anterior ventral chaetae; C, tip of posterior ventral chaeta; D, dorsal

pectinate chaeta; E, tips of dorsal hair chaetae, with plumosity shown for one hair. Scale bar for A 200 um.

penes within thin-walled muscular penis sacs

attached to the dorsolateral body wall by

numerous muscle fibres. Penes and penis sacs

with cuticle no thicker than on body wall (ie

without penis sheaths). Male pores in line with

ventral chaetae on anterior half of XI, within a

common transverse depression of the ventral body

wall. Spermathecal ampullae variable in size

(770-1350 um long), each containing 1 or 2 long

thin spermatozeugmata (length of only one whole

spermatozeugmata measured, 660 x 45 um, but

others 80 um wide in cross-section, indicating

greater length). Spermathecae with stout ducts

(300-370 um) that are constricted ectally, leading

to pores anterior on X slightly medial to line of

ventral chaetae. Spermathecal chaetae long (510-

670 um) and thin (10 pm) with grooved tips, ectal

half lying in well-developed chaetal sacs. The sacs

tall (220-300 um) and muscular, with inner lining

tissue that can be everted with the tip of the chaeta

EMBOLOCEPHALUS YAMAGUCHII FROM SOUTH AUSTRALIAN STREAMS 143

Vill = IX X

FIGURE 3. E. yamaguchii. Reconstruction of genitalia (with organs drawn in the same plane for simplicity) from

serial sections of A, SAM E3096 and B, SAM E3097. Scale bar for A 200 um.

to form a large papilla and with muscular layer

extending to cover the ental end of the chaeta. Sacs

with a gland attached dorsally, varying in extent of

development (between and within individuals).

Spermathecal chaetal sacs located asymmetrically,

both opening slightly ventral to the line of somatic

ventral chaetae, 1 immediately posterior to

spermathecal pore on one side and the other close

to 11/12 in front of the sperm funnel on other side.

Female funnels posterolateral on XI.

144

Remarks

The presence of simple-pointed chaetae on

segments II to VIII, separate openings for the

spermathecal ducts and spermathecal chaetae,

dorsolateral ridges on anterior segments and

asymmetrical location of the spermathecal chaetae

clearly ally these Australian specimens with the

Japanese species E. yamaguchii rather than with

any of its congeners. We initially considered that

the Australian specimens were either conspecific

with E. yamaguchii or represented a very similar

sister-species. The latter was seen as more likely

since E. yamaguchii is known only from Lake

Biwa in Japan and other Embolocephalus seem to

have fairly restricted Holarctic distributions.

However, a thorough comparison of E.

yamaguchii (from the description in Ohtaka 1994

and material seen by us) and the Australian

specimens led to the conclusion that there was no

morphological justification for a new species

description. The only differences between the

Australian and Lake Biwa specimens are the size

of the worms and the size of some genital organs.

None of the Australian specimens reach the

maximum length of 35 mm measured for the E.

yamaguchii neotypes (Ohtaka 1994), but most

Lake Biwa specimens are less than 30 mm

(Ohtaka pers. comm.). The vasa deferentia of the

Australian specimens are narrower than those (20—

24 um wide entally, 40 pm ectally) recorded for

E. yamaguchii by Ohtaka (1994) and the ectal

portion is not so enlarged relative to the ental

portion. However, the vasa deferentia of Lake

Biwa specimens measured by us (as narrow as

13 pm entally to a maximum of 25 um ectally)

more closely match measurements from the South

Australian material. Other components of the

genitalia (atria, spermathecal ampullae and

spermathecal chaetae) are slightly larger in the

Australian specimens than recorded by Ohtaka

(1994). The raised epidermal pad was not noted

for E. yamaguchii by Ohtaka (1994) but was

present on Lake Biwa material seen by us and is

visible in Fig. 4B of Ohtaka (1994). This feature

is not known for other Embolocephalus species.

These specimens represent the first records of

this species and genus in Australia and one of

the few records of the genus from southern

continents. The known distribution of this

species (Lake Biwa in Japan and the Mount

Lofty Ranges in South Australia) is exceptional

within the Australian tubificid fauna. Other

tubificids found in Australia are either endemic

or cosmopolitan (occurring in most, if not all,

other continents).

AM PINDER & PK McEVOY

?Spirosperma sp.

Material examined

All specimens immature. WSL consultants

voucher AN30: 2 in alcohol, Old Namoi River at

Bullerawa Station, (Duncan’s Junction) New

South Wales, 30°18’05”S 149°04’00”E, 27 Jun

2000. AWT: 1 specimen, Georges River at

Cambridge Avenue, New South Wales, 33°58’S

150°54’E, 1 Apr 1996; 1 specimen, Peach Tree

Creek, at Weir Reserve, New South Wales,

33°45’S 150°41’E, 17 Jul 1996; 1 specimen,

Second Ponds Creek downstream of Rouse Hill

Sewage Treatment Ponds, 33°40’S 150°55’E,

1 Apr 1996. AP: 1 whole-mounted on slide,

Plenty River at Lower Plenty, Victoria, 37°44’S

145°06’E, 10 Apr 1994, coll. S. Schreiber

(formerly Monash University); Glenelg River at

Rocklands, Victoria, 37°14’S 141°57’E, 3 Jun

1994, coll. S. Schreiber. NMV F81865: 1 in

alcohol, LaTrobe River at Moe-Willow Grove

Road Bridge, Victoria, 38°11°18"S 146°15’12”E.

NMV F81864: 1 in alcohol, Curdies River,

Victoria, 38°20’S 143°08’E, 7 Feb 1992, coll. G.

Quinn (formerly Monash University). AWQC

voucher 5665: 1 in alcohol, Pilby Creek outlet

(AWQC site 13095), South Australia, 33°59’S

140°53’E, 24 Nov 1998, coll. D. Schulze and T.

Venus.

Brief description

Prostomium retractable within mouth. Body

wall densely papillate, with foreign material

A C

FIGURE 4. Spirosperma sp. A, ventral chaetae of

anterior to mid-body segments; B, ventral chaeta of

posterior segment; C, tips of dorsal crotchet chaetae.

EMBOLOCEPHALUS YAMAGUCHII FROM SOUTH AUSTRALIAN STREAMS 145

attached to most papillae. Some papillae with

foreign matter on lateral walls only. Without rings

of distinctly larger papillae. Ridges of enlarged

epidermal cells absent on anterior segments.

Ventral chaetae (Fig. 4) all bifid, 3 to 6 per

bundle, with upper teeth 2 to 3 times as long as

lower teeth in anterior and mid-body segments,

1.5 times as long as lower teeth in posterior

segments (Fig. 4). Ventral chaetae of first few

segments usually slightly longer and thicker than

the rest. Dorsal bundles with 1 to 3 smooth hair

chaetae and 1 to 3 very small crotchet chaetae

with finely bifid ends and 1 or 2 intermediate

teeth (Fig. 4).

Remarks

This species lacks simple-pointed chaetae

anteriorly and so has been tentatively identified as

a Spirosperma rather than an Embolocephalus, but

mature specimens are required to confirm the

generic affiliation. It appears to be widespread in

rivers of southeastern Australia.

ACKNOWLEDGMENTS

Specimens of E. yamaguchii were collected as part

of the Monitoring of River Health Initiative funded by

the Land and Water Resources Research and

Development Council and Environment Australia.

Funding from the Wildlife Conservation Fund of the

Department for Environment and Heritage (South

Australia) enabled identification to species of

oligochaetes from the above project. Laboratory and

microscope facilities were made available by Stuart

Halse (Department of Conservation and Land

Management). Specimens of Spirosperma were made

available by Australian Water Technologies, Alena

Glaister at Monash University and Kylie Swingler at

WSL Consultants (specimens from a Department of

Land and Water Conservation NSW project). Akifumi

Ohtaka (University of Hirosaki, Japan) kindly

provided material of E. yamaguchii from Lake Biwa

and engaged in useful discussions with the authors.

Gordon Thomson (Murdoch University, Perth)

performed the serial sectioning and Jane McRae

(Department of Conservation and Land Management)

assisted with photography.

REFERENCES

Brinkhurst, RO. 1991. A phylogenetic analysis of the

Tubificinae (Oligochaeta: Tubificidae). Canadian

Journal of Zoology 69: 392-397.

Brinkhurst, RO. 1981. A contribution to the taxonomy

of the Tubificinae (Oligochaeta: Tubificidae).

Proceedings of the Biological Society of Washington

94: 1048-1067.

Brinkhurst, RO & Marchese, M. 1989. ‘Guide to the

freshwater aquatic Oligochaeta of South and Central

America’. Asociacion Ciencias Natureles del Litoral:

Santo Tome, Argentina.

Brinkhurst, RO & Wetzel, MJ. 1984. Aquatic

Oligochaeta of the world: Supplement. A catalogue

of new freshwater species, descriptions and revisions.

Canadian Technical Report of Hydrography and

Ocean Sciences 44: 1-101.

Dumnicka, E. 1983. Tubificidae (Oligochaeta) from

subterranean waters, with description of two new

genera. Bijdragen tot de Dierkunde 53: 255-261.

Holmquist, C. 1978. Revision of the genus Peloscolex

(Oligochaeta: Tubificidae) 1. Morphological and

anatomical scrutiny; with discussion on the generic

level. Zoologica Scripta 7: 187-208.

Holmquist, C. 1979. Revision of the genus Peloscolex

(Oligochaeta: Tubificidae) 2. Scrutiny of the species.

Zoologica Scripta 8: 37-60.

Ohtaka, A. 1994. Redescription of Embolocephalus

yamaguchii (Brinkhurst, 1971) comb. nov.

(Oligochaeta: Tubificidae). Proceedings of the

Japanese Society of Systematic Zoology 52: 34-42.

Pinder, AM & Brinkhurst, RO. 1994. ‘A Preliminary

Guide to the Identification of the Microdrile

Oligochaeta of Australian Inland Waters’.

Cooperative Research Centre for Freshwater

Ecology: Albury.

Pinder, AM & Brinkhurst, RO. 2000. A review of the

Tubificidae (Annelida: Oligochaeta) from Australian

inland waters. Memoirs of Museum Victoria 58: 39-

75.

Timms, BV. 1978. The benthos of seven lakes in

Tasmania. Archiv fiir Hydrobiologie 81: 422-444.

EARLY ABORIGINAL FISHING TECHNOLOGY IN THE LOWER

MURRAY, SOUTH AUSTRALIA

PHILIP A. CLARKE

Summary

The early Aboriginal fishing technology of the Lower Murray region of South Australia is described

and compared with other areas in south-eastern Australia where fish was a dominant dietary

component of hunters and gatherers. This is a study of cultural geography, the chief concern being a

description of hunting and gathering techniques and their significance to Aboriginal occupation of

the landscape.

EARLY ABORIGINAL FISHING TECHNOLOGY IN THE LOWER MURRAY,

SOUTH AUSTRALIA

PHILIP A CLARKE

CLARKE, PA. 2002. Early Aboriginal fishing technology in the Lower Murray, South

Australia. Records of the South Australian Museum 35(2): 147-167.

The early Aboriginal fishing technology of the Lower Murray region of South Australia is

described and compared with other areas in south-eastern Australia where fish was a dominant

dietary component of hunters and gatherers. This is a study of cultural geography, the chief

concern being a description of hunting and gathering techniques and their significance to

Aboriginal occupation of the landscape.

PA Clarke, Science Division, South Australian Museum, North Terrace, Adelaide, South

Australia 5000. Manuscript received 7 December 2001.

INTRODUCTION

Aboriginal fishing technology in the Lower

Murray region is discussed from the pre-European

period to the early years of European settlement.

A major aim is to describe how hunting and

gathering techniques used by Lower Murray

people made their region culturally distinctive

(Fig. 1).! This region is defined as the area from

Cape Jervis in the west, east to Wellington on the

Murray River and south to Kingston, taking in the

whole of the Lower Lakes, Coorong and

associated coastal belt. It is estimated that just

prior to European settlement, the region supported

a population of 5000 Aboriginal people, although

this figure was probably seriously affected by two

early waves of smallpox (Brown 1918: 230;

Campbell 2002: 119-133; Clarke 1994: 57-63;

1995: 156, footnote 1; Gale 1969). The

descendants of these people, many of whom still

live in the region, generally call themselves

Ngarrindjeri (Berndt & Berndt 1993; Clarke 1994;

Hemming & Jones 2000; Jenkin 1979).?

In the Lower Murray region the Murray River

flows from Murray Bridge to Lake Alexandrina

through an open valley cut across a very low and

flat limestone karst plain, which is less than 30

metres above sea level (Fenner 1931: 81-83;

Twidale 1968: 148-149, 383-384). Below

Wellington, the river becomes two large lakes

(Alexandrina and Albert) and a series of channels

in the form of a delta, eventually exiting behind

scattered islands at the Murray Mouth.’ Here, the

river meets the Coorong, which drained the South

East region of South Australia before European

intervention.* The sea, winds and tides combine to

drive the river back with heavy sand dune

systems, called Sir Richard and Younghusband

Peninsulas (northwest and southeast parts,

respectively).° From the point of view of the early

Aboriginal inhabitants, the delta of the Lower

Murray provided the region with many kilometres

of shoreline for hunting and gathering activities.

The climate of the Lower Murray region is

influenced by the powerful ‘controls’ of its

temperate latitude, the proximity of the sea and

the relief of the land (Fenner 1931: 125; Howchin

1909: 142; Penney 1983: 85-93; Schwerdtfeger

1976: 75-86). Upstream, long sections of both the

Murray River and its main tributary, the Darling

' Hunting and gathering practices in the region since the 1940s are discussed elsewhere (Clarke 2002).

? The Ngarrindjeri ( = Narrinyeri) were formerly made up of descent groups who spoke one of several dialects, such as Ramindjeri, Yaraldi

( = Jarildekald) and Tangani ( = Tanganekald).

* Lake Alexandrina is called ‘Lake Victoria’ on some early official maps (Cockburn 1984: 7).

‘The Coorong was formerly known as the ‘South East Branch’ in reference to Lake Alexandrina, which it joined at Pelican Point (Cuique [R.

Penney] in the South Australian Magazine, September 1842, vol. 2: 18-23).

Due to the action of the ocean currents, the location of the river exit into the sea is constantly moving; at present it is migrating northwards towards

Goolwa at the rate of several metres per year (F. Tuckwell, pers. comm.). The complete disappearance of Barker Knoll at the Mouth as early as

1859 (Linn 1988: 78) indicates that some movement is a natural feature.

148

KANGAROO

ISLAND

PA CLARKE

VICTOR

HARBOR

& No MURRAY

Cun MOUTH POINT

& MCLEAY

COORONG

SOUTHERN

OCEAN

INGSTON

NORTHERN

TERRITORY

WESTERN

AUSTRALIA

FIGURE 1. The Lower Murray cultural region.

EARLY ABORIGINAL FISHING TECHNOLOGY

River, flow through semi-arid regions.

Nevertheless, the Lower Murray region is entirely

contained within the high rainfall area of South

Australia, receiving 350-750 mm per year. It

comes under the rainfall shadow of the Mount

Lofty Ranges to the west, with precipitation also

increasing near the coast. Annual average

temperature throughout the region is less than

18°C, with the greatest range of temperatures

being during the summer months (Fenner 1931:

65, 126; Griffin & McCaskill 1986: 50-51; Laut

et al 1977).

SOURCES OF ABORIGINAL ENVIRONMENT USE DATA

The literature of Aboriginal hunting and

gathering technology for the region is based on

four main sources of data — the archaeological

record, the historical ethnographic record,

scientific analysis of the properties of naturally

occurring substances and contemporary research

with Aboriginal people. This paper deals mainly

with historical ethnographic sources of

information, which includes artefacts collected

from living people. Archaeological investigations

provide evidence of the importance of fish in the

diet of coastal/riparian Aboriginal groups, with an

indication of the material cultural items and the

main species fished.° Biological analysis of human

bone also assists in determining the pre-European

diet (Pate 1997, 1998, 2000). Scientific methods

of analysis of food sources, such as those

determining the pharmacological and nutritional

properties, can illustrate their potential human

uses. The literature investigating the usefulness of

Australian fish as food and medicine includes

Brand Miller et al (1993: 222-223), Clarke (1989:

3) and Isaacs (1987: 153-164). Nevertheless,

cultural perceptions influence the potential use of

plants and animals, with not all available

resources being fully utilised. To understand how

Lower Murray Aboriginal people used their

environment, and thereby moulded their cultural

landscape, we must at present rely primarily on

records made by early European observers,

supplementing this with ethnographic data from

contiguous riparian/coastal regions.

Although useful data on pre-European plant

use, such as species identification, method of use

and seasonality, can be obtained from

149

contemporary fieldwork with Aboriginal people,

there are significant post-European changes in

Lower Murray Aboriginal culture and in the

physical landscape itself. For instance,

information received from Lower Murray

Aboriginal people in recent times on bush foods

used during the last 50 to 60 years indicates far

less use of roots than before (Clarke 1988: 64). In

contrast, other indigenous foods such as fish,

waterfowl, kangaroos, emus and berries continued

to be used in restricted contexts. Nevertheless, the

introduction of foreign species, such as European

carp, and environmental changes in the waterways

have decreased numbers of some Australian fish

species. The availability of European-type foods,

especially flour, potatoes and canned foods,

obtained from missions, farm stations and towns

led to a decline in many indigenous food sources.

The decrease in ‘bush food’ consumption was

particularly marked for those species requiring

significant labour expenditure. Other foods were

replaced by European varieties with more

favourable properties. The bulrush root, for

example, contains a great deal of fibre that makes

eating difficult. Unlike earlier Aboriginal foods,

many European sources were available at all times

of the year because they were imported.

The detailed recording of Aboriginal culture in

the Lower Murray region started with the German

Heinrich A. E. Meyer, who ran a mission at

Encounter Bay during the late 1830s and early

1840s (Meyer 1843, 1846). From him we obtain a

Ramindjeri-speaker’s view of Lower Murray

culture. The Englishman George Taplin

established an Aboriginal mission at Point

McLeay on the south-western shore of Lake

Alexandrina in 1859. In the 1870s he published

two main books on Ngarrindjeri ( = Narrinyeri)

culture, based on records in his journals (Taplin

1859-79, 1874, 1879). Taplin used Ramindjeri

sources as a guide to his research, although local

Aboriginal groups living around Point McLeay

were mainly Yaraldi-speakers. He described

people who were feeling the impact of European

expansion and the considerable changes it had

brought upon them and the Lower Murray

landscape, particularly at the northern end. One of

Taplin’s initiatives to improve their circumstances

was to establish a local fishing industry for

Aboriginal people at the mission (Jenkin 1979:

97-98, 110-111).’

(1978) and Pretty et al (1983),

There are many entries in Taplin’s journals concerning the establishment of the fishing industry. The important references are 21 September 1859,

18 October 1859, 25 November 1859, 22 December 1859, 11 January 1860, 7 February 1860, 15 February 1860 and 1 March 1860.

150

Adding to the missionary records in the 19th

century are the recordings of colonists such as

George French Angas (1847a, 1847b), William A.

Cawthorne (1844 [1926]), Dirk M. Hahn (1838-

39), Richard Penney (1840-43)§ and William

Wells (1852-55). During this century, various

anthropologists studied Lower Murray Aboriginal

culture: Ronald M. Berndt (1940; with Catherine

H. Berndt 1993), Alison Harvey (1939ms, 1943)

and Norman B. Tindale (1930-52, 1931-34,

1934-37, 1935, 1937, 1938, 1938-56, 1941,

1981, 1987; with Mountford 1936; with Pretty

1980). Their accounts are rich in detail concerning

past hunting and gathering practices, filling many

gaps that were left in the early historical record.

They also contain examples of post-European

innovations by indigenous people. Largely due to

the interests and efforts of the Director of the

South Australian Museum, Edward C. Stirling,

and Ethnology Curator, Norman B. Tindale, the

South Australian Museum possesses a large

collection of early artefacts from the Lower

Murray region.® In the 1980s the present author

commenced studying Aboriginal relationships to

the physical landscape (Clarke 1985a, 1985b,

1986a, 1986b, 1987, 1988, 1989, 1994 chapter 4,

1998, 1999b, 2001a). The region and Aboriginal

relationship to it also featured in the Hindmarsh

Island Bridge controversy (Bell 1998; Stevens

1995; Wilson 1998). The attention that scholars

have directed towards the Lower Murray has

meant that it is ethnographically one of the best-

described regions in southern Australia. The

biases within this literature are discussed

elsewhere (Clarke 1994 section 11.3, 1999a: 149-

151, 2001a: 19-20).

Ear_y ABORIGINAL FISHING TECHNOLOGY

In the pre-European period the high Aboriginal

population along the river frontage and the coastal

zone of South Australia was in sharp contrast to

the sparse population in the interior. In the Murray

Basin region the distribution of people was

estimated to be 0.4-0.5 square kilometres for each

PA CLARKE

individual, in contrast to 31-88 square kilometres

required in Central Australia (Jones 1974: 326;

Lawrence 1968: 72-73; Maddock 1972 [cited

Rose 1987: 22-23]; Meggitt 1962 [cited Abbie

1976: 46-47]; Smith 1980: 68-90; Tindale 1981:

1860). It has been demonstrated that the physical

environment of the Lower Murray formed both a

natural and cultural region (Clarke 1994). Stable

isotope research on human bone indicates that, in

general, neither people nor foods were moving

from the Murray Bridge area to the adjacent

Murray Mouth and Coorong (Pate 1998). The

Lower Murray therefore formed a separate hunting

and gathering region. Nevertheless, some aspects

of the material culture and natural resource use by

Aboriginal people here was similar to that of

riverine and coastal communities described from

elsewhere in Australia (Hallam 1975; Lawrence

1968: 85-122; Lourandos 1997: 195-243).

In the 1930s Tindale mapped many of the

favourite fishing places and lookouts in southern

South Australia, from informants such as Louisa

Eglinton (Narangga woman), Milerum (Clarence

Long, Tangani man) and Albert Karloan (Yaraldi

man).'° In the Tangani language, elevated areas

used as ‘watch places’ were called popaldi,

whereas in Ramindjeri they were koinkoinj."

Older men generally used these when looking out

for fish shoals and bird movements, as well as for

monitoring the movements of their neighbours by

observing the smoke from their campfires. In

comparison to adjacent regions, such as the

Murray Mallee and Mount Lofty Ranges, the

Lower Murray landscape offered Aboriginal

people abundant opportunity to make fish a more

significant food source.

Before the arrival of Europeans, Aboriginal

people in southern Australia did not widely use

the fishhook and line (Curr 1883: 110; Davies

1881: 129; Eyre 1845, vol. 2: 266-267; Massola

1956; Meyer 1846 [1879: 192]; Olsen 1991: 5).

Nevertheless, the use of bone bi-points or fish

gorges (muduk) and fishing lines has been

recorded from along the Murray River (Flood

1999: 48; Gerritsen 2001; Pretty 1977: 321-322).

Similar items have been recorded from the

* Newspaper articles by Richard Penney between 1840 and 1843 are reprinted in the Journal of the Anthropological Society of South Australia 1991:

1-107.

° From 1988 to 1999, a major exhibition on Aboriginal culture in the Lower Murray, called ‘Ngurunderi, a Ngarrindjeri Dreaming’, was open at the

S.A. Museum (Hemming & Jones 2000).

'° Tindale Aboriginal Site Maps of southern South Australia, South Australian Museum Archives, Adelaide. Also see Tindale (1935-60: 15).

"Tindale (no date, ‘Milerum’, Stage A, #9, Archives, South Australian Museum, Adelaide).

2 Tindale (1934-37: 285) recorded a description by Karloan (Karlowan) of fishhooks being made from bone, gum and stick, but this was possibly a

post-European development.

EARLY ABORIGINAL FISHING TECHNOLOGY 151

Gippsland and Geelong areas of southern Victoria

(Smyth 1878 vol. 1: 391). Shell fishhooks have

also been recovered from coastal shell middens in

eastern Victoria (Mulvaney & Kamminga 1999:

292).

After Europeans arrived, Aboriginal people in

the Murray River and Lower Murray regions

adopted new fishing techniques and appeared to

have made their own version of the fishhook.!? In

1938 Tindale recorded Albert Karloan who said:

Our fish hook was made from a bone of a

kangaroo tied like a real fish hook to another

piece which was the point but our old folks used

the fish spear all the time; nothing was as good

as the real fishing spear! We walked along the

banks & got fish in the reeds (Tindale 1930-52:

42).

European twine, hooks and lines were also

handed out at the Point McLeay Mission (Taplin

Journals: 18 September 1862). Fishing techniques

used in the Lower Murray region included netting,

spear and club fishing, trapping and opportunistic

collecting.

NET-FISHING

The Ngarrindjeri people referred to the act of

netting fish as ngerin (Taplin 1879: 130). Marine

net fishing often involved the coordination of a

large number of people. Worsnop provided a vivid

account of Aboriginal people sea fishing in the

19th century. He records:

In Encounter Bay I have seen the natives fishing

almost daily. Two parties of them, each provided

with a large net, square in form, with a stick at

either end, and rolled up, swam out a certain

distance from the shore, and then spread

themselves out into a semicircle. Every man

would then give one of the sticks round which

his piece of net was rolled to his right hand man,

receiving another from his left hand neighbour,

bringing the two nets together, thus making a

great seine. They now swam in towards the

shore, followed by others of their number, who

were engaged in splashing the water and

throwing stones, frightened the fish and

prevented their escape from the nets (Worsnop

1897: 90-91).

George French Angas painted two men fishing

with a seine net at Second Valley, south of

Adelaide, in 1844 (Angas 1847a: Plate XXI).

Cawthorne provides a similar scene, in

watercolour, at Second Valley, dated c.1842

(figure in Hemming & Jones 2000: 9), as does

Snell in his sketches, dated 14 July 1850, at Yorke

Peninsula (Snell 1849-59 [1988: 128]). Eylmann

(1908: 375 & Plate XXXIV fig. 3) published a

drawing of two Ngarrindjeri men using a long

seine net, with a third man helping to drive the

fish in. It is likely that several nets were

sometimes connected together with supporting

sticks.'* In situations where nets were not

available, branches could be used to drive the fish

up on the beach (Angas 1847b: 112). A former

colonist wrote to Tindale stating that in the Port

Germein area, north of Adelaide:

On occasions a wall of prickly bushes was built

in about three feet [one metre] of water and

rolled shorewards, the ends converging to an arc.

The fish which were impaled or caught were

extracted by the pushers and thrown to others

following behind (Parkes 1936, cited in Tindale

1935-60: 48).

The ‘wall’ of branches, comprised of saltbush

and mangrove, was called winda and required two

people to roll it and two more for picking out the

fish (Parkes 1936, cited in Tindale 1935-60: 96).

Net fishing in fresh water required fewer people

than sea fishing, with small nets used to catch fish

living amongst reeds and logs. Nets were essential

items of Lower Murray material culture for fishing

in the lagoons of the Lower Lakes, Coorong,

Murray River and in the creeks of the southern

Fleurieu Peninsula. In the Encounter Bay area,

Aboriginal women also caught tadpoles from

claypans with fine meshed nets and cooked them

in large ‘mutton-fish’ shells (Worsnop 1897: 83).

There was some degree of specialisation in the

nets used by Lower Murray people. Harvey (1943:

111) described three main types of nets used by

Yaraldi-speaking people of the Lower lakes. Nets

with a small mesh, jatrumi [pronounced

‘yatrumi’], were used when fishing for kanmuri

(mullet, Aldrichetta forsteri). The nets with a

larger mesh, neri, were used for tukkeri (bony

bream, Nematalosa erebi). The big drum nets,

dongari, were mainly for catching pondi (Murray

cod, Maccullochella peeli) found under large logs

"In 1932 Tindale (1953: 42) recorded from a European named Arthur White that Aboriginal people living along the river had used a double-pointed

wooden fishhook, Tindale (no date, ‘Milerum’, Stage A, #3, Archives, South Australian Museum, Adelaide) also claimed that they used a fishing

toggle made from bone as a form of fishhook.

‘* This technique was also used in southern Yorke Peninsula (Tindale 1936: 57).

[52

submerged in the river.’ The drum nets were

made with sticks in the form of a rectangular

pouch (Harvey 1943: 111).

The Tangani people of the Coorong used

specialised nets, such as the jataruma

[pronounced ‘yataruma’] to catch mullet and the

donggari to catch mulloway (Argyrosomus

hololepidotus) and other big fish (Tindale 1934—

37: 226). They also used the ngeiri net, which

was made from the same sedges that were

collected for basket and mat making. All nets used

by Tangani people in the pre-European period

were for hauling (Tindale 1934-37: 226). The

Tangani used a hooped fishing net, termed

kandarangku, with a mesh of 2.5 cm, for larger

fish (Tindale 1951: 258). Apparently the term is a

play on words meaning ‘widow catcher’, as

kandari is applied to coarse rope and rangku

refers to a widow.'® Murray River people around

Moorundie used the same type of net,

kanderunku, to catch ponku (Murray cod).'7 The

Kingston people used a net called pinang kanji to

round up fish in shallow waters (Tindale 1931-

34: 89). Angas provided a small watercolour

painting of a pouch-like net ‘for taking very small

fish from Lake Bonney’ (Angas 1847a: Plate LI)

in the South East of South Australia.

Aboriginal people in southern South Australia

probably did not use set nets until well after

European settlement. Tindale’s main Lower

Murray informant, Clarence Long (Milerum),

claimed that set nets were rarely used in the

Coorong when he was younger, and not at all

before then (Tindale 1934-37: 226). This suggests

that Europeans had introduced the practice to the

local Aboriginal people. From available accounts,

sinkers and floats were not used in association

with net fishing during the pre-European period

of southern South Australia. The record of Murray

River people using clay from wasp nests to make

sinkers for set nets is probably a post-European

innovation.'* Tindale (1935-60: 17) suggested that

vegetation placed along the top of the net to

prevent fish from jumping out of the water might

have helped keep the net vertical.

It is likely that in the pre-European period most

PA CLARKE

fishing nets in southern South Australia and

Victoria were fabricated from two-ply string made

from fibre obtained from the roots and stems of

the bulrush (Typha species) (Beveridge 1883: 42;

Krefft 1862-65: 361; Tindale 1935-60: 17; Zola

& Gott 1992: 8-9, 62). Europeans also know this

plant as flag, cumbungi and reed mace, whilst

contemporary Ngarrindjeri people refer to it as

manangkeri or manakeri (Clarke 2001b). It was a

major food source, with the chewed remains of

the roots being generally in good supply in

Aboriginal camps (Clarke 1988: 69-70, 72;

Tindale & Mountford 1936: 497). Tindale stated

that:

. mature rhizomes were chewed and fibres

made into string, the chewing for this purpose

being a duty of women. Older women’s teeth

were often worn to the gum line by their

constant chewings.!?

He also suggested that the proximity of bulrush

grounds would have made a good riverside camp.

In 1894 an Aboriginal man from Point McLeay,

Jacob Harris, provided a detailed written account

of net making from ‘rushes’, which were probably

bulrushes. He said that before Europeans arrived,

Aboriginal people in the Lower Murray region

used to:

make our nets almost the same as the Europeans

did, the meshes were the same, the only

difference being that yours [Europeans] were

made of twine etc., while ours from rushes. The

rushes had to be steamed or cooked or whatever

you like to call it, first a hole was dug, say a

couple of ft. [ = 60 cm], then a fire was placed

in it, after it had burned almost to ashes some

damp or wet grass was put on the coals, then the

rushes, so that the steam arising from the grass

would steam or cook ... put in about evening and

left until morning, it was then taken out and

divided among the camp to chew or suck the

juice out of it, which is very sweet, it was then

made or twisted into a line for to be made into a

net. ... the meshes were the same [as European

nets].?°

Angas stated that the marine fishing nets were

‘composed of chewed fibres of reeds, rolled upon

1934.

Descendants from the Yaraldi and Maraura people from the Lower Lakes and Murray River regions made S.A. Museum drum net, A21338, in

16 Tindale (no date, ‘Milerum’, Stage A, #3, Archives, South Australian Museum, Adelaide).

"Tindale, ‘Ngaiawung vocabulary’, Archives, South Australian Museum, Adelaide. Also see Scott (cited Tindale 1951: 258).

18 Tindale (no date, ‘Milerum’, Stage A, #3, Archives, South Australian Museum, Adelaide).

9 Tindale (no date, ‘Milerum’, Stage A, #3, Archives, South Australian Museum, Adelaide).

2 J. Harris letters (D6510(1)14,15, Mortlock Library, Adelaide). Tindale (1934-37: 164) received a similar description of the process from his

Ramindjeri informant, Reuben Walker.

EARLY ABORIGINAL FISHING TECHNOLOGY

the thigh, and twisted into cord for the purpose’

(1847a: Plate XXI).?' Emu oil was sometimes

rubbed into the fibre while it was being made into

two-stranded cord (Tindale 1931-34: 60). Fibre

treated in this manner makes the string more

supple, while maintaining its strength and

durability. Tindale (1951: 257) maintained that

Aboriginal cordage from all areas of Australia was

always composed of single or two-ply twists,

never three or more. Nevertheless, there some

were exceptions. For instance, West (1999: 20,

23-28) describes the making of three-ply rope in

Arnhem Land and possibly Tasmania.

After European settlement, changes in the

Aboriginal diet would have led to a significant

decrease in the availability of fibre from bulrush

roots. Fibre used by Lower Murray people to make

string for nets also came from an unidentified

plant known in the local Aboriginal language as

calgoonowrie (MacPherson, no date: 10). A plant

recorded as used for making Coorong nets was

kuka (native flax, Linum marginale).”* In 1932 an

informant from the Maraura people of the Upper

Murray of South Australia, Frank Fletcher, said

that gill nets and drag nets were both made from

the pungur rush (Tindale 1953: 23).73 Harvey

(1943: 109-110, 112) stated that the stems of

jalkeri (knobby club rush, Isolepis nodosa) were

used for making nets.“ But field investigations

have indicated that the stems of this rush are pithy

and not suitable for making string or baskets. The

stems of mangatu (spiny sedge, Cyperus

gymnocaulos) would have been better; this

species is still used by local Aboriginal people

who make baskets and mats from it.** Nets made

from sedge stems were also made for dry uses,

such as catching ducks.”°

The technique used to make nets in southern

South Australia has been variously described as

153

‘knotted netting’ or ‘knotted looping’ (Davidson

1933: 257-259, 269-272, figs 1, 9-10; West

1999: 30-33, 49). The Tangani people called the

mesh of the net, mandar, which reputedly meant

‘the eyes’; the netting knot or tie was mulbakan

and the act of making a net was lagulun (Tindale

1934-37: 226). The last term is a reference to the

threading motion of making the knot being like

‘spearing’.*’ In the Ramindjeri language of

Encounter Bay, /akk-in was recorded as ‘spearing;

weaving (as rushes for a basket)’ (Meyer 1843:

74). The mesh sizes for nets in southern South

Australia were variable, in the range 1—2.5 cm

with the Kongarati Cave fragments, 2.5 cm

recorded for the Coorong hoop net and 8 cm for

netting wrapped around a desiccated body found

at Fromm Landing, Murray River.”

SPEAR AND CLUB FISHING

During the day, men caught fish such as

malawe (mulloway, Argyrosomus hololepidotus)

by standing motionless in the river or in their

canoes (Hemming & Jones 2000: 9; Meyer 1846

[1879: 192-193]; Smith 1930: 230-231; Snell

1849-59 (1988: 182]; Tindale 1934-37: 7).

Fishers attracted the fish by their shadows and

stabbed them with hand-held spears. Large spears

made from Callitris pine with two to three

wooden prongs lashed to the head were used for

spearing mulloway and Murray cod.” Being large,

these spears were also used as punting poles when

fishers were travelling on bark canoes and rafts

(Hemming & Jones 2000: 9). It is likely that these

spears were hidden near the area of their use when

the fishing season was over. In the Lower Lakes

area fishermen frightened fish from their hiding

places by a large noise, created by thrusting one

2! A supporting reference is Angas (1847b: Plate LVI). An example of a net made from Typha fibre is A2000, collected from the Lower Lakes district.

2 Tindale (no date, ‘Milerum’, Stage A, #3, Archives, South Australian Museum, Adelaide).

The term pungur may relate to the punggurtko specimen of Juncus species collected by Tindale from Swan Reach, 5 August 1964 (A68584, South

Australian Museum). If so, then the plant referred to for net-making was probably of the same species. Similarly, a related term, puungort, is a

Westem Victorian Aboriginal term for the basket sedge, Carex tereticaulis (Robson 1986).

This plant was formerly known as Scirpus nodosus,

Examples of River Murray fishing nets made from sedges (Cyperus species) in the South Australian Museum include specimens A17529, A21338,

A26250, A45090, The term mangatu was recorded from Aboriginal sources in the 1980s (Clarke 2001b).

7 See figure in Hemming & Jones (2000: 13). Satterthwait (1986: 39-40) provided an overview of the use of nets for catching waterfowl in

Aboriginal Australia.

In the Ngarrindjeri language of the Lower Lakes, /akelun was recorded to mean ‘spearing’ (Yallop & Grimwade 1975: 33-34, 85-86).

Tindale (1951: 258) reported on specimens A65091, A25351 and A20616, respectively. Smyth (1878; 389-390) discussed mesh sizes for Victorian

nets.

In 1936 Tindale collected a 3.1 m-long fishing spear (Museum specimen A26093) made of pine from the Coorong, This specimen has two prongs

lashed to the head and was called punkularipuri ma:wantj in the Tangani language. In 1844 G. French Angas painted a close-up of a large spear

with three prongs lashed on (Angas collection, Archives, South Australian Museum, Adelaide — see Angas 1847b: Plate XXX). This watercolour

is reproduced in Hemming & Jones (2000: 9).

154

of these pronged spears into the water (Unaipon

1924—25 [2001: 19]). The compressed air caught

between the prongs rose to the surface with a loud

report. There is an account of spear-fishing

competitions held from November to April among

the Piltindjeri group of Yaraldi-speaking people

living along the southeastern shore of Lake

Alexandrina, with the fish caught being presented

to senior people in the community (Smith 1930:

231-236; Unaipon 1924-25 [2001: 19-24]). In

this instance, it required skill in fish tracking,

involving the detection of movements in

pondweed, reeds and water ripples.

At night, a fire was used to attract fish to be

struck by spear or club (Angas 1847b: 112).

Angas painted a club called a wadna, which was

‘used by the Port Lincoln natives, to throw at fish

whilst swimming’ (Angas 1847a: Plate LI).

Similar bent elongated clubs from the Lower

Murray are also in the Museum collection. Clubs

like these and described as boomerangs were

reported as used along the Coorong (Olsen 1991:

5). Sometimes a bark canoe was used as a

platform to fish from, with a fire contained by a

clay hearth in the middle that also served to cook

the catch (Angas 1847b: 54, 101, 107; Meyer

1846 [1879: 193]).*%° Mobility across water was

important for the fisherman in order to reach areas

favoured by particular species of fish. Therefore,

rafts made from reed stems (Phragmites australis)

and grasstree flower-stalks (Xanthorrhoea

species) would also have been important items of

fishing material culture in the Lower Lakes and

along the river lagoons.*' Although relatively

common along the Murray River and in the Lower

Lakes, bark canoes were rarely seen in the

Coorong lagoon, and even then only for bringing

in trade items.** Red gum trees (Eucalyptus

camaldulensis) required for bark are found only

in permanent freshwater regions, such as along

creeks in the southern Fleurieu Peninsula and

along the Murray River. There is no evidence that

any form of watercraft was ever used for sea

fishing, although short trips were made on reed

rafts or floats to nearby rocky islands in order to

kill seals there (Tindale 1941: 241).

PA CLARKE

OPPORTUNISTIC HARVESTING

There are documented cases of Aboriginal use

of fish poisons in small lagoons of the upstream

reaches of the Murray River bordering northern

Victoria (Curr 1883: 110). Nevertheless, Taplin

doubted that Ngarrindjeri people possessed

knowledge of the use of poisonous plants (Taplin

1879: 47). When large numbers of fish died for

natural reasons, such as changes in salinity in the

river or lake, Aboriginal people quickly gathered

them (Eyre 1845, 2: 266). Before the construction

of the barrages across the Murray Mouth

separating the Coorong from Lake Alexandrina,

elderly Aboriginal informants interviewed in the

1980s claimed that there were sometimes rapid

changes in the water from fresh to saline (Clarke

2002). This tended to kill a large number of fish,

providing an abundant but temporary source of

food. At other times, the incoming salt water

drove certain species, such as Murray cod,

upstream and suspended fishing in the lakes until

it receded (Olsen 1991: 8; Taplin Journals:

19 May 1860). Bony bream also die off seasonally

in large numbers and, according to Ngarrindjeri

informants in the 1980s, they had provided a

temporary windfall of food (Clarke 2002).

FisH-TRAPS AND ENCLOSURES

Fish-traps and enclosures ranged from slight

modifications to natural features of the landscape

to special purpose-built structures (Mulvaney &

Kamminga 1999: 34-35). For example, in some

areas Aboriginal people were able to use the

narrowing of the channels between swamps as

traps in which to place netted bags. In his diary

Wells records:

Just now [July] there is fish to be had out of the

swamps, and we got one of the native females to

make us a net. They are made of a long kind of

soapy platt [sic.], but flat and round, quite as

large as an umbrella top. It is then doubled not

unlike a huge ankle boot, and at the two ends

there is a small opening. It is then raised from

the bottom in such a way that the fish when

*° The South Australian Museum has a bark canoe (A6443) from Avoca Station on the Darling River that contains an early 20th century mock-up

made by museum artisans of the fireplace for display. The original fireplace was made in a mud-lined wooden container (A53554).

*! Jacob Harris, an Aboriginal man at Point McLeay writing in 1894, claimed that the use of the ‘rude kind of raft’ made from reeds predated the use

of bark canoes in the Lower Murray (J. Harris letters, D6510(L) 14,15, Mortlock Library, Adelaide). The South Australian Museum holds raft

specimens (A14632, A14633) made by a Yaraldi woman, Amy Johnson, in 1930.

* This statement derived from accounts by Meintangk woman, Ethel Watson, and Tangani man, Clarence Long (Milerum) (Tindale 1931-34: 69, 87).

EARLY ABORIGINAL FISHING TECHNOLOGY 155

caught cannot return. The net is then fastened

with sticks in a narrow channel where one

swamp runs into another and all fish going with

the stream are caught. These fish the blacks call

Coogolthee. The nets are laid overnight and by

break of day next morning we have plenty

brought us for breakfast.”

On the frontier of British colonisation, wild

foods and Aboriginal labour were often crucial to

the survival of European settlers (Clarke 1996).

Examples of more active manipulation of the

environment were the construction of long

trenches by Aboriginal people to concentrate fish,

in the long term saving them much labour

expenditure. For instance, Smith records that in

the Murray region:

In the low-lying country, alongside of the river,

trenches are dug two or three hundred yards [180

or 270 metres] long and from four to five feet

[1.2 to 1.6 metres] deep. When the Murray

becomes flooded it overflows its banks to the

extent of a mile or more [1.6 kilometres or more]

on each side, and frequently the Murray cod, the

bream, the butterfish, and other fish are living in

this water. When the waters become low through

evaporation and soakage the fish are easily

caught ... They wade into the shallow water and

scoop the fish into baskets made especially for

this purpose (Smith 1930: 229).

These channels were often relatively large

modifications of the landscape that would have

had a significant impact upon the local watertable

(Lourandos 1997: 219-22),

Another modification of the landscape for

fishing was the building of stone and wooden

fish-traps. Some fish-traps, probably the

predominantly wooden ones, were known by the

Ngarrindjeri name, ku:yitaypari.** The Yaraldi

people in the Lower Lakes did not make such

structures, preferring to use fishing spears instead

(Tindale 1931-34: 73-75). In the Coorong fish

were harder to spear due to the prolific

waterweed, so the Tangani relied heavily upon

stone fish-traps, talaipar (Tindale 1931-34: 73—

75). Areas of shallow water in the Coorong, such

as at the Yungurumbar crossing place, were

considered good places for making such stone

fish-traps (Tindale 1931-34: 71).** The Tangani

generally placed their fish-traps made from

limestone blocks along the landward shore of the

Coorong (Tindale 1974: 61-62). This was

presumably because the water here is shallow and

covers a limestone shelf, in contrast to the

seaward side of the Coorong lagoon, which is

deep and sandy. Tindale provided a detailed

record of the construction and use of fish-traps or,

as his informants called them, ‘fishing-stations’

(Tindale 1931-34: 73-75, 1934-37: 5). The fish-

traps were generally made in the shape of a V,

with one of the walls, termed nganangkure or

ngalde, connecting with the bank. The wall that

extended into the Coorong lagoon was sometimes

up to 30 metres long. The basketware trap was

placed at a gap in the base of the V. The species

of fish seasonally caught in the traps included

kongoldi (probably congolli), njindumi (an eel

species), palengoi (unknown species), lapalap and

therugarai (unknown fishes, not found in the

Lower Lakes) and teri pateri (‘sand mullet’)

(Tindale 1931-34: 75, 1934-37: 5).*° No attempt

was made to drive the fish in, relying solely on

their natural movements.

At Noonamena, on the mainland side of the

Coorong near Meningie, the tops of silted over

fish-trap formations can still be seen. Aboriginal

people interviewed by the present author in the

early 1980s said that these fish-traps were

naturally formed stone structures that were

previously modified and maintained with stones

and pieces of wood. According to Ngarrindjeri

man Jack Koolmatrie, mullet travel northwards up

the Coorong during the day (Ely 1980). Upon

reaching the trap, fish would swim into the wide

mouth of a horseshoe-shaped line of rocks. The

foundation of this was a natural reef of exposed

rock with all gaps except one narrow exit shored

up with logs and boulders. The mullet were forced

to travel through this narrow exit where a net or

basketry container put in their path would catch

them.

Elsewhere in southern Australia, some of the

fish-traps have been described in the literature as

% Wells, 16 July 1853, The species of fish referred to here is probably kungali (congolli, Pseudaphritis urvilli) (Clarke 2001b).

* Yallop & Grimwade (1975: 55) list ku:yiti as ‘rushes, sticks’, and ku:yitaypari as ‘fish-trap, barrier of sticks’.

** The Yungurumbar crossing place is probably that between Rabbit Island and Junggurungbar (pronounced Yunggurungbar), a hill on the

Younghusband Peninsula (see Tindale 1938, fig.1).

The eel species referred to here is probably Anguilla australis. The palengoi was said by Tindale to be equivalent to the Yaraldi pelingi. This is

probably the pelanggi that Berndt & Berndt (1993: 308) described as ‘mudfish’ or ‘butterfish’. The Japalap may possibly have been a species of

Galaxias, In the case of teri pateri or ‘sand mullet’, Tindale (1931-34: 74) said that its Yaraldi name is weialapi. This is probably the same word as

the ‘welappi’ or ‘mullet’ recorded by Taplin (1879: 131). Eckert & Robinson (1990: 19) consider that this refers to the ‘jumping mullet’ (Liza

argentea).

156

weirs.*” According to Edward J. Eyre, Aboriginal

people seasonally gathered at the channels around

Moorundie that connected the river flats with the

Murray River:

making a weir across them with stakes and grass

interwoven, [would] leave only one or two small

openings for the stream to pass through. To these

they attach bag nets, which receive all the fish

that attempt to re-enter the river. The number

procured in this way in a few hours is incredible.

Large bodies of natives depend upon these weirs

for their sole subsistence, for some time after the

waters have commenced to recede (Eyre 1845,

2: 253).

This practice occurred during early December

when the Murray River floods had already reached

their highest point and were beginning to recede.

Further south, in the shallow waters and swamps

of Lake Frome near Burr Range, small fish were

caught in weirs (Angas 1847b: 174). Also in the

South East, near Rivoli Bay, Angas recorded:

On some of the swamps the natives had built

weirs of mud, like a dam wall, extending across

from side to side, for the purpose of taking the

very small mucilaginous fishes that abound in

the water when these swamps are flooded

(Angas 1847b: 155).

In the pre-European period, Aboriginal

modification of the riverine landscape for fishing

must have been considerable. It was oral history

among old river boat captains in the 1980s that

before the lock system was introduced, remains of

Aboriginal built fish-traps were known hazards to

paddle-steamers when the water level was low (T.

Sim, pers. comm.). Hahn recorded that in the

Hahndorf area of the Mount Lofty Ranges, the

Aboriginal people would:

build a dam into the river, high enough to let

about a foot [31 centimetres] of water stream

over it. Because of this dam, the fish in their run

must come close to the surface of the water,

where the savages stand in readiness to spear

them (Hahn 1838-39 [1964: 133]).

PA CLARKE

In coastal zones weirs of brushwood

constructed at mouths of creeks caught fish left by

receding tides (Angas 1847b: 112). Some of the

trenches, traps and weirs were designed to catch

bait for much larger fish. For example, near

Martin Well on the Coorong, drains 100m in

length were constructed by Aboriginal people to

catch small fish, called lap-lap, for bait (Worsnop

1897: 106).** These were netted in fine close mesh

nets. These weir and trench constructions appear

similar in design to the much larger earthworks at

Lake Condah in western Victoria.”

Outside the Lower Murray, but elsewhere in

southern South Australia, fish-traps have been

recorded in the Port Lincoln estuarine area

(Martin 1988; Mountford 1939), as well as at the

mouth of Pedlar Creek south of Adelaide

(Stapleton & Mountford, no date). In the case of

the latter, the Tangani man, Milerum, knew of

these traps as he and his family camped there,

when he was a child, on their way to Adelaide via

the coast.“ There were also fish-traps made from

large water-worn boulders at Moana Cove, also

south of Adelaide, but these were in poor

condition when Tindale inspected them in the

1920s.*' Some traps could be dragged out of the

water when fish were not required. For instance,

Charles Sturt stated that on his 1830 expedition

down the Murray River he:

observed some cradles, or wicker frames, placed

below high water-mark, that were each guarded

by two natives, who threatened us violently as

we approached. In running along the land, the

stench from them plainly indicated what they

were which these poor creatures were so

anxiously watching (Sturt 1833, 2: 165).

Small dams or ‘pounds’ might not only be

structures for catching fish, but were also used to

keep part of the catch alive for future use. At the

Point McLeay Aboriginal Mission, ‘fish pounds’

made of stakes were in common use by Aboriginal

people during George Taplin’s period there

(1859-79). According to Lower Murray man

7 For example, Beveridge (1883: 48); Curr (1883: 110) and Pretty et al (1983: 119). Tindale listed ‘fish weir (set in creek)’ as ake in a Murray River

dialect (Tindale, ‘Ngaiawung vocabulary’, Archives, South Australian Museum, Adelaide).

This fish, lap-lap, was possibly a species of Galaxias. Lapps Lapps was recorded to mean ‘small fish’ in the Booandik language (Smith 1880: 3).

Wells (1852-55) listed lap-lap as ‘very small fish’. Tindale (1931-34: 74) said that lapalap were a Coorong fish species not known in the Lower

Lakes.

* For a description of the Lake Condah eel-traps, see Coutts et al (1978); Flood (1995: 240-245; 1999: 216-220); Hemming (1985); Robson (1986)

and Worsnop (1897: 104-106). The S.A. Museum has a basketry trap, A6431, collected from Lake Condah, c.1910 (see figure in Hemming &

Jones 2000: 9).

“Tindale (no date, ‘Milerum’, Stage A, #3, Archives, South Australian Museum, Adelaide).

‘| Tindale (no date, ‘Milerum’, Stage A, #1, Archives, South Australian Museum, Adelaide).

“2 The ‘fish pounds’ were recorded by Taplin on the shore of Lake Alexandrina (Journals, 11-12 October 1859; 8 November 1859; 11 November

1859; 28 November 1859; 10 January 1860; 21 November 1861; 20 March 1862).

EARLY ABORIGINAL FISHING TECHNOLOGY 157

Lindsay Wilson, interviewed by the present author

in the early 1990s, the practice of fish storage in

wooden enclosures in the lake at Point McLeay

continued until the 1930s. During the early

twentieth century along the Coorong, European

fishermen kept alive the bream they had caught by

putting them in ‘pounds’ made of tea-tree stakes,

until the weather was suitable for transporting

them to the market (Evans 1991: 40).

CULTURAL ASPECTS

Aboriginal people along the Murray River

considered that their Dreaming Ancestors created

the wetlands that provided them with abundant

sources of food. For instance, Natone, an

Aboriginal man from the South Australian section

of the Murray River, claimed that a blind woman,

Noreela, had created the environs of the river

(Bellchambers 1931: 112, 125). Starting from

Lake Victoria, Noreela, with two young children

to guide her, made the river by driving back the

sea. She travelled like a ‘drunken bee’; her

meandering course meant that the river was very

long. This lengthening of her journey was

considered greatly desirable, as it increased the

number of fishing and hunting grounds, with a

lagoon at each elbow. The fossils jutting out of

cliffs along the Murray River were said by

Aboriginal people to be the remains of fish killed

and eaten by Noreela and her children. Another

account, still told by Ngarrindjeri people in the

1980s, was of a Thukabi Ancestor, who was a

large river turtle that came down the Murray

River:

Thukabi [a large river turtle] came down [from

the Darling district] through the desert looking

for a place to lay its eggs. As she went, the drag

of its tail made the river, its flipper carved out

the lagoons and banks. You can see where it

went. When it got to the lake, it pushed itself

into the sea (H. Rankine cited Clarke 1994:

114).

In all these accounts, the present course of the

Murray River is explained by the actions of

Ancestral Beings.*?

In the Ngarrindjeri Dreaming of the Lower

Murray, Ngurunderi chased a large Murray cod,

Pondi, down the Murray River, widening its banks

in the process, until it was eventually speared near

the Murray Mouth (Berndt & Berndt 1993: 224;

Clarke 1995: 148-149; Hemming & Jones

2000).* Whilst Ngurunderi was at the Murray

River entrance into Lake Alexandrina, a water

spirit known as the Mulgyewonk tore holes in his

nets, which prevented him fishing for his family

(Tindale & Pretty 1980: 50). There are accounts

of Aboriginal legends from further upstream in

Murray-Darling Basin where the dreaded river

spirit who fed on boys was a large Murray cod

(Sinclair 2001: 120-121, 252). Ngurunderi made

the fishing grounds and lookouts for the Lower

Murray people, which is told in song (Tindale

1931-34: 259).

Fish species also appear elsewhere in Lower

Murray mythology. For instance, Yamakawi

(Shark Ancestor) had a prominent role in the

Kondoli (Whale Ancestor) Dreaming of

Encounter Bay, which explained the origin of fire

(Clarke 2001a: 24-25). There is also a Dreaming

myth relating predominantly to freshwater fishing

technology, which took place in the region from

Lake Hindmarsh in western Victoria to Lake

Alexandrina in the Lower Murray (Harvey

1939ms, 1943; Tindale 1934-37: 65-69). In this

account the drum-nets of the pelicans were

transformed into large pouches under their bills.

In the Dreaming of Limi (= Lime), the love of

eating fish is involved in explaining the creation

of the Inman and Hindmarsh Rivers of the

southern Fleurieu Peninsula (Berndt & Berndt

1993: 311; Meyer 1846 [1879: 202]).*° The

Skyworld landscape contains a fish too, with a

constellation of stars being Nunganari, the

Stingray (Berndt & Berndt 1993: 164, fig. 25;

Clarke 1997: 131).*°

Economically important fish species were

considered by the Lower Murray people to have

been created by Ancestors during the creation

period. For instance, in a Ramindjeri account of

the Kondole (Whale) Dreaming, the Kuratje and

Kanmari Ancestors became small fish when they

ran in the sea:

From the late 1980s, an animated version of this myth was given at the Signal Point Museum, Goolwa.

“Sinclair (2001; 120) speculated that the relatively large size that some cod grow to, up to 1.8 metres, enhanced Aboriginal beliefs of the beast’s

mythical qualities.

4S Berndt & Berndt (1993; 311) suggested that Limi was a Stingray or Carpet Shark Ancestor. Meyer (1843 part 2: 75) listed ‘Lime’ as a ‘kind of

seal’. Limi and his family were eventually transformed into a group of prominent rocks in the sea at Victor Harbor near the Bluff (Meyer 1846

[1879: 202]).

4° According to Tindale (no date, ‘Milerum’, Stage A, #1, Archives, South Australian Museum, Adelaide), the Tangani considered that there were

seven stars shaped like a stingray.

158

The latter was dressed in a good kangaroo skin,

and the former only a mat made of seaweed,

which is the reason, they say, that the kanmari

{mullet, Aldrichetta forsteri] contains a great

deal of oil under the skin, while the kuratje

(Western Australian salmon, Arripis truttaceus]

is dry and without fat (Meyer 1846 [1879:

203]).4”

In other Dreaming accounts, the Murray cod,

Pondi, was cut into smaller pieces by either

Ngurunderi or Waiyungari, depending on the myth

version, and thrown back into the water to become

different types of fish (Clarke 1995: 148; 1999b:

53-54).

The Ramindjeri version of the Ngurunderi myth

epic, recorded from Reuben Walker in 1935, starts

from a large lake near mountains somewhere to

the northeast of the Lower Murray and involves a

large fish which is not a Murray cod:

Ngurunderi came to the Lake and speared the

fish which, made off with it at a great speed and

cut a deep track right down to Lake Alexandrina.

Ngurunderi followed and noticed that the river

thus formed was without fish. So he stood on the

bank and broke the bark of the red gum tree

(wuri) up into shreds & threw it into the water

and said; you are Murray cod. This must be true

because, when you cut open a Murray cod you

can see a tree like mark of blood vessels on the

walls of its body; it is just like a gumtree.

Ngurunderi then threw in pujulanki bush (Native

wild grape? Cryptandra hispidula) and this

became catfish (Tandanus tandanus). He threw

in peppermint gum bark (tentumi) and it became

the giri (Terapon bidyanus Mitchell). After he

had made all the best fishes he threw in a piece

of refuse and it became the Bony bream (tukari,

Fluvialosa richardsoni (Caselman)).*®

In relation to the blood vessels, it is worth

noting that along the Murray European fishers

have an analogous belief which interprets the tree-

like markings on cod swim bladders as ‘skin

maps’ that show the birth place of the individual

fish (Sinclair 2001: 124-127). Tindale recorded

from a European settler the following Dreaming

account involving Matumeri who:

.. chased a large fish from the sea entrance of

the Murray Mouth up through the Lake until he

caught it out off Poltallock [Poltaloch Station]

where he killed it and pulled it to pieces and

threw the pieces all about and they turned into

small fishes and that is how the different sorts of

PA CLARKE

fish came (RD Anderson 1934 [cited in Tindale

1934-37: 175]).

The Ancestor mentioned here is probably

Matamai, who was Ngurunderi’s son (Clarke

1995: 146). The theme of an Ancestor tearing up

large beings to make many smaller species also

occurs in relation to kangaroos according to one

account of the Waiyungari mythology (Clarke

1999b: 54).

Aboriginal people considered themselves as

having a role in the continuation and wellbeing of

their environmental resources. Ceremonies were

sometimes performed in order to increase fish

supply. For instance, Howitt relates:

There is a spot at Lake Victoria [ = Lake

Alexandrina], in the Narrinyeri [ = Ngarrindjeri]

country, where when the water is, at long

intervals, exceptionally low, it causes a tree-

stump to become visible. This is in charge of a

family, and it is the duty of one of the men to

anoint it with grease and red ochre. The reason

for this is that they believe that if it is not done

the lake would dry up and the supply of fish be

lessened. This duty is hereditary from father to

son (Howitt 1904: 399-400).

The illicit involvement between Waiyungari and

Nepeli’s wives in the Ngarrindjeri Dreaming was

perceived as the cause of poor fishing in early

spring each year. The arrival of the Young Men

(Orion) and the Women (Pleiades) constellations

in September was considered to help turn this

around (Berndt & Berndt 1993: 164; Clarke

1999b: 57). The flowering of certain plants may

also have been an indication of the arrival in the

Lower Murray of certain species of fish. This was

the case at Marion Bay in Yorke Peninsula, where

the prolific flowering of tea-trees was a sign to

the Narangga people that the mullet fish were

soon to come in large numbers.” Here, it was

claimed that initiation ceremonies were held then

to take advantage of this seasonally abundant food

source.

The abundance of fish would have allowed for

a larger Aboriginal population in the Murray

Basin in comparison to the surrounding regions.

The Lower Murray was particularly rich in fish

resources, involving marine, estuarine and

freshwater species (Eckert & Robinson 1990;

Evans 1991; Glover 1983; Olsen 1991; Sim et al

2000). Angas said ‘The Milmenduras subsist

“Identification of these fish names is given by Eckert & Robinson (1990: 19-20). Note that kuratye is equivalent to kuratje.

“Tindale (1930-52: 119). [Italics by the present author.]

“Reminiscences recorded by E. Davies in the Mail newspaper, Adelaide, on 25 March 1952.

EARLY ABORIGINAL FISHING TECHNOLOGY 159

chiefly on fish, and though extremely wild and

treacherous, present some of the best specimens

of the Aboriginal Inhabitants, as regards physical

appearance’ (Angas 1847a: Plate XI). Angas also

commented that ‘On the S. E. coast and along the

shores of the Murray and Lakes Alexandrina and

Albert the natives live chiefly upon fish, and

waterfowl’ (Angas 1847a: General Remarks).*°

Aboriginal people considered that some of the

spirits with whom they shared the landscape also

liked eating fish. For example, Lower Murray

people believed that the dreaded river spirit, the

Mulgyewonk, was attracted to the smell of fish

and once captured a young boy who was washing

fish oil from his hands on the edge of the lake

(Clarke 1999a: 157; Harvey 1939ms).

From the daily account available in Taplin’s

Journals, it is clear that fish, termed mame in

general, were a favourite food item in the Lower

Murray region. Ngarrindjeri people considered

that the dominance of fish in their diet set them

apart from at least some of their neighbours. They

gave cultural significance to the fact that when

their babies tried to speak, their first word was

mam (Taplin Journals: 10 October 1861). Adults

proudly interpreted this as the infant’s desire to

eat fish. The fish entrails, ngarakuni, were

considered good eating, being grilled over the

coals with the edges of the fillet curled up to catch

the juices.*' Special sticks, wunupi, were used as

fire tongs to remove food, such as fish, from hot

coals. Fish remains are often found in Aboriginal

middens on the banks of the Lower Lakes and

Murray River (Luebbers 1978, 1981, 1982; Pretty

et al 1983: 117-118; Tindale 1930-52: 67).

Some Aboriginal groups in southern South

Australia were noted by their neighbours for

having a diet dominated by fish. This is shown by

a recorded remark by Parnkalla people of

northeastern Eyre Peninsula that their Port Lincoln

neighbours, the Nauo, had ‘an offensive breath,

being fish eaters’ (Schiirmann 1844, 1: 7). In the

Lower Murray, Murray cod oil was rubbed on

initiates (Tindale 1930-52: 139). The Tangani

people of the Coorong had a song ‘ridiculing men

who refused to lend their fishing net’ (Tindale

1934-37: 267).

The material culture of the Lower Murray

people reflected their fishing background. Old

fishing nets were used in the Murray Basin for

wrapping human bodies that had already been

desiccated (Sheard et al 1927: 173; Tindale 1951:

258; Tindale & Mountford 1936: 495, 499). In the

Lower Murray, fishing nets were often among the

personal items placed in burial bundles lodged on

tree platforms (Hackett 1915: 29). Stingray tail

barbs or ‘nails’ were used in ngildjeri sorcery

(Berndt & Berndt 1993: 260). In the Murray River

area, Tindale recorded the medicinal use by

Nganguruku people of Eucalyptus leaves, tindunj,

with fish fat. Apparently they would ‘infuse

leaves in bark dish over hot ashes, mix with fish

fat (liver) for colds’.** Angas painted an object

that he described as ‘Kaikoonga — Bones of a fish

found in the Murray, worn as a head ornament, in

the same manner as the teeth of the kangaroo’

(Angas 1847a: Plate XXX). String-games or ‘cats-

cradles’ played by Lower Murray and South East

women and children often featured ‘fishing nets’

(Tindale 1931-34: 88).

The wide distribution of recorded modifications

to the landscape to assist in capturing or storing

fish in the Lower Murray and neighbouring

coastal and riverine areas indicates that their use,

in pre-European times, was a major subsistence

strategy. Lourandos has described some

earthworks by Aboriginal people in temperate

southern Australia as artificial drainage systems

operating to flush fish from swamps into channels

set with traps (Lourandos 1997: 219-221, 227).

This form of swamp management coped with

excess water during floods and helped retain water

in times of drought. This was part of a subsistence

pattern that allowed for a larger and semi-

sedentary human population, in comparison to

other regions. The material culture and diet of

Aboriginal people living in the south would

therefore have significantly differed from groups

to the north and particularly those situated some

distance away from major bodies of water. Fish

‘| Harvey (1939ms) and Tindale (1930-52; 248-249) sketched the manner in which fish were cut up and have given the Yaraldi names for the pieces.

% Tindale & Mountford (1936: 496), See photograph in Hemming & Jones (2000: 17).

*} N_B. Tindale specimens (A68579 — Eucalyptus oleosa,; A68585 — E. foecunda; A75835 — unknown Eucalyptus species) collected ‘3 miles north of

Swan Reach’, 5 August 1964, Clarke (1989: 3) has a similar record relating to mulloway liver.

“ In 1930 Tindale collected a string-game (A14962), called ‘fishing net’, from Amy Johnson (Yaraldi people, Lower Lakes). A string-game

(A66733), described as a Tangani ‘fish net game’ from the Coorong, was possibly a replica made by Tindale in 1934 (see Hemming & Jones 2000:

19). In 1930 DS. Davidson and N.B. Tindale collected at Swan Reach a string-game (A14958), called a ‘net’, made by Jerry Mason (Yiraruka

people, Murray River).

160

storage in pounds and mud pools was a pre-

European Aboriginal practice for managing the

windfall/drought situation of food gathering.

Another Lower Murray technique to extend the

use of fish food in pre-European times was to dry

fish on racks (Berndt & Berndt 1951: 29).

Because of the highly seasonal nature of fishing,

it is likely that some of the stone and wood

structures found in southeastern Australia,

identified by archaeologists as fish or eel-traps,

would be better described as fish pounds, being

for storage rather than capture.

Among the Lower Murray people, many of their

totemic familiars, the ngaitji, were species of fish

(Berndt & Berndt 1993: 306-312). This was also

the case for totemic groups along the Murray

River (Tindale 1953: 37, 49). In their daily life,

menstruating women in the Lower Lakes and

Murray River areas were forbidden to eat fish

(Berndt & Berndt 1993: 124-126, 141; Eyre 1845,

2: 295; Harvey 1939ms). These women were not

allowed to go near the water at all, as it was

considered that the success of men fishing would

be spoiled. On one occasion in the past, a large

incursion of seawater brought a considerable

number of poronti (sea mullet, Mugil cephalus)

into the Coorong lagoon (Tindale 1931-34: 119).

In spite of their abundance, amongst the Tangani

people only old men were allowed to eat them.

The mulloway would swim into the Coorong

lagoon to spawn under the limestone cliffs

(Tindale 1931-34: 119). In this condition only old

men were allowed to spear them. Similarly, if

young men ate pelenggeri fish (unknown species),

they would prematurely become baldheaded and

grey (Tindale 1934-37: 39).

In the Lower Lakes no Yaraldi women, except

the elderly, were allowed to eat catfish (Harvey

1939ms). Similarly, young Yaraldi girls were not

allowed to eat big-bodied kai:kuanggi

(‘freshwater bream’ — possibly black bream,

Acanthopagrus butcheri). Women during

menstruation were not allowed to go near water or

to eat fish caught with nets, as the Yaraldi

believed it would drive the fish away. In recorded

versions of the Ngurunderi Dreaming, his fleeing

wives cooked and ate tukkeri (bony bream) fish.*

The importance of this particular episode is

explained in various ways, depending on the

particular account. These are that the wives were

PA CLARKE

breaking a food prohibition by eating this fish

species; that the fat exploded in the fish, making

them sacred to Ngurunderi; and that the wives

incurred Ngurunderi’s wrath by giving him the

smaller of the two fish they had cooked. The

unifying theme in all versions is that this fish was

not to be eaten by women.

In the Murray River region more restrictions

were placed upon females until past the age of

child bearing than upon males of the same age

(Eyre 1845, 2: 293). Teichelmann stated that in

southern South Australia, food prohibitions were

such that women with children were prohibited

from eating certain food and that they generally

lived upon vegetables (Teichelmann 1841: 7). The

categories of people with least amount of food

prohibitions appear to have been infants and the

elderly. Eyre said that in the Murray River area

‘No restrictions are placed upon very small

children of either sex, a portion being given to

them of whatever food their parents may have.

About nine or ten years appears to be the age at

which limitations commence’ (Eyre 1845, 2: 293-—

295). He also stated that old men and women were

able to eat most things. All prohibitions would

have been based on cultural logic, even if the

reasons were obscured from those practising the

ritual. People generally excluded from

prohibitions were those not sexually or

economically active. It was the power to produce,

either physically or spiritually, at particular life

stages that was perceived as making people

sensitive to influences potentially harmful to

group harmony. In a sense, an individual’s

position in the society could be defined by what

the person could eat and what economic activities

they could engage in. The major categories

determining prohibitions appear to have been age,

gender and initiation status.

Aboriginal people could, at certain times,

exercise a degree of choice in the food they lived

on. For instance, some Lower Murray animal

foods such as fish, emu and kangaroo meat were

highly favoured foods when available. Yet

vegetable foods such as roots were probably the

mainstay when meat was not easily obtainable

(Clarke 1988: 73-74). A report from the

Adelaide-based Statistical Society in 1842

illustrates the seasonality of Aboriginal food in

the southern areas.°° The report notes that in

55 Accounts summarised from Berndt & Berndt (1993: 224, 435), Clarke (1995: 149, 1999b: 54) and Tindale (1934-37: 285).

s6 ‘Transactions of the Statistical Society. Report on the Aborigines of South Australia.’ Register newspaper, 8 January 1842. Thomson (1939) also

considered the seasonal aspects of Aboriginal culture.

EARLY ABORIGINAL FISHING TECHNOLOGY 161

spring mainly vegetables and grubs were eaten.

With the commencement of summer, fish were

obtained, as were kangaroos, emus, lizards and

the eggs and young of birds. During the hottest

part of the year possums and acacia gum were

procured, while in autumn berries and nectar were

available. In the winter a variety of roots were

consumed, as were possums and other animals.

The coastal zones of southern South Australia

were rich in natural resources, particularly food

such as fish, molluscs and coastal berries. Meat

from occasional whale strandings was also an

attractive coastal food source (Clarke 2001a).

Although many of these foods were available for

the greater part of the year, the onset of winter

made the coast a harsh zone in which to live.

Partly for this reason, ‘salt water’ Aboriginal

groups in southern South Australia would have

moved according to season between inland and

the coast. Stable isotope analysis of human bone

indicates that ‘salt water’ groups did not penetrate

up the Murray River beyond the boundaries of the

Lower Murray cultural region (Pate 1997, 1998,

2000). The pattern in the Adelaide region was a

general movement away from the coast in late

autumn, so that more substantial shelters could be

built in the protected Mount Lofty Ranges

foothills (Clarke 1991: 58-59; Ellis 1976: 116~

117; Ross 1984: 5; Tindale 1974: 60-61). The

historical record shows similar early patterns for

coastal groups from the Lower South East of

South Australia (Foster 1983: 23-43). Seasonal

movements, although an aspect of the Aboriginal

relationship to the physical environment, are

essentially dictated by the ‘cultural landscape’.

A seasonal population movement occurred

among at least some Lower Murray Aboriginal

groups. In winter the ‘salt water’ Tangani people

camped along the mainland side of the Coorong

lagoon, where firewood was plentiful and shelter

from weather available.*’ The fish-traps

maintained there, where the water is shallow,

provided a reliable source of food. During

summer these Tangani people camped on the

Younghusband Peninsula between the Coorong

and the Southern Ocean, giving them easy access

to coastal foods such as marine fish and berries.

There were also political reasons for the

movements, with the actions of neighbours

impacting on Lower Murray groups. For example,

one of the disadvantages of camping on the

mainland side of the Coorong was that here the

37 Tindale (1938: 21, 1974: 61-62).

Tangani were open to attack from the Ngarkat

people, who normally ranged in mallee areas to

the east of the Lower Murray. During harsh

summers the Ngarkat people were forced towards

the Murray River and Lower Lakes when their

water supplies dwindled, but the Tangani

considered that they were not likely to be attacked

during the winter. The locations of most Lower

Murray seasonal camps are not known, although

more archaeological research may improve our

present knowledge. For ‘fresh water’ Lower

Murray people, such as the Yaraldi-speaking

groups living along the edge of Lake Albert and

Lake Alexandrina, the yearly movement was

probably from the lakeshore to nearby inland

areas, in order to maximise food supplies and

comfort (see Fig. 2). During warmer months

lakeside camps were cooler as well as being close

to freshwater food sources. In contrast, during

winter, campsites in close proximity to large

bodies of water were more exposed to cold

weather. Back from the lakeside, the forests

provided natural windbreaks and had more

firewood and hut building materials available. The

prominence of aquatic technology used by the

Lower Murray people would have given them

little interest in the remote and harsh inland

regions.

CONCLUSION

Resource usage by early Lower Murray

Aboriginal people was a function of the broader

environmental and regional patterning of the

landscape. Aboriginal people in the Lower Murray

were not randomly dispersed over the landscape;

Ngarrindjeri people were restricted to the riparian/

marine areas that were consistent with their

material culture. They were, among other things,

regionally organised according to their perception

and use of the natural resources. The Lower

Murray people considered their relationship with

the environment to be an active one. Not only did

they physically manipulate their resources, they

also considered themselves to be influenced and

organised by the environment. The material

culture of the Lower Murray people, although

having some aspects in common with water-based

subsistence cultures in the South East and the

Murray River regions, was distinctive. They were

largely a cultural group confined to the riparian/

162 PA CLARKE

alll)

pa — FOREST SHRUBLAND

WOODLAND RY — GRASSLAND

E= | - open scrus MM ~ SUCCESSION

Kingston \od} ACG Y

Coastal Groups Inland Groups

Winter camps Winter camps

Summer camps Summer camps

COASTAL WOODLAND

SUCCESSION

SOUTHERN COORONG LAKE MENINGIE

OCEAN ALBERT

FIGURE 2. Pre-European vegetation of southern South Australia (after Boomsma & Lewis 1980: map) and

Aboriginal camping zones in the Lower Murray.

EARLY ABORIGINAL FISHING TECHNOLOGY

coastal habitats of the southern coastal region of

South Australia. Their hunting and gathering

practices help to define this cultural region.

Approaches in cultural geography, focusing on the

cultural construction and perception of the

landscape, are well suited to the study of

Aboriginal environmental knowledge.

163

ACKNOWLEDGMENTS

A draft of this paper appeared in the author’s PhD

thesis, which was supervised by C Anderson, P Smailes

and K Garbett. T Sim checked the scientific names of

the fish species.

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FOSSIL LIZARDS FROM THE PLIOCENE CHINCHILLA LOCAL FAUNA,

QUEENSLAND, WITH DESCRIPTION OF A NEW SPECIES

M. N. HUTCHINSON & B.S. MACKNESS

Summary

The lizard fauna of the Middle Pliocene Chinchilla Local Fauna consists of members of the

Gekkonidae, Agamidae, Varanidae and Scincidae. A new species of the scincid lizard genus Tiliqua

is described, based on a complete right dentary. The new species combines relatively unspecialised

dentition, most similar to that of T. nigrolutea, with large size and relatively gracile proportions,

with a relatively slender coronoid process and deeply concave margin between the coronoid and

angular processes. An additional scincid dentary is probably referable to the genus

Cyclodomorphus. Cranial remains of a species of the gekkonid genus Diplodactylis are described,

representing a group of species within Diplodactylus that have a vestigial jugal. An agamid dentary

and two varanids of differing sizes are also described.

FOSSIL LIZARDS FROM THE PLIOCENE CHINCHILLA LOCAL FAUNA, QUEENSLAND,

WITH DESCRIPTION OF A NEW SPECIES

MN HUTCHINSON & BS MACKNESS

HUTCHINSON, MN & MACKNESS, BS. 2002. Fossil lizards from the Pliocene Chinchilla

Local Fauna, Queensland, with a description of a new species. Records of the South Australian

Museum 35(2): 169-184.

The lizard fauna of the Middle Pliocene Chinchilla Local Fauna consists of members of the

Gekkonidae, Agamidae, Varanidae and Scincidae. A new species of the scincid lizard genus

Tiliqua is described, based on a complete right dentary. The new species combines relatively

unspecialised dentition, most similar to that of 7. nigrolutea, with large size and relatively

gracile proportions, with a relatively slender coronoid process and deeply concave margin

between the coronoid and angular processes. An additional scincid dentary is probably referable

to the genus Cyclodomorphus. Cranial remains of a species of the gekkonid genus

Diplodactylus are described, representing a group of species within Diplodactylus that have a

vestigial jugal. An agamid dentary and two varanids of differing sizes are also described.

Mark N Hutchinson, Department of Herpetology, South Australian Museum, North Terrace,

Adelaide, South Australia 5000. Brian S Mackness*, School of Biological Sciences, University

of New South Wales, Kensington, New South Wales 2052. *Current address: PO Box 560,

Beerwah, Queensland 4519, Australia. Manuscript received 17 April 2002.

The Chinchilla Sand was named by Woods

(1960) for a sequence of weakly consolidated grey

to yellowish and light brown sands, ferruginised

heterogeneous conglomerates, grits, sandy clay

and clays. These outcrops range from shallow

beds to sections several metres deep along a

40 km stretch of the Condamine River valley,

approximately 240 km ENE of Brisbane.

The Chinchilla Sand has yielded a diverse array

of fossil remains, the Chinchilla Local Fauna

(Mackness, Wilkinson & Wilkinson 1999),

Mammals have been the best studied group and

the fauna recovered comprises macropodids

(Bartholomai 1963, 1966, 1967, 1973, 1975,

1976; Flannery & Archer 1983), phascolarctids

(Archer 1977), thylacoleonids (Archer & Dawson

1982; Bartholomai 1962; Woods 1956),

peramelids (Mackness et al 2000), dasyurids

(Archer 1982; Bartholomai 1971; Dawson 1982;

Wroe & Mackness 1998, 2000a, 2000b), a

thylacine (Mackness et al submitted), murid

rodents (Godthelp 1990) and a molossid bat

(Hand et al 1999). At least three lineages of birds

are also known, an emu, a rail and a duck (Olson

1975, 1977; Patterson & Rich 1987). The living

lungfish Neoceratodus and plotosid catfish have

also been recorded (Kemp & Molnar 1981;

Mackness et al 1999). Reptile remains described

so far pertain to larger taxa, such as crocodilians

(Palimnarchus), turtles and madtsoiid snakes

(Bartholomai & Woods 1976; Gaffney 1981;

Gaffney & Bartholomai 1979; Mackness &

Scanlon 1998; Woods 1960, 1962). The only

lizards to date have been Hecht’s (1975)

identification of a number of large varanid fossils

from the Chinchilla Local Fauna as Megalania sp.

The Chinchilla Local Fauna appears to

biocorrelate with the Kanunka Local Fauna of the

Tirari Desert, South Australia (Tedford et al 1992)

and the Spring Park, Bow and Big Sink Local

Faunas (Mackness et al 2000). On the basis of the

Kanunka Local Fauna having a

magnetostratigraphic age of approximately 3.4

million years, the Chinchilla Local Fauna is

estimated to be between late Early Pliocene and

Middle Pliocene in age (Tedford et al 1992).

Lizard fossil remains have received relatively

little attention in reports on Australian fossil

faunas, due both to a paucity of material and a

lack of data concerning osteology. In some

lineages of lizards, such as agamids (Covacevich

et al 1990) and varanids (Mackness & Hutchinson

2000), the restricted amount of morphological

divergence within the living fauna makes

identification problematic. For the anatomically

diverse skinks and gekkonoids, this picture is

steadily changing for the better as new finds focus

attention on specific areas of anatomy. In this

170

study we point out some useful character states of

gekkonid skulls that permit attribution to both

major lineages and more restricted species groups.

We also build on the knowledge of one of

Australia’s most characteristic lizard faunal

elements, the bluetongue skinks of the genus

Tiliqua (Shea & Hutchinson 1992) to describe a

new and distinctive species.

MATERIALS AND METHODS

The specimens described here were recovered

through quarrying or wet sieving of sediments

from ‘Wilkinson’s Quarry’, Chinchilla,

Queensland, a site that has been continuously

worked by Cec and Doris Wilkinson for over 11

years. The sediments are primarily fluviatile in

nature and represent a number of depositional

events. Most fossils in these units occur as

isolated pieces and all represent Pliocene taxa

(Mackness, Wilkinson & Wilkinson 1999).

Specimens were examined using a Wild M3Z

stereomicroscope with eyepiece micrometer and

drawing tube. The fossils are registered in the

collection of the Queensland Museum, Brisbane

and were compared with skeletal material in the

collection of the South Australian and Western

Australian Museums. Comparative scincid

specimens included all living species of Tiliqua

and representatives from all species groups in the

genera Cyclodomorphus and Egernia. The

morphology of the gekkonid fossils dictated the

gekkonid specimens that would be the most likely

candidates for comparison, the ‘unspecialised’

small- to medium-sized diplodactylines. We also

examined representatives of those genera of

gekkonines that occur in the Australian region and

the adjacent Melanesian—Indonesian region, as

well as a scattering of specimens from across the

global diversity of the subfamily Gekkoninae. In

addition, published diagrams and discussions of

gekkonid cranial osteology were consulted to

ensure that our selection was representative,

mainly aimed at justifying some of the general

statements we make regarding major patterns of

variation that differentiate gekkonine and

diplodactyline skulls and mandibles. Specimens

examined are listed in Appendix 1. Terminology

for bones follows Estes et al (1988).

Measurements

Measurements of varanid vertebrae and the

large Tiliqua species were made using Vernier

callipers accurate to 0.05 mm, while smaller

MN HUTCHINSON & BS MACKNESS

specimens and teeth were measured with a

calibrated microscope eyepiece. Vertebral

measurements are summarised below and largely

follow Smith (1976). Statistical analyses of these

measurements are provided in Appendix 2; Tables

1 and 2.

Vertebral length (Pr-Po) — the greatest distance

from the anterior edge of the prezygapophysis to

the posterior edge of the postzygapophysis.

Vertebral width (Pr-Pr) — the maximum width

of the vertebra between the lateral edges of the

prezygapophyses.

Vertebral width (Po-Po) — the maximum width

of the vertebra between the lateral edges of the

postzygapophyses.

Centrum minimum width (BW) — the smallest

distance across the centrum.

Condylar width (CW) — the greatest transverse

diameter of the condyle.

Abbreviation for specimen numbers: AR:

University of New South Wales Research

Collection; SAM: South Australian Museum;

WPC: Wilkinson Private Collection.

SYSTEMATICS

Order SQUAMATA Oppel, 1811

Family AGAMIDAE Gray, 1827

Unidentified material

Material examined

A left dentary (WPC 1354), bearing mid to rear

section of tooth row.

Characters

Agamid reptiles are distinguished in having a

dentition combining one to three anterior

pleurodont teeth followed by acrodont teeth. Other

agamid features are summarised by Estes (1983).

Description

The specimen is from a moderately sized

individual, perhaps of skull length of approximately

30 mm, and bears nine acrodont teeth. The

posterior of the dentary is largely missing although

the facet for the coronoid is still visible. No mental

foramina are discernible on the specimen.

Remarks

Covacevich et al (1990) discussed several

problems in identifying fragmentary agamid

remains beyond family level.

QUEENSLAND PLIOCENE FOSSIL LIZARDS 171

pr. dors.

pr. st.

musc.

D pr. cor. E fac. cor.

F fac. cor.

f. men.

FIGURE |. A-F Diplodactylus cf steindachneri. A, right maxilla (QM F30573) in lateral view and B, medial view.

C, left parietal (QM F30574) in dorsal view. D-F right mandible (QM F30572). D, lingual view of dentary. E,

lingual view of articular + surangular. F, articulated mandible in labial view. Abbreviations: ch. sh. — choanal shelf:

f. men. — mental foramina; fac. art. — articular facet; fac. cor. — coronoid facet; fac. j. — jugal facet; fac. spl. —

splenial facet; musc. — insertion point for dorsal neck musculature; pr. ang. — angular process; pr. cor. — coronoid

process; pr. dors. — dorsal process; pr. st. — supratemporal process; pr. sur. — surangular process; pr. vom. —

vomerine process; rap — retroarticular process; spl. n. — splenial notch. Scale bar = 5 mm.

Family GEKKONIDAE Gray, 1825 Material examined

Right maxilla (QM F30573); left parietal (QM

Subfamily DIPLODACTYLINAE Underwood, F30574); partial right mandible (QM F30572),

1954 consisting of a dentary and fused surangular plus

articular.

Diplodactylus Gray, 1825

Description

Diplodactylus cf steindachneri Maxilla. The maxilla (QM F30573, Fig. 1A-B),

(Fig. 1A-F) a right element, is almost complete, with slight

172

erosion of the edges on the margin of the dorsal

process. Total length 6.1 mm. A slender,

dorsoventrally flattened vomerine process extends

medially from the anteromedial end of the bone.

The facial portion of the maxilla is gently arched

along its dorsal margin. Although the dorsal

margin is slightly damaged, there is no

posterodorsally directed frontal process. The

margin bordering the nasal opening is almost

semicircular. The outer surface of the maxilla

bears two series of foramina; one consisting of a

row of six openings just above the teeth, the

posteriormost the largest; and a second consisting

of three relatively large openings at the level of

the anteroventral corner of the orbit. The

subocular ramus of the maxilla is robust and

relatively short and deep. It does not taper to a

point, as is usual in gekkonoids; the outline of the

caudal extremity of the specimen is bifurcated,

with the dorsal fork larger than the ventral. The

internal surface of the maxilla bears a prominent

choanal shelf, which is constricted at its mid-point

and again anteriorly before projecting as the

vomerine process. The inner face of the suborbital

region has a small facet for a vestigial jugal on the

dorsal fork, below which is a longer facet for the

ectopterygoid.

There are 33 tooth loci, most with intact teeth.

The teeth are small, cylindrical in section and

tapering to acute points, with an apical cusp and

lingual cusp separated by a narrow occlusal

groove; this morphology is regarded as typical and

plesiomorphic for gekkonids (Sumida & Murphy

1987). The individual teeth are uniform in size

and vary only slightly in shape.

Parietal. The left parietal (QM F30574) (Fig.

1C) is complete except for slight erosion along

the medial margin. In common with many living

species, the medial edge of the bone may not

have been fully ossified. Beginning at the bone’s

anteromedial apex, the anterior margin describes

a shallow sinusoidal curve posterolaterally,

terminating at the obtusely pointed apex of the

articulation point with the postorbital. The

caudal margin of the body of the parietal is

almost straight-edged and faces caudolaterally.

This margin bears a shallow caudally opening

pocket, which would have been the insertion

point for the superficial neck musculature. The

supratemporal process is relatively slender and

tapers to an acute point. Ventrally, the bone

bears a descending flange parallel to the lateral

margin, the flange descending to a pointed

projection (epipterygoid process) about halfway

along. Overall length of the specimen is 4.2 mm;

MN HUTCHINSON & BS MACKNESS

width (measured at the apex of the postorbital

articulation) 1.9 mm.

Adult mandible (QM F30572, Fig. 1D-F) is

represented by a complete dentary and nearly

complete compound bone consisting of the fused

articular and surangular bones. The angular,

splenial (or the composite angulosplenial typical

of diplodactylines) and coronoid are absent.

Dentary. Total length (taken as a straight line

from symphysis to tip of angular process) 6.5 mm.

There are 36 teeth or tooth loci, the teeth being

similar in size and shape to those on the maxilla,

with the most anterior teeth having somewhat

more recurved tips.

The dental sulcus is well defined by a lingual

parapet that is evident as far as the posteriormost

tooth. The posterior limit of the bone has three

processes, which define two embayments. The

uppermost (coronoid) process terminates just

behind the last tooth and has facets on its lingual

and labial aspects for the coronoid bone. The

middle (surangular) and lower (angular) processes

would have terminated at about the level of the

apex of the dorsal process of the coronoid bone.

The labial face bears four mental foramina, the

posteriormost lying at the level of the 24th tooth.

On the lingual face, the splenial notch extends

forward to the level of the 27th tooth, and there is

a facet extending anteriorly to the notch,

suggesting that the splenial terminated at about

the level of the 22nd tooth. There is no bony

intramandibular septum exposed within the

splenial notch.

Comparisons

The identification of these specimens as

gekkonoids is based on several characters that are

uniquely combined in this clade. The morphology

of the articular and retroarticular region of the

mandible shows an articular facet facing

posterodorsally and the retroarticular process as

slender and ventrally positioned. There is

complete overgrowth by the dentary of the groove

for Meckel’s cartilage. The teeth are small, finely

pointed and numerous. The parietal is poorly

ossified, paired and lacks a pineal foramen. The

elements are those of a small (approx 45 mm

SVL) generalised gecko.

Studies of the phylogenetic relationships among

the Gekkota (Bauer 1990; Donnellan et al 1999;

Estes et al 1988; Kluge 1967a, 1967b, 1987) agree

that the Australian region has three of the four

major gekkotan lineages: the Gekkoninae,

Diplodactylinae and Pygopodidae. The remaining

gekkotan clade, the Eublepharidae, is found no

QUEENSLAND PLIOCENE FOSSIL LIZARDS

closer than Borneo (the genus Aelurascalabotes).

To further establish the affinities of the Chinchilla

specimens, we have noted several characters that

appear to be useful in diagnosing membership of

particular gekkotan lineages, based on the most

commonly recovered elements: mandibles,

maxillae, frontals and parietals.

Maxilla. The shape of the maxilla differs

markedly between most gekkonines and the rest

of the gekkotan lineages. Authors describing this

area of the face generally emphasise bony

contacts, using the dichotomy of either a

prefrontal-nasal contact (shown by most

diplodactylines) or a frontal—maxilla contact

(shown by most gekkonines) (Fig. 2). In most

Australian gekkonines (Cyrtodactylus, Gehyra,

Heteronotia, Nactus), and in most other

gekkonines examined, the maxilla arches

posteromedially to contact the frontal, terminating

in a frontal process that greatly reduces the

exposure of the prefrontal and excludes the

prefrontal from the nasal. In the remaining

Australian gekkonine Christinus and in

Diplodactyline

173

diplodactylines, the maxilla lacks this frontal

process and the prefrontal includes an

anteromedial process that contacts the nasal,

usually excluding it from the maxilla. In

eublepharids (Aelurascalabotes [Grismer 1988],

Hemitheconyx [Rieppel 1984]) the prefrontals

bear anteromedial processes but they are excluded

from the maxilla due to well-developed

anterolateral processes of the frontal; the maxilla

is similar in shape to that of the diplodactylines.

Pygopods are variable but most tend to show a

gekkonine-like posterodorsal expansion of the

maxilla to contact the frontal. The maxilla of the

Chinchilla gecko has a low, gently arched dorsal

process with no frontal process.

The Chinchilla fossil shows a distinctive feature

of the maxilla that suggests a particular

relationship within the Diplodactylinae. In lateral

view its subocular ramus shows a relatively deep,

bifurcate posterior termination, a characteristic

also seen in a few species of the genus

Diplodactylus. In these species the notched tip of

the terminal portion of the maxilla is correlated

Gekkonine

FIGURE 2. Comparison of the anterior part of the skull in a diplodactyline (Diplodactylus stenodactylus) and an

Australian gekkonine (Gehyra dubia) gecko showing the usual patterns of contact between the frontal (f), nasal (n),

maxilla (mx) and prefrontal (prf) bones. Areas in grey show the anteromedial (nasal) process of the prefrontal

(diplodactyline), and the frontal process of the maxilla (gekkonine). D. stenodactylus also shows a vestigial jugal

(j), compared with the usual degree of development of this bone in most other geckkotans as shown by G. dubia.

174

a rr > FF:

“WUT y- yor IY Uy ee

MN HUTCHINSON & BS MACKNESS

FIGURE 3. Right maxilla (slightly anterodorsal in aspect) and sketch of dorsal view of right orbit in three species

of Diplodactylus, showing reduction of the jugal (grey), A, D. granariensis (R29135); B, D. damaeus (R24553);

and C, D. steindachneri (ROS153). Approx. length of maxillae 6-7 mm, not to scale.

with reduction of the jugal bone reported by

Kluge (1967b). The two species reported as

having the greatest reduction of the jugal (D.

steindachneri and D. stenodactylus) have the most

pronounced development of the bifurcate shape

(Fig. 3A—C) and are essentially identical in

appearance to the Chinchilla maxilla.

Frontal. No frontal has been recovered at

Chinchilla, but future finds should be identifiable

as gekkonine if there are three facets on its

anterior portion (for the nasal, maxilla and

prefrontal) or otherwise if there are only two

facets. The relatively long nasal processes of

eublepharid frontals might also be diagnostic for

that clade.

Parietal. Many gekkotan taxa have distinctive

parietal morphologies. However, this bone shows

marked ontogenetic variation in its degree of

ossification, the elaboration of muscle attachment

points on its lateral and posterior margins, and the

robustness of the supratemporal process. The most

distinctive feature of the Chinchilla specimen is

the concavity for the neck musculature on the

posterior margin. This arrangement is not typical

of most geckos, where the musculature simply

attaches to the posterodorsal surface and edge of

the parietal, or to the posteroventrally deflected

trailing edge, generally with no more than a

depression or a low ridge to mark the point of

attachment. A more-or-less well-defined pocket is

present in some members of the genus

Diplodactylus (D. stenodactylus, D. steindachneri,

D. byrnei, intraspecifically variable in D.

damaeus) but is absent from other Diplodactylinae

examined and was not seen in the gekkonines.

Pygopod parietals are much slenderer than those

of other gekkotans, the midline length being at

least twice the width (cf less than 1.5 times the

width).

Mandible. Some diplodactyline and gekkonine

taxa as well as pygopods (Hutchinson 1997) have

unique modifications of the mandible that are

diagnostic for individual genera, but many

gekkotan taxa have mandibles that are

superficially similar in overall shape. In these

generalised mandibles, the retroarticular process

provides a key to the two major gecko

subfamilies. In gekkonines, the retroarticular

QUEENSLAND PLIOCENE FOSSIL LIZARDS 175

process is generally spoon-like in shape, its dorsal

surface forming an obvious, concave bowl. In

most diplodactylines (Saltuarius and some

Strophurus are exceptions) and in pygopodids, the

retroarticular process is rod-like, with an oval,

flattened or shallowly concave cross-section and

terminated by a club. The morphology of the

dorsal surface of the retroarticular process in

eublepharids has not been described (published

diagrams show only the shape in lateral or ventral

view). The Chinchilla specimen has the clubbed,

diplodactyline type of retroarticular process.

The fossil maxilla and mandible therefore not

only show an _ overall similarity with

diplodactylines, but also share several discrete

character states that indicate specific

diplodactyline affinities. The characteristics that

distinguish the fossils are those that are uniquely

combined in the genus Diplodactylus.

The Chinchilla fossils were found as

disarticulated elements but their size and

preservation, especially the mandible and maxilla,

are consistent with having come from one animal.

Based on the possession of a bifurcate terminus to

pr. cor.

pr. ang.

sym

iaf

the posterior ramus of the maxilla and the

presence of a ‘pocket’ on the trailing edge of the

parietal, they are referred to D. steindachneri. It is

one of three species of Diplodactylus still found

in the region (the other two are D. vittatus and D.

tessellatus, Ingram & Raven 1991), and is the only

one of these three species to have a vestigial jugal

and bifurcate maxillary terminus. The other

species with reduced jugals, members of the D.

stenodactylus group, are found in the deserts of

central and western Australia, the closest to

Chinchilla being D. immaculatus of western

Queensland.

Family SCINCIDAE Gray, 1825

Subfamily LYGOSOMINAE Mittleman, 1952

Tiliqua Gray, 1825

Tiliqua wilkinsonorum sp. nov.

(Fig. 4A-B)

f. men.

S$. sur.

can. inf. aiv.

ims

spl. n.

FIGURE 4. A-B Tiligua wilkinsonorum n. sp. QM F30567 Holotype right dentary. A, labial; B, lingual.

Abbreviations: can. inf. alv. — inferior alveolar canal; cr. sym. — symphysial crest; f. men. — mental foramina; fac.

cor. — coronoid facet; iaf — inferior alveolar foramen (= anterior inferior alveolar foramen); ims — intramandibular

septum; mames — limits of attachment for external adductor musculature; pr. ang. — angular process; pr. cor. —

coronoid process; spl. n. — splenial notch; s. sur. — surangular suture; sym. — symphysis. Scale bar = 5 mm.

176

Material examined

Holotype: a right dentary (QM F30567).

Type Locality

Wilkinson’s Quarry site, Chinchilla area,

southeastern Queensland. The specimen comes

from a fossil-bearing unit within the quarry that

lies unconformably on top of a layer of fine sand.

The sediments are primarily fluviatile in nature

and represent a number of depositional events.

Age

Chinchilla Local Fauna, late Early to Middle

Pliocene (Tedford et al 1992).

Diagnosis

A large species, dentary tooth row length

33 mm, differing from other Tiliqua by the

following combination of characters: all tooth

crowns pointed and retaining an occlusal ridge,

the crown scarcely wider than the shaft; coronoid

process of the dentary relatively slender, narrower

than long, its shape correlated with a curved and

concave, rather than angular or straight, dentary—

surangular suture connecting the base of the

coronoid process with the tip of the angular

process.

Description

A right dentary, bearing a complete row of 20

teeth or tooth loci. The specimen is almost intact,

being slightly damaged at the apex of the splenial

notch and on the ventral edge adjacent to a

repaired crack through the jaw at the level of the

15th tooth. Total length, from the anteriormost

point of the symphysis to the tip of the angular

process, 47.8 mm. Depth at level of 16th tooth

(excluding tooth), 7.8mm. Maximum depth of

jaw, measured vertically from the apex of the

coronoid process, 13.1 mm. Maximum width,

measured in occlusal view at the level of the 16th

tooth, 8.0 mm.

Meckelian groove closed, no trace of suture. A

pronounced crest runs caudally from the

symphysis along the ventrolingual margin of the

jaw, gradually merging with the body of the

dentary at about the level of the eighth tooth.

Apex of the splenial notch at about the level of

the 12th or 13th tooth, slight uncertainty due to

the broken edge of the inferior alveolar foramen

at the apex of the notch. Labial surface with

pronounced, arcuate adductor muscle scar that has

its apex level with the last tooth. Angular process

projects 15 mm beyond the level of the last tooth.

Dorsal margin of angular process merges with the

MN HUTCHINSON & BS MACKNESS

ventral margin of the coronoid process along a

smooth concave curve that has its apex at the level

of the tooth row. Coronoid process well

developed, flattened, but relatively small

compared with other Tiliqua. A series of mental

foramina starts at the level of the 11th tooth and

runs anteriorly to the symphysis. These foramina

are mostly arranged as a series of superposed

pairs, with a total of about nine in the series.

Intramandibular septum fused ventrally along

its length to the ventral lamina of the dentary,

completely separating the inferior alveolar canal

from the Meckelian canal. Caudal margin of

intramandibular septum with a deep, curved

notch, bounded ventrally by a caudally projecting

prong.

Tooth row, in occlusal view, almost straight,

gently curving medially anterior to the level of the

sixth tooth. Labial wall of dental sulcus tall and

robust. Lingual wall low but distinct anteriorly,

reducing caudally and disappearing by about the

level of the 16th tooth. Lingual face of dentary

below the dental sulcus vertical, sharply distinct

from the tooth row. Dentition pleurodont. The

first of 20 teeth is broken off and the 10th, 12th

and 19th loci are empty, but all other teeth are

present and well preserved. The teeth vary

distinctly in size, becoming larger progressing

caudally, with the maximum reached in positions

13 to 16, the subsequent teeth then diminishing in

size. As a guide to the changes in tooth size, tooth

6 is 3.5mm high by 1.3mm wide (measured

normal to the lingual aspect) while tooth 14 is

4.3mm high by 2.4mm wide; thus, height

increases by 23% and width by 85%. In lingual

view, the tooth crowns are acutely pointed

anteriorly, becoming obtusely pointed by about

the 13th tooth. The crown of each tooth is

margined by an occlusal ridge that separates its

lingual and labial surfaces. In mesial view the

tooth crowns have a parabolic curved outline

labially, but the lingual surface is flatter,

descending almost vertically from the apical ridge

and forming an angular contact with the rounded

labial surface. On the largest teeth, several low

striae run vertically down from the apical ridge on

both labial and lingual surfaces of the crown.

Comparisons

The combination of the closed Meckelian

groove, enlarged ‘cheek’ teeth with more than one

large tooth posterior to position 10, and an

enlarged and flattened coronoid process is unique

and diagnostic for the bluetongue lizards of the

genus Tiliqua (Shea 1990). The related genus

QUEENSLAND PLIOCENE FOSSIL LIZARDS 177

Cyclodomorphus has only a single enlarged cheek

tooth, while other lygosomines with a closed

Meckelian groove lack such obvious dental

differentiation.

Most species of Tiliqua, however, have a more

specialised dentition than 7. wilkinsonorum. The

plesiomorphic tooth crown morphology in skinks

includes a convex outer surface, curving

occlusally to a pair of apical ridges separated by a

groove (M. H. pers. obs.; Sumida & Murphy

1987), while the lingual surface is flattened and

oriented almost vertically, forming an angular

contact with the apical ridge. In lingual and mesial

views the crown is acutely pointed. In contrast to

this, species of Cyclodomorphus and Tiliqua have

teeth modified for durophagy (Estes & Williams

1984). Most species of Tiliqua, including T.

gigas, T. multifasciata, T. occipitalis, T. pusilla,

T. rugosa and T. scincoides, have cheek teeth with

expanded crowns, and have lost or greatly reduced

the plesiomorphic asymmetry of the crown. In

occlusal view the crown is circular in shape, and

when viewed mesiodistally the crown profile, a

convex curve, is similar in shape lingually and

labially. The apex of the tooth has a central

conical projection with pronounced striae

radiating over the crown from this central point.

Most individuals lack an occlusal ridge, although

many 7. occipitalis and some T. rugosa have a

remnant on the tooth apex. Tiliqua adelaidensis

has quite different dentition in which the enlarged

cheek teeth are labiolingually compressed, with a

pronounced cutting edge formed by the apical

ridge. Tiliqua nigrolutea is the living species that

shows the closest dental similarity to T.

wilkinsonorum. The teeth show little expansion of

the crowns, which are pointed and retain an

obvious occlusal ridge that demarcates the crown

into lingual and labial aspects. These two species

show an asymmetric tooth crown profile in

mesiodistal view, with a more steeply descending

lingual surface. Compared with T. nigrolutea, the

teeth of T. wilkinsonorum differ only in being

more robust and having the crowns of the largest

teeth more expanded and obtusely pointed. These

differences could be attributed to allometry or

individual variation in tooth proportions, which is

considerable in living Tiliqua (M. H. pers. obs.).

However, other characteristics of the fossil

specimen differentiate it from both T. nigrolutea

and other Tiliqua species.

The dentary is from a very large Tiliqua. Using

the average ratio of tooth row to mandible length

(0.48) that applies in extant Tiliqua, the tooth row

length of 7. wilkinsonorum of 33 mm extrapolates

to an approximate mandible length of 68 mm.

This size is approached only by the largest males

of T. rugosa and T. scincoides (G. M. Shea pers.

comm.) and suggests that 7. wilkinsonorum may

have had a snout-vent length close to 400 mm and

a mass in excess of one kilogram. In spite of this

size, the dentary of T. wilkinsonorum is notably

gracile. This is most clearly seen in the shape of

the coronoid process and the surangular suture

running from the base of the coronoid process to

the tip of the angular process. In all extant Tiliqua,

the coronoid process is dorsally and caudally

expanded. This expansion is masked to some

extent because the base of the coronoid process is

absorbed by caudal expansion of the trailing edge

of the dentary such that the surangular suture is

not deeply incised into the labial lamina of the

dentary. The precise course of this suture varies

both inter- and intraspecifically (Fig. SA-I). In T.

nigrolutea, T. occipitalis and T. rugosa, this

suture usually runs vertically from the coronoid

process and then angles caudoventrally to the tip

of the angular process of the dentary. In T.

adelaidensis, T. gigas, T. multifasciata and T.

scincoides, the suture tends to run caudoventrally

in an almost straight line from the base of the

coronoid process. In none does it trace the

excavated curve seen in T. wilkinsonorum (and

other skinks). In T. wilkinsonorum the coronoid

process is less expanded than in any extant

species, with the possible exception of T. gigas,

but is more clearly defined ventrally by the

anterior sweep of the concave surangular suture.

In living Tiliqua species, the expanded coronoid

process and adjacent parts of the dentary increase

the strength of this region of the mandible and

increase the area of attachment for the jaw-closing

tendon of the m. adductor medius externus

superficialis (Haas 1973). The relative slenderness

of the coronoid process and the concave

surangular suture in 7. wilkinsonorum are closer

to the morphology seen in other skinks, including

Egernia, the genus that probably includes the

sister group of the Tiliqua lineage.

In summary, 7. wilkinsonorum is one of the

largest known specimens of Tiliqua, and has less

specialised teeth than all but 7. nigrolutea and a

more slender (plesiomorphic) coronoid process of

the dentary than all but T: gigas. The morphology

of the teeth and the relative gracility of the caudal

end of the dentary suggest that 7. wilkinsonorum

was not as specialised for durophagy as most

other large Tiliqua.

Tiliqua wilkinsonorum is the third species of

Tiliqua to be found in the Tertiary of Australia

178 MN HUTCHINSON & BS MACKNESS

FIGURE 5S. Dentary variation in Tiliqua. Detailed views showing variation evident in the degree of expansion of the

coronoid process of the dentary and the shape of the dentary-surangular suture. All to same scale except 7.

adelaidensis (inset); both scale bars = 5mm. A, T. wilkinsonorum. B, T. rugosa R27028. C, T. adelaidensis

R40738. D, T. nigrolutea RO2725. E, T. nigrolutea R47698. F, T. occipitalis R35758. G, T. scincoides R27039. H,

T. gigas R11419. I, T. multifasciata, R35757.

and the second extinct species. Shea and

Hutchinson (1992) described a very small species,

T. pusilla, from the early Middle Miocene of

Queensland, and specimens indistinguishable

from a living species, T. scincoides, have been

recorded from the Pliocene of South Australia

(Pledge 1992) and Queensland (Mackness &

Hutchinson 2000).

cf. Cyclodomorphus Fitzinger, 1843

Material examined

A partial dentary (QM F30568).

Description

The specimen represents about one-third of a

left dentary, lacking the anterior portion, anterior

to the last six teeth and lacking the coronoid and

angular processes. It is not a juvenile, based on

several cycles of tooth replacement being evident.

The groove for the Meckelian cartilage is

completely obliterated by the dentary. The third-

last tooth is markedly enlarged. The tooth anterior

to this is absent, but its locus is smaller, while the

other remaining teeth are smaller again. A single

large mental foramen is present on the labial

surface of the jaw at the level of the fourth-last

tooth. The teeth, both enlarged and small, have

blunt, slightly laterally compressed crowns with

virtually no apical ornamentation such as cusps,

grooves or striae.

Comparison

The fossil appears to have only one markedly

enlarged cheek tooth, with a second moderately

enlarged tooth anterior to it. Among extant

Australian skinks, only members of the Tiliqua

lineage combine a closed Meckelian groove and

enlarged durophagous cheek teeth (Shea 1990).

QUEENSLAND PLIOCENE FOSSIL LIZARDS 179

The dentitions of the two genera of this lineage,

Tiliqua and Cyclodomorphus, differ in that adults

of Tiliqua have several enlarged cheek teeth while

those of Cyclodomorphus usually have only one

strongly enlarged tooth. On this basis as well as

its small size and overall shape, the Chinchilla

specimen is more similar to Cyclodomorphus than

it is to Tiliqua. However, Tiliqua adelaidensis and

the extinct T. pusilla show that the small species

of Tiliqua may have only two enlarged cheek

teeth, so that the distinction on tooth pattern alone

is not entirely conclusive. Other attributes

distinguishing the two genera, including the shape

of the symphysial region, are not preserved on the

specimen. In the shape of the crowns and lack of

occlusal striae, the fossil is unlike any species of

either Cyclodomorphus or Tiliqua examined in

this study, but it is too fragmentary to either

allocate to a genus with certainty or to warrant

description as a new taxon.

Family VARANIDAE Gray, 1827

?Megalania sp.

Material examined

Five isolated dorsal vertebrae (WPC 115, 116,

2409, 2014, 3322), three isolated dorsal fragments

(WPC 2040, 3555, 3556), four isolated caudal

vertebrae (WPC, 1429, 1573, 2364, 2524).

Characters

The genus Megalania is characterised in part

by having massive thoracic and lumbar

vertebrae with weakly developed zygosphenes

(absent in typical Varanus) as well as small

depressed neural canals. The adult teeth of

Megalania are large and slightly recurved

distally. The anterior cutting edge is rounded

and serrated distally. The posterior cutting edge

is thin, blade-like and serrated along its entire

length (Hecht 1975).

Description

The dorsal vertebrae are much more massive

than those of any extant varanid but compare well

in morphology with those from an extant varanid

Varanus varius (AR 7641) as well as from large

fossil varanids from the Bluff Downs Local Fauna

(Appendix 2; Tables 1 and 2). The Chinchilla

specimens are 62% larger than those

measurements supplied for Australia’s largest

extant varanid, the perentie Varanus giganteus by

Smith (1976) and 20% larger than those for the

large fossil varanid from Bluff Downs Local

Fauna (Mackness & Hutchinson 2000).

Remarks

A single maxilla (QM F874) from the

Chinchilla Sand was referred to Varanus dirus

(Hecht 1975), a taxon originally described by De

Vis (1889) on the basis of an isolated tooth from

the Pleistocene Kings Creek locality. Hecht (1975)

synonymised V. dirus with Megalania prisca.

Four isolated vertebrae (two caudals and two

dorsals) have also been collected from the

Chinchilla Sand and, although Hecht (1975)

referred them to Megalania, he also suggested that

this material may represent a separate Pliocene

species.

The assignment of the larger varanid vertebrae

described in this paper to ?Megalania is done

purely on the basis of convention. Opinion is

divided as to the current status of the genus, with

Hecht (1975) and Molnar (1990) concluding it is

valid but Estes (1983) and Lee (1996) suggesting

that Megalania should be synonymised with

Varanus. A large collection of varanid fossils

from Pliocene localities is presently under study

by the authors.

Varanus sp.

Material examined

Two isolated dorsal vertebrae (WPC 118,

1430), one isolated caudal vertebra (WPC 3557).

Characters

These vertebrae are identified as varanid by

their distinctive condyle—cotyle articulations. The

dorsal vertebrae also show the characteristic

constriction of the centra anterior to the condyle

seen in Varanus.

Description

The vertebrae show characteristic varanid

morphology with a range of centra lengths

(Appendix 2; Table 3) that indicate a medium-

sized goanna of the size of Varanus gouldii.

Remarks

Although Wilkinson (1995) suggests some

characters for separating varanid species on the

basis of vertebral morphology, many of these rely

on the neural spine being present. This feature is

missing on all three fossil vertebrae. Interspecific

and/or intraspecific variation in varanid vertebrae

have yet to be fully tested; therefore, characters

180

identified by Wilkinson (1995) must be used with

caution.

DISCUSSION

The composition of the lizard fauna from the

Chinchilla Sand Local Fauna, at least at family

level (Appendix 2; Table 4), is similar to that of

the Bluff Downs Local Fauna. This is the only

other Australian Pliocene locality with a

significant lizard record (Mackness & Hutchinson

2000). It differs from Bluff Downs in the generic

representatives of each family, but the significance

of this is unknown; it could merely be stochastic,

reflecting the very small sample sizes at each

locality. The palaeoenvironment was primarily

fluviatile for Chinchilla and lacustrine and

fluviatile for Bluff Downs.

The exact nature of the Chinchilla

palaeoecology has yet to be determined. While

there may have been a significant wetland

component, as suggested by the presence of

various aquatic taxa such as turtles (Gaffney 1981;

Gaffney & Bartholomai 1979) and waterbirds

(Olson 1975, 1977), there is also evidence that the

environment was highly seasonal (Mackness et al

1999; Wroe & Mackness 2000). The presence of

arboreal species such as Koobor (Archer 1977)

and an unnamed phascolarctomorph (Mackness et

al 1994) would be consistent with a complex and

mature forest, but the composition of the

mammalian fauna indicates a lack of undoubted

MN HUTCHINSON & BS MACKNESS

closed-forest species. Further, the predominance

of grazing kangaroos and diprotodontids suggests

that grasslands were also part of the environment.

The presence of an extinct reptile species at

Chinchilla is noteworthy given the slow rate of

faunal turnover in reptile populations (La Duke

1991). Tiliqua wilkinsonorum represents a

distinctive but extinct species of lizard, a

relatively uncommon observation for the Pliocene

(Estes 1983). A conclusion that can be drawn

from the presence of such ‘fossil endemics’ is that

they signal ecological or geographical factors at

Chinchilla during the Pliocene that are no longer

present in recent environments.

ACKNOWLEDGMENTS

The authors thank Cec and Doris Wilkinson for their

tireless efforts in recovering fossil remains from the

Chinchilla Local Fauna over the past 12 years and for

making the lizard material available for us to study.

Glenn Shea, Jim Mead, Michael Archer and Suzanne

Hand provided helpful comments on the manuscript.

The study of the Chinchilla material was supported in

part by an ARC Program Grant to M Archer; a grant

from the Department of Arts, Sport, the Environment,

Tourism and Territories to M Archer, S Hand and

H Godthelp; a grant from the National Estate Program

Grants Scheme to M Archer and A Bartholomai; and

grants in aid to the Riversleigh Research Project from

the University of New South Wales, Wang Australia

Pty Ltd, ICI Australia and the Australian Geographic

Society.

REFERENCES

Archer, M. 1977. Koobor notabilis (de Vis), an unusual

koala from the Pliocene Chinchilla Sand. Memoirs of

the Queensland Museum 18: 31-35.

Archer, M. 1982. Review of the dasyurid (Marsupialia)

fossil record, integration of data bearing on

phylogenetic interpretation, and suprageneric

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QUEENSLAND PLIOCENE FOSSIL LIZARDS 183

APPENDIX 1

Osteological material used for comparisons. Specimens are from the Herpetology Section of the

South Australian Museum and the Western Australian Museum (WAM prefix).

Gekkonines

Christinus marmoratus (R35562; R39934),

Cosymbotus platyurus (R36789), Cyrtodactylus

louisiadensis (R14002), Gehyra australis

(R34136), G. dubia (R42872), G. sp. (2n = 44)

(R44595), G. catenata (R35561), G. oceanica

(R08687), G. purpurascens (R36377), G. robusta

(R34222), Gekko gecko (R35559), G. japonicus

(R07435), G. vittatus (R55978) Gonatodes

humeralis (R40088), Gymnodactylus darwinii

(R40090), Hemidactylus frenatus (R35564),

Heteronotia binoei (R09751 A, R33565,

R39935), Nactus cheverti (RO9740 B), Paroedura

picta (R55105), Phelsuma madagascariensis

(R40025), Ptyodactylus hasselquisti (R49804),

Tarentola annularis (R40024), Thecadactylus

rapicauda (R40099-100).

Diplodactylines

Bavayia ornata (RO6752), Crenadactylus

ocellatus (RO3113 C), Diplodactylus byrnei

(R13514), D. conspicillatus (R19967), D.

damaeus (R20017, R24553) D. ‘granariensis’

(SA populations) (RO2010, R26494, R29135), D.

immaculatus (R42676), D. pulcher (R26383), D.

steindachneri (RO5153, R52746), OD.

stenodactylus (RO7592, R21240, R26777), D.

tessellatus (RO3876, R40938), D. vittatus

(R35568), Hoplodactylus pacificus (R11025),

Nephurus asper (R35567), N. laevissimus

(R14987 A), N. levis (RO0309 A, R27044—-45), N.

milii (R55422, R57083), N. stellatus (R32297),

Oedura lesueurii (R33597), O. marmorata

(R42893), O. monilis (R35563), O. tryoni

(R33583), Phyllurus platurus (R35054),

Rhynchoedura ornata (R35827), Saltuarius

swaini (R29205), Strophurus ciliaris (R10697,

R35566), S. intermedius (R14325 C).

A range of pygopod skulls was also examined

(see Hutchinson 1997 for listing).

Egernia group skinks

Corucia zebrata (R35765), Cyclodomorphus

maximus (WAM R77193, WAM R77637), C.

michaeli (R35682), C. gerrardii (R35761,

R47699), C. melanops elongatus (R03231,

RO3856 G, R35681), C. venustus (R38021),

Egernia coventryi (R35686, R47693-94), E.

cunninghami (R35680, R35763), E. depressa

(R03433 H), E. formosa (WAM R65803), E.

hosmeri (R22510), E. inornata (RO7234, RO7238,

RO7245 A, R35687), E. kingii (WAM R36376,

WAM R89269-70), E. luctuosa (WAM R36019),

E. major (R27043, R35762), E. multiscutata

(R08469, R25245), E. napoleonis (R35692,

R51076, WAM R45350), E. pilbarensis (WAM

R78945), E. pulchra (R40026, WAM R71884), E.

richardi (R10841), E. saxatilis intermedia

(R43961), E. stokesii (RO2560, R41913), E.

striata (RO7179, RO7198), E. striolata

(R02909 B, R16593, R26891, R38019), E. whitii

(R02726, R35690, R27042, R35689, R45316,

R34886, R35688, R35691), Tiliqua adelaidensis

(R40738, R40745, R43412), T. gigas (R11419),

T. multifasciata (R27041, R35757), T. nigrolutea

(R0O2725, R27048, R27050, R47698), T.

occipitalis (RO2724, R25369, R27047, R35758),

T. rugosa (R02563[2], RO2564, R27026, R27028,

R02801, R25615, R27027, R27029, R27584,

R31855, R35760), T. scincoides (RO2561,

R2703640, R35759, R38020, R43962, R27094).

184 MN HUTCHINSON & BS MACKNESS

APPENDIX 2

TABLE 1. Measurements (mm) of ?Megalania dorsal vertebrae from Bluff Downs and Chinchilla Local Faunas

compared with Varanus giganteus. Measurements as defined in Methods. Range (mean + standard deviation). Data

for V. giganteus taken from Smith (1976), for Bluff Downs from Mackness & Hutchinson (2000).

Specimen No Pr-Po BW/Pr-Po CW/Pr-Po Pr-Pr/Pr-Po

V. giganteus 20 24.5-27.1 (25.740.14) 0.54-0.64 (0.58+.005) — 0.55—0.63 (0.59+.006) — 0.88-1.01(0.92+.007)

Bluff Downs 7 24.5-27.1 (25.7+40.14) 0.54-0.64 (0.584.005) 0.55—-0.63 (0.59+.006) — 0.88-1.01(0.92+.007)

Chinchilla 4 24.5-27.1 (25.7+0.14) 0.54-0.64 (0.58+.005) — 0.55—0.63 (0.59+.006) 0.88-1.01(0.92+.007)

TABLE 2. Measurements (mm) of individual ?Megalania vertebrae. Measurements as defined in Methods.

Specimen Pr-Po Pr-Pr Po-Po BW CW COW CEL

WPC116 39.4 33.2 39.3 20.5 26.1 27.6 25.0

WPC2014 38.9 46.4 41.4 - - 30.4 -

WPC2409 48.3 57.3 49.5 26.4 29.4 31.6 28.2

WPC3322 37.2 43.5 40.4 17.6 23.2 25.2 26.4

WPC3555 56.8 - - - - - -

WPC3556 44.3 - - - -

(Mean/SD) (41.6+6.6) (47.6466) 42.6446) (21.5445) (262+43.1) (28.7429) (265+ 1.6)

TABLE 3. Measurements (mm) of centra of dorsal vertebra of fossil Varanus sp. from Chinchilla Local Fauna

(WPC), Bluff Downs Local Fauna (QMF) and extant Varanus varius (AR7641).

Specimen Measurement Specimen Measurement

WPC118 12.2 AR7641 (a) 15.3

WPC1430 16.5 AR7641 (b) 15.3

QM F7774 14.2 AR7641 (c) 15.4

QM F23238 10.5 AR7641 (d) 15.4

QM F23659 10.8 AR7641 (e) 15.6

QM F23683 13.8 AR7641 (f) 15.7

TABLE 4. Comparison of lizard taxa recovered from the Queensland Pliocene (Chinchilla and Bluff Downs Local

Faunas. Source: Mackness & Hutchinson 2000). Code: present *, absent x, endemic form “.

Taxa Chinchilla Bluff Downs

Gekkonidae

cf Heteronotia x °

Diplodactylus 2) x

Agamidae

unidentified material ° °

Varanidae

?Megalania ° °

Varanus ° °

Scincidae

Tiliqua 7 °

Cyclodomorphus ° x

Egernia x °

Eulamprus x °

FIRST REPORT OF THE CUPRESSACEAN CONES IN THE EOCENE

EYRE FORMATION OF SOUTH AUSTRALIA

N. S. PLEDGE

Summary

Conifer impressions are a very minor component in the various ‘silcrete flora’ assemblages found in

the interior of South Australia. Previously, no fruiting bodies had been found, although numerous

foliage species have been recognised. This note describes impressions of the first woody ovulate

cones found, discovered in a small block of silicified sandstone near Lake Hart, west of Woomera.

They are distinctive and do not appear to bear close affinity to any modern Australian species, but

resemble cones of the African Widdringtonia.

FIRST REPORT OF CUPRESSACEAN CONES IN THE EOCENE

EYRE FORMATION OF SOUTH AUSTRALIA

NS PLEDGE

PLEDGE, NS. 2002. First report of cupressacean cones in the Eocene Eyre Formation of South

Australia. Records of the South Australian Museum 35(2): 185-191.

Conifer impressions are a very minor component in the various ‘silcrete flora’ assemblages

found in the interior of South Australia. Previously, no fruiting bodies had been found, although

numerous foliage species had been recognised. This note describes impressions of the first

woody ovulate cones found, discovered in a small block of silicified sandstone near Lake Hart,

west of Woomera. They are distinctive and do not appear to bear close affinity to any modern

Australian species, but resemble cones of the African Widdringtonia.

NS Pledge, South Australian Museum, North Terrace, Adelaide, South Australia 5000.

Manuscript received 19 April 2002.

Besides Athrotaxis, the modern Australian flora

contains three genera of cupressacean conifers:

Callitris, Actinostrobus and Diselma. Recent

morphological and molecular work by Gadek et al

(2000) shows that the latter are closely related to

an African genus Widdringtonia, within a

monophyletic group, the subfamily Callitroideae,

comprising all Southern Hemisphere

cupressaceans except Athrotaxis.

Fossil conifers, including members of the

Cupressaceae, are sparse in the Australian fossil

record (Hill & Carpenter 1989). Many reported

specimens have occurred as impressions in fine-

grained sediments, with consequent difficulty in

interpreting fine structure, but organic remains

also occur, eg in Tasmania, and have yielded

valuable material (ibid). This has more than

doubled the number of genera in the Australasian

region, with taxa that occur today in New Guinea,

New Zealand and elsewhere, as well as several

extinct genera (Hill & Brodribb 1999).

Plant impressions in silicified sandstone have

been known from the Woomera area / Arcoona

Plateau for many years (Chapman 1937), and as

far as Stuart Creek to the north and Clayton

Station on the Birdsville Track to the northeast.

Most of the moulds are of angiosperm leaves

(Rowett 1997), a few of which have been

described (Chapman 1937; Greenwood et al

2001). Conifer vegetative shoots are a minor

unobtrusive component, studied by Nunn (1964)

and Offler (1969, 1984). In the 1970s localities

were found yielding natural moulds of fruiting

bodies, mostly of myrtaceous origin (Ambrose et

al 1979; Lange 1978). A few specimens referable

to fruit of Proteaceae (Banksia sp.) and

Casuarinaceae have been noted (Greenwood et al

2001; this writer, unpubl.) but, until now, no

coniferous cones have been recognised, despite

the widespread occurrence of vegetative shoots.

Nunn (1964) recognised 21 different conifer

morphotypes, some of which could be related to

modern species from the Australasian region. She

stressed the problems and uncertainties of

identifying the species.

These plant fossil impressions occur in a

number of isolated, discrete localities over an area

of thousands of square kilometres, often in

apparent channel deposits, eg at Nurrungar, Island

Lagoon, near Woomera (pers. obs. 1969), in what

has been regarded as Eyre Formation s.]. Their

age has long been in dispute (Ambrose et al 1979;

Callen & Lange 1986; Chapman 1937;

Greenwood et al 1990; Wopfner et al 1974) but

recent discoveries at Nelly Creek, Lake Eyre

South, have at last related them to deposits datable

by palynological studies to the middle Eocene

(Alley et al 1996; Callen & Cowley 1995;

Christophel et al 1992) within the Eyre Formation.

Variations in the contained fossil floral

assemblages suggest that the different localities

probably have slightly different ages.

MATERIAL AND METHODS

In 1979 members of the Woomera Natural

History Society discovered plant impressions near

186

sMarree

t:..

Po Ne

L orrens

L. Hart o ok

& ve 6

7 g

AUSTRALIA

Adelaide

Ca)

FIGURE 1. Locality map, Lake Hart and other ‘silcrete

flora’ localities in South Australia.

Lake Hart in the Woomera Prohibited Area (Fig.

1). Subsequently, L. Marsh and T. Nurenberg

presented a slab of quartzite to the South

Australian Museum (SAM P22732; Fig. 2)

bearing the impressions of a few leaves and seven

fruit-like bodies. Silicone rubber casts (Fig. 3)

taken from these moulds disclosed fruits strongly

reminiscent of Callitris ovulate cones. Closer

examination, however, showed that, instead of the

six subequal valve scales (2 triplets) seen in

Callitris, these fruit had two unequal pairs of

valves (Fig. 4). Re-examination of the collection

of silcrete flora material in the Museum failed to

uncover any further specimens; SAM P22732 is

therefore unique.

Comparison was made with dried specimens in

the South Australian State Herbarium and with

trees growing at the Mount Lofty Botanic

Gardens.

NS PLEDGE

DESCRIPTION

Order CONIFERALES

Family CUPRESSACEAE Neger

Genus Incertae sedis

The cones are globose to ellipsoidal in shape,

with a slightly conical distal apex. The ellipsoidal

shape may be due to slight compression during

burial, but the flattening is always the same

relative to the smaller valve scales and symmetry

of the cone, and compression is therefore an

unlikely cause. Dimensions vary from about

17 mm diameter and a length of up to 22 mm for

the globose forms, to 15-22 mm diameter x

20 mm in length for the ellipsoidal specimens. In

apical view the pair of larger valves are at the

ends of the apparent ellipse. The smaller,

narrower, valves meet apically along a contact line

of about 5 mm, thus separating the pair of larger

valves (Fig. 4). Although apparently mature, the

cones have a relatively smooth surface, with none

of the roughness that characterises the opened

cones of many Callitris species. However, one

cone, at least, shows slight apically convergent

ribs on the valves. This is reflected in an

impression of what appears to be a decorticated

cone (Fig. 3), having exposed ribs that have the

same symmetry and relationships as the valves.

None of the fruits has split to release its seeds, so

the extent of the sutures separating the valves may

not be fully expressed. Nevertheless, the sutures

extend slightly more than halfway towards the

base. The valves each have a near-apical, small

(roughly 1 mm), circular to elliptical scar,

arranged symmetrically on the fruit (Fig. 4). The

origin or purpose of these scars is unclear,

although they are in the same position as the

‘spurs’ on the valves of some species of Callitris

(Baker & Smith 1910: 47) and species of

Papuacedrus, Widdringtonia and Tetraclinis

(pers. obs.; Hill & Carpenter 1989; McIver 2001).

The base and attachment are seen in one

specimen, showing a petiole expanding gradually

to about 5 mm diameter at its junction with the

fruit, where there is an expanded leaf scale below

the smaller valve (Fig. 4). Another specimen

seems to show a 7-8 mm diameter subcircular

scar on the base, where the stem attached.

Comparisons

Initial comparisons were with species of

Callitris. The obvious difference is in the number

CUPRESSACEAN CONES IN THE EOCENE EYRE FORMATION 187

oo S

FIGURE 2. A, the fossil slab, SAM P 22732. B, positive cast of part of the slab, replicating the original appearance

of the leaf litter at the time of burial. Scale in mm.

188 NS PLEDGE

of valves in the cones, Callitris having three large

valves alternating with three smaller ones, all

ending acutely. Amongst living Australian

conifers, there is only one species having two

pairs of valves in the same decussate geometry as

the fossils: Diselma archeri from Tasmania.

However, its ovulate cones are tiny, barely larger

in diameter than the shoots they terminate (pers.

obs.). The living Papuacedrus from the island of

New Guinea also has two pairs of valves, but its

cones too are small and conical (Hill & Carpenter

1989).

A passing note in Baker and Smith (1910)

directed attention to several non-Australian

genera. Diselma had once been synonymised with

Fitzroya from Chile and Patagonia, just as

Callitris had been joined with Tetraclinis from

Mediterranean North Africa and Widdringtonia

FIGURE 3. Positive silicone rubber cast of the better fossil cones, showing 1, cone with apical view of sutures and

valves with terminal scars; 2, cone with attached stem and basal leaf-scale; 3, cone with basal attachment scar; 4, a

decorticated cone; 5, foliage fragment. Scale in mm.

CUPRESSACEAN CONES IN THE EOCENE EYRE FORMATION 189

spp. from southern Africa. I was not able to see

fruits of Fitzroya, but am informed that they are

nothing like the fossils (R. Hill, pers. comm., July

2002). Through the help of Ms H. Vonow at the

South Australian Herbarium, I examined dry

specimens of Diselma archeri, Tetraclinis

articulata, Widdringtonia cupressoides, W. whytei

and others and Papuacedrus papuana, and living

Widdringtonia spp. in the Mount Lofty Botanic

Gardens. There is an obvious similarity between

these species and the fossil cones in the number

and symmetry of the valves. However, Tetraclinis

can be removed from consideration on the basis

of two characters: the spurs near the tips of its

valves are quite enlarged, particularly on immature

cones (there is obviously some allometric growth

involved here), and the foliage structure is quite

different from the fossil shoot associated with the

cones. Offler (1969) indicates that Papuacedrus

foliage is widely represented in her material, and

—

FIGURE 4. Detail of cones; silicone rubber cast, showing 1, cone with apical view of sutures and valves with terminal

scars, 2, cone with attached stem and basal leaf-scale, 3, cone with basal attachment scar, 4, foliage fragment.

190

it has been recognised in the Eyre Formation of

the Poole Creek Palaeochannel near Lake Eyre

South (Alley & Pledge 2000: 69). Fossil foliage

has also been recorded from various Oligocene

and Miocene sites in Tasmania (Hill & Brodribb

1999; Hill & Carpenter 1989). However,

Papuacedrus differs considerably from the fossils

in the tear-drop shape and small size of the cones,

and in the presence of bract-like appendages

halfway up each cone scale (Hill & Carpenter

1989).

There are two genera considered closely related

to Papuacedrus, namely Libocedrus from New

Zealand and New Caledonia, and fossil in

Tasmania, and Austrocedrus from South America

and also fossil in Tasmania (Hill & Carpenter

1989). They have ovulate cones of the same order

of size as Papuacedrus (and therefore are much

smaller than the Lake Hart fossils) but are similar

in having a structure subapically on the abaxial

surface of each cone scale: a spine in the case of

Libocedrus and a minute tubercle in Austrocedrus

(ibid). However, the cone scales of Austrocedrus

are all apically acute with none truncated; therefore,

that genus can probably be ruled out of contention.

The woody ovulate cones of the Widdringtonia

spp. vary between species in both size and

morphology. Like the fossils, they have two

unequal pairs of valve scales, the upper pair

having truncated apices that therefore meet along

a contact of several millimetres. Of those

examined, cones of W. cupressoides (Fig. 5)

resemble the fossils most closely, although they

tend to be smaller and have more prominent spurs

on the valves, and the overall shape is more tulip-

like, with a depressed base. Other species show a

rougher, more rugose and warty or tuberculose

surface on the valves than do the fossils, but this

may be a reflection of the maturity of the fruit.

The Lake Hart fossils differ markedly from the

North American Cretaceous species

Widdringtonia americana (McIver 2001),

primarily in the development of a prominent umbo

or boss on each valve scale in the latter species.

The vegetative shoot (Figs 3, 4) associated with

the fossil cones is poorly preserved (the grain-size

of the sediment approaching that of the smaller

features), and cannot be guaranteed to relate to

the cones. It seems to have the same square cross-

section and short decussate, scale-like leaves as

Diselma archeri (Clifford & Constantine 1980).

This is unlike Widdringtonia cupressoides, which

has shoots of rounder cross-section and longer

decussate leaves. The leaves are apparently

arranged in opposite pairs, as in Libocedrus and

NS PLEDGE

cee gauamcwmmmenecemmeras

—

FIGURE 5. Female cones of Widdringtonia

cupressoides (L.) Endlicher, specimens from the South

Australian Herbarium collections.

Papuacedrus (Hill & Carpenter 1989).

Unfortunately, no other specimens from this site

are available to support or deny this association,

and the locality has reportedly been stripped by

commercial interests for decorative stone.

CONCLUSIONS

It is apparent that these impressions represent

the fruiting bodies of a previously unknown

Australian species of conifer, and it is suggested

that this shows closest affinities with species of

Widdringtonia, Diselma and Papuacedrus.

Without corroborative evidence in the form of

organically associated foliage and pollen, it is not

possible to specify which taxon the cones

represent; no identification is proposed, but it

could be a new extinct genus. Widdringtonia

today inhabits seasonally dry to semi-arid regions

of southern Africa (Hill & Brodribb 1999), as do

Callitris spp. Such an environment is implied

(Alley et al 1996) for part of the Eyre Formation.

In view of the fact that Offler (1969) found no

CUPRESSACEAN CONES IN THE EOCENE EYRE FORMATION 19]

trace of either Callitris or Actinostrobus foliage in

the ‘silcrete flora’ of the Woomera area, that no

cones of Callitris have subsequently been found

in the large collections made since her study, and

that there have been only a few finds of fossil

Callitris foliage in southeastern Australia (Hill &

Brodribb 1999), it is suggested that these typical

Australian genera evolved subsequently, possibly

from a form like the fossil taxon described here.

ACKNOWLEDGMENTS

I thank Ms L Marsh and Mr T Nurenberg for

donating the specimen to the South Australian Museum;

Ms H Vonow, South Australian Herbarium, who

assisted with access to preserved dry conifer specimens;

and Prof. R Hill and a reviewer who read an early draft

and provided some very useful references.

REFERENCES

Alley, NF, Krieg, GW & Callen, RA. 1996. Early Eyre

Formation, lower Nelly Creek, southern Lake Eyre

Basin, Australia: palynological dating of macrofloras

and silcrete, and palaeoclimatic interpretations.

Australian Journal of Earth Sciences 43: 71-84.

Alley, NF & Pledge, NS. 2000. The plants, animals and

environments of the last 280 (290) million years. Part

2 in WJH Slaytor (ed). ‘Lake Eyre South’. Royal

Geographical Society of South Australia, Lake Eyre

Monograph Series 5: 35-82.

Ambrose, GJ, Callen, RA, Flint, RB & Lange, RT.

1979. Eucalyptus fruit in stratigraphic context in

Australia. Nature 280 (5721): 387-389.

Baker, RT & Smith, HG. 1910. ‘A Research on the

Pines of Australia’. Government Printer: Sydney.

Callen, RA & Cowley, WM. 1995. Billa Kalina Basin.

In JF Drexel, WV Preiss & AJ Parker (eds). ‘The

geology of South Australia’, vol. 2, The Phanerozoic,

pp. 195-197. Geological Survey of South Australia,

Bulletin 54.

Callen, RA & Lange, RT. 1986. Billa Kalina Basin. Jn

“The Lake Eyre Basin — Cainozoic Sediments. Fossil

Vertebrates and Plants, Landforms, Silcretes and

Climatic Implications.’ Australasian Sedimentology

Group Field Guide Series No 4. Geological Society

of Australia: Sydney.

Chapman, F. 1937. Descriptions of Tertiary plant

remains from central Australia and from other

Australian localities. Transactions of the Royal

Society of South Australia. 61: 1-16.

Christophel, D, Scriven, LJ & Greenwood, DK. 1992.

An Eocene megafossil flora from Nelly Creek, South

Australia. Transactions of the Royal Society of South

Australia 116(2): 62-76.

Clifford, HT & Constantine, J. 1980. Ferns, Fern Allies

& Conifers of Australia. University of Queensland

Press: Brisbane.

Gadek, PA, Alpers, DL, Heslewood, MM & Quinn, CJ.

2000. Relationships within Cupressaceae sensu lato:

A combined morphological and molecular approach.

American Journal of Botany 87: 1044-1057.

Greenwood, DR, Callen, RA & Alley, NF. 1990. The

correlation and depositional environment of Tertiary

strata based on macrofloras in the southern Lake

Eyre Basin, South Australia. South Australian

Department of Mines and Energy, Report Book 90/

15.

Greenwood, D, Haines, PW & Steart, DC. 2001. New

species of Banksieaeformis and a Banksia ‘cone’

(Proteaceae) from the Tertiary of central Australia.

Australian Systematic Botany 14: 871-890.

Hill, RS & Brodribb, TJ. 1999. Southern conifers in

time and space. Australian Journal of Botany 47:

639-696.

Hill, RS & Carpenter, RJ. 1989. Tertiary gymnosperms

from Tasmania: Cupressaceae. Alcheringa 13: 89-

102.

Lange, RT. 1978. Carpological evidence for fossil

Eucalyptus and other Leptospermae (subfamily

Leptospermoideae of Myrtaceae) from a Tertiary

deposit in the South Australian arid zone. Australian

Journal of Botany 26: 221-233.

McIver, EE. 2001. Cretaceous Widdringtonia Endl.

(Cupressaceae) from North America. International

Journal of Plant Science 162(4): 937-961.

Nunn, CE. 1964. The plausibly coniferous shoots from

the Mt Eba-Haggard Hill-Jack’s Hill fossil plant

assemblages of South Australia. Unpublished BSc

(Hons) thesis, University of Adelaide.

Offler, CE. 1969. The external morphology of extant

and fossil vegetative shoots as a basis for

palaeobotanical studies (exemplified by a study of

silicified Tertiary floras from central South

Australia). Unpublished PhD thesis, University of

Adelaide.

Offler, CE. 1984. Extant and fossil Coniferales of

Australia and New Guinea. Part 1: a study of the

external morphology of the vegetative shoots of the

extant species. Palaeontographica Abteilung B:

Palaeophytologie 193: 18-120.

Rowett, A. 1997. Earthwatch °96. MESA Journal 5: 27-

29.

Wopfner, H, Callen, RA & Harris, WK. 1974. The

Lower Tertiary Eyre Formation of the southwestern

Great Artesian Basin. Journal of the Geological

Society of Australia, 21(1): 17-51.

DESCRIPTIONS OF A NEW GENUS AND TWO NEW SPECIES OF

VIVIPARID SNAILS (MOLLUSCA: GASTROPODA: VIVIPARIDAE) FROM

THE EARLY CRETACEOUS (MIDDLE-LATE ALBIAN) GRIMAN CREEK

FORMATION OF LIGHTNING RIDGE, NORTHERN NEW SOUTH WALES

ROBERT J. HAMILTON-BRUCE, BRIAN J. SMITH & KAREN L. GOWLETT-HOLMES

Summary

Fossil gastropods belonging to the family Viviparidae (Caenogastropoda (Prosobranchia)) are

described from the Early Cretaceous (middle-late Albian) Griman Creek Formation of Lightning

Ridge, northern New South Wales. From our research, this is the earliest definitive record of non-

marine gastropods from Australia and among the oldest viviparid material in the world recorded to

date. A new genus and two new species are described, and other material attributable to the extant

genus Notopal is recorded. Implications for the current distribution of the Viviparidae are

addressed.

DESCRIPTIONS OF A NEW GENUS AND TWO NEW SPECIES OF VIVIPARID SNAILS

(MOLLUSCA: GASTROPODA: VIVIPARIDAE) FROM THE EARLY CRETACEOUS

(MIDDLE-LATE ALBIAN) GRIMAN CREEK FORMATION OF LIGHTNING RIDGE,

NORTHERN NEW SOUTH WALES

ROBERT J HAMILTON-BRUCE, BRIAN J SMITH & KAREN L GOWLETT-HOLMES

HAMILTON-BRUCE, RJ, SMITH, BJ & GOWLETT-HOLMES, KL. 2002. Descriptions of a

new genus and two new species of viviparid snails (Mollusca: Gastropoda: Viviparidae) from

the Early Cretaceous (middle-late Albian) Griman Creek Formation of Lightning Ridge,

northern New South Wales. Records of the South Australian Museum 35(2): 193-203.

Fossil gastropods belonging to the family Viviparidae (Caenogastropoda (Prosobranchia))

are described from the Early Cretaceous (middle—late Albian) Griman Creek Formation of

Lightning Ridge, northern New South Wales. From our research, this is the earliest definitive

record of non-marine gastropods from Australia and amongst the oldest viviparid material in

the world recorded to date. A new genus and two new species are described, and other material

attributable to the extant genus Notopala is recorded. Implications for the current distribution

of the Viviparidae are addressed.

*Robert J Hamilton-Bruce, South Australian Museum, North Terrace, Adelaide, South

Australia 5000; Brian J Smith, Queen Victoria Museum, Wellington Street, Launceston,

Tasmania 7250, Australia; Karen L Gowlett-Holmes, CSIRO Division of Marine Research,

GPO Box 1538, Hobart, Tasmania 7001, Australia; *corresponding author — email: hamilton-

bruce.robert@ saugov.sa.gov.au, Manuscript received 29 July 2002.

Opalised fossils have long been known from

the Lower Cretaceous (middle—late Albian)

freshwater deposits of the Griman Creek

Formation, Lightning Ridge, New South Wales

(Smith 1999). To date, much of the published

work has focused on fish (Kemp 1991; Kemp &

Molnar 1981) and terrestrial tetrapod remains

(Archer et al 1985; Flannery et al 1995; Molnar

1980a, 1980b, 1991; Molnar & Galton 1986; Rich

et al 1989) with relatively few reports (Dettman et

al 1992; McMichael 1957; Smith 1999)

summarising the vast quantities of invertebrate

material recovered from the area. The recent

opportunity to examine Lightning Ridge

gastropod specimens (brought to our attention by

Ben Kear, South Australian Museum, as part of

an ongoing assessment of vertebrate and

invertebrate fossil material from the locality) from

the collections of the Australian Museum, Sydney,

and several private individuals has prompted a

systematic appraisal of the gastropods within the

material recovered. This study describes

specimens attributable to the family Viviparidae

(Caenogastropoda (= Prosobranchia)), including a

new genus and two new species. Indeterminate

material belonging to the extant genus Notopala

is also recorded, and the implications for

distribution of extant viviparid taxa are discussed.

The Viviparidae is a cosmopolitan group of

freshwater caenogastropods, characterised by

medium- to large-sized turbiniform shells which

possess a rounded body whorl; moderately high,

pointed spire; wide, round aperture; and sub-

spiral, horny operculum (Smith 1992). The current

Australian distribution of the group is limited to a

few species occurring in the large drainage basins

that span much of the arid centre and northern

tropical regions of the continent.

The fossil record for Viviparidae is known from

the Jurassic to Recent (Viviparus Montfort, 1810),

with a tentative report based on an internal shell

(?Bernicia Cox, 1927) mould (possibly of marine

origin) from the Lower Carboniferous of England

(Brookes-Knight et al 1960). Within Australia

there are few records of fossil viviparids.

Etheridge (1902) described the earliest potential

Australian taxon, Viviparus (?) alba-scopularis

Etheridge (also noted by Newton (1915)) from the

Aptian marine deposits of the Doncaster Member

(Wallumbilla Formation sensu Burton & Mason

1998), White Cliffs, New South Wales. This

specimen is currently under examination by the

authors and at present is regarded as being of

unclear viviparid affinity. Cotton (1935a) erected

194

a species of Notopala (N. wanjacalda) from late

Pleistocene sediments along the Murray River

near Sunnyside, South Australia, and also noted a

second taxon (Notopala sp.) from the same

deposits, which showed strong similarity to the

extant N. hanleyi (Frauenfeld, 1862). Viviparid

snail shells from Early Cretaceous deposits in the

Lightning Ridge area were recorded but not

described by Dettman et al (1992) and Smith

(1999); the latter also recorded possible

representatives of the Naticidae, Thiaridae and

Ellobiidae. Few other Australian non-marine

gastropod fossils (all of Tertiary age) have been

recorded (Archer et al 1994; Arena, 1997;

Chapman 1937; McMichael 1968).

MATERIALS AND METHODS

All specimens described herein are derived

from the Lightning Ridge opal fields (exact mine

localities from which these specimens originated

are unknown), Surat Basin, northwestern New

South Wales. The opal-bearing sediments in this

area form part of the Griman Creek Formation, a

unit dated as middle—late Albian in age (Dettman

et al 1992). The deposit predominantly reflects a

coastal plain facies (Burger 1988; Dettman et al

1992) situated in an Early Cretaceous high latitude

zone (~70° S, Embleton 1984). Invertebrate

fossils, plant root impressions and vertebrate

taphonomy suggest a freshwater estuarine to

lacustrine setting (Dettman et al 1992; Molnar

1980a). Palaeoclimatic indicators imply strongly

seasonal conditions with sea-level isotopic

palaeotemperatures in the Eromanga Basin / Surat

Basin regions ranging from 11.9°C (northeast) to

16.3°C (southwest) (Dettman et al 1992; Stevens

& Clayton 1971).

Designation of parietal fold position on the

specimens was achieved by positioning the fold

within the aperture on a 360° compass setting

when a vertical line through the axis, juxtaposed

to the column, is intersected by a horizontal line

corresponding to the midpoint of the aperture.

The adapical axis above the point of intersection

is taken as zero degrees. The diameters of all

shells were measured following the method of

Boycott (1928) and are defined as ‘the greatest

dimension that can be found starting with the

edge of the lip to a point on the opposite side of

the shell on the last whorl’. To enable

extrapolation of the numbers of missing whorls,

the incremental angle of the shells was judged

by drawing a line so as to touch each of the

RJ HAMILTON-BRUCE, BJ SMITH, KL GOWLETT-HOLMES

present whorls across each side of the shell; the

point at which they met was assessed as being

approximately the original apex of the shell (Cox

1960). All specimens are deposited in the

Australian Museum (AM), Sydney. All

specimens were prepared using a diamond drill-

bit attached via a flexible lead to a high-speed

rotary motor. Shell measurements were made to

the nearest 0.05 mm using dial calipers.

SYSTEMATICS

Class GASTROPODA

Subclass ORTHOGASTROPODA

Superorder CAENOGASTROPODA

Order ARCHITAENIOGLOSSA

Superfamily AMPULLARIOIDEA

Family VIVIPARIDAE Gray, 1847

Diagnosis

Medium to large dextral, turbiniform shells,

body whorl rounded, spire moderately high,

pointed; aperture wide, round, parietal fold

present or absent; operculum horny, subspiral

(modified after Smith 1992).

Remarks

The above diagnosis follows Smith (1992),

modified to accommodate the presence of a

parietal fold in the new genus described below.

Viviparid snails are, as their name suggests,

viviparous (live bearing) and are found in both

lotic and lentic systems throughout the world

(Browne 1978). Within Australia the family is

currently represented by the extant native genera

Notopala, Larina and Centropala (Smith 1992).

A species accidentally introduced from Asia,

Bellamya heudei guangdungensis (Kobelt, 1906),

recorded by Shea (1994) as established in New

South Wales, is not considered part of the

Australian fauna in this study. The Australian

members of the family have undergone substantial

revision Over many years (Cotton 1935a, 1935b;

Sheldon & Walker 1993; Stoddart 1982); the use

of intraspecific shell variation and morphométric

data by recent authors has resulted in a substantial

reduction in the numbers of accepted species.

Shell colouring and pattern and the form of the

operculum, which are important characters in

EARLY CRETACEOUS VIVIPARID SNAILS 195

determining generic placement in this family, are

characters lost in most fossils.

The present specimens from Lightning Ridge

are over 100 million years old (Albian [108—

97.5 Ma]) and totally opalised. While sculpturing

on some specimens has been preserved, colour

bands and opercula have not; therefore, only

structural shell characters could be used to assign

them to taxa. Two major factors have been taken

into account before deciding to place them in

Viviparidae — the shell morphology, which fits

within the currently accepted diagnosis of the

family with only slight modification (the presence

of the parietal fold in the two new species); and

the freshwater depositional environment from

which they originated. Whilst all the specimens

can be accommodated within the family

Viviparidae, those possessing a parietal fold

cannot be assigned to any currently recognised

genus and are considered to belong to a new

genus which is described below. Direct

comparisons between species, living or fossil, are

few (Brown 1980); however, due to the lack of

colour patterns and opercula as well as soft parts

in these fossils, the only remaining methods are

based on morphology. Therefore, we have applied

parts of the morphometric data gathered by

Sheldon and Walker (1993) to justify the

placement of these specimens within Viviparidae,

and in one case in the extant genus Notopala, and

to offer a method of comparison between extant

and extinct species. None of the data comparisons

are in any way intended to suggest relationships

between members of either the Notopala sp.

presented here or the new genus and any extant

species.

Genus Albianopalin gen. nov.

Diagnosis

Shell dextral, thick, solid, globose to

subglobose, ventricose, three-five whorls,

subumbilicate; aperture subovate, large,

approximately equal to height of spire; parietal

fold present, simple. Operculum unknown.

Type species

Albianopalin benkeari sp. nov.

Etymology

From the combination of Albian, n. referring to

age of the Lower Cretaceous opal-bearing deposits

of the Griman Creek Formation, and opalin,

Middle English, from opalus, Latin, alteration of

Greek opallios, for ‘opal’.

Remarks

While this new genus has similarities to

Notopala (Cotton, 1935b), Albianopalin gen. nov.

is easily separated from Notopala and all other

genera within the family by the presence of a

parietal fold in the basal lip, currently a feature

unique to this genus within the family.

FIGURE 1. AMF122185 Albanopalin benkeari sp. nov. in A, apertural and B, dorsal views.

196

FIGURE 2. AMF122185 Albanopalin benkeari sp. nov.

in apertural view showing angle of parietal fold.

Albianopalin benkeari sp. nov. (Figs 1, 2,

Appendix)

Diagnosis

With the features of the genus; parietal fold

simple, 70° from vertical axis.

Type specimen

Holotype: AMF122185 (Figs 1, 2, Appendix)

Locality and horizon

Lightning Ridge opal fields (precise site within

the opal fields unknown), northern New South

Wales, Surat Basin, Griman Creek Formation,

middle—late Albian.

Description

Shell 17.7 mm high, 16.4 mm maximum

diameter, dextral, turbiniform, subglobose.

Teleconch with three complete whorls and broken

parts, pointing to the possible presence of further

whorls. Whorls impressed. Relatively evenly

spaced spiral prosocyrt ornamentation present on

many areas of teleconch. Aperture large

(10.85 mm high), round; basal outer lip varicose

with no evidence of eversion. A single,

triangulated parietal fold (3.0mm long, 1.0 mm

wide, 2.0 mm deep) present on upper surface of

basal lip, deepest side facing adaperturally (Fig.

2);

RJ HAMILTON-BRUCE, BJ SMITH, KL GOWLETT-HOLMES

Etymology

Named for Ben Kear, for his assistance in this

study.

Remarks

Holotype unique. The position of the parietal

fold separates A. benkeari sp. nov. from the

other new species described below. The

holotype specimen is undistorted, but the spire

is incomplete. Extrapolation from the spire

angle indicates that there may have been at least

four and as many as five complete whorls

originally. Being opalised, the state of

preservation of the holotype is very good, with

much of its detailed surface ornamentation

preserved.

Albianopalin lizsmithae sp. nov. (Figs 3, 4,

Appendix)

Diagnosis

With the features of the genus; parietal fold

simple, 112° from the vertical.

Type specimen

Holotype: AMF122186 (Figs 3, 4, Appendix)

Locality and horizon

Lightning Ridge opal fields (precise site within

the opal fields unknown), northern New South

Wales, Surat Basin, Griman Creek Formation,

middle—late Albian.

Description

Shell, 15.1 mm high, 12.9 mm maximum

diameter, dextral, turbiniform, subglobose.

Teleconch with two complete whorls and broken

parts, pointing to the possible presence of further

whorls. Whorls impressed. This particular

specimen has undergone some dorsoventral

compression which has resulted in distortion,

especially at base of final whorl. Aperture large

(9.95 mm high), round; basal outer lip varicose

with no evidence of eversion. A single, basal

parietal fold (3.75 mm long, 2.3 mm wide,

2.2 mm deep) is present on surface of basal lip,

deepest side facing adaperturally (Fig. 4). Spiral

prosocyrt ornamentation present on parts of

teleconch, especially proximal to aperture.

Etymology

We name this species for our colleague

Elizabeth (Liz) Smith of the Australian Museum

(AM), for her assistance in this study.

EARLY CRETACEOUS VIVIPARID SNAILS 197

FIGURE 3. AMF122186 Albanopalin lizsmithae sp. nov. in A, apertural and B, dorsal views.

FIGURE 4. AMF122186 Albanopalin lizsmithae sp.

noy. in apertural view showing angle of parietal fold.

Remarks

Holotype unique. The position of the parietal

fold separates A. lizsmithae sp. nov. from A.

benkeari sp. nov. as described above. The

holotype has undergone moderate dorsoventral

compression, resulting in the specimen appearing

stretched. Like A. benkeari sp. nov., the spire is

incomplete, and extrapolation from the spire angle

indicates that there originally may have been at

least four and as many as five complete whorls.

Being opalised and having damage to parts of the

surface of the teleconch, the state of preservation

is good enough to preserve some of its detailed

surface ornamentation.

Genus Notopala (Cotton, 1935b)

Type specimen

Type species: Paludina hanleyi (Frauenfeld,

1864) by original designation.

Diagnosis

Shell dextral, globose—-conic, subumbilicate,

five whorls, ventricose to angulate below the

periphery; aperture subovate, large, about equal to

height of spire; parietal fold absent; operculum

corneous.

198 RJ HAMILTON-BRUCE, BJ SMITH, KL GOWLETT-HOLMES

FIGURE 5. AMF13001 Notopala sp. in A, apertural and B, dorsal views.

Remarks

These specimens are extremely significant.

Until their discovery, none of the currently living

native Australian viviparid genera were known

from deposits older than Plio-Pleistocene. This

temporal range is now extended back to the Lower

Cretaceous (middle—late Albian). It is also

interesting to note that another living species, the

lungfish Neoceratodus forsteri, has been recorded

from the Griman Creek Formation of Lightning

Ridge (Kemp & Molnar 1981).

Notopala sp. (Fig. 5, Appendix)

Referred material

AMF13001 (Fig. 5), AMF122166—-AMF 122184

(not figured herein).

Locality and horizon

Lightning Ridge opal fields (precise site within

the opal fields unknown), northern New South

Wales, Surat Basin, Griman Creek Formation,

middle-late Albian.

Description

Shell dextral, turbiniform, subglobose.

Teleconch with two to three complete

impressed whorls; the incremental angle of the

shell indicates that there were originally four or

five whorls. Aperture large, round; basal outer

lip varicose with no evidence of eversion.

Remarks

There is considerable variation in size, state of

preservation, degree of corrosion and/or damage to

the shells, opalisation, amount of matrix deposition

on the shells, completeness of the aperture and

presence/absence of ornamentation. We have not

described these specimens as a new species at this

stage. We feel that the exact taxonomic position of

this material compared with other species in the

genus requires further study. The preceding table

(Table 1), diagram (Fig. 6, modified from Sheldon

& Walker 1993) and graph (Fig. 7) show

measurements of shell characters for each of the

living Australian species of Notopala, and illustrate

morphometric similarity of the specimens of

Albianopalin nov. sp. as well as Notopala sp.

described here to existing members of the genus.

However, the specimens of Notopala sp. vary

greatly in their physical condition, and it has not

been possible to take the full range of

measurements from each of the 20 specimens being

studied. The reason for the inclusion of this data is

not to demonstrate any relationship between extant

species and the specimens from Lightning Ridge,

but to show that the physical characteristics of the

new material (Albianopalin sp. nov. and Notopala

sp.) fit within the parameters for inclusion in the

family Viviparidae.

EARLY CRETACEOUS VIVIPARID SNAILS 199

Umbilical

width

(UMB)

Aperture

width

(APW)

Shell

width

(SHW)

FIGURE 6. Diagram illustrating measurement parameters used in the morphometric analysis (modified from

Sheldon & Walker 1993).

APL SHW UMB APW ALL

Pos 7 ee a

3.0

2.0

1.0

4.00 6) o ® 6 8 6.

0.0

N. waterhousii

[_] Notopala sp.. gen.

Paka Sp.. ¥ He N. essingtonensis

"Banded" Shells

E= Notopala sublineata (Cooper) tes, alisoni ' .

Ul) Notopala sublineata (Darling) Wl 4. benkeari nov. sp.

ENN. hanleyi Wl 4. lizsmithae nov. sp.*

| = Standard deviation * single sample

FIGURE 7. Histogram showing results of morphometric analysis. Taxa include living species of Notopala (modified

from Sheldon & Walker 1993), Notopala sp. (AMF13001, AMF122185-122184), Albianopalin benkeari

(AMF122185) and A. lizsmithae (AMF122186).

200

RJ HAMILTON-BRUCE, BJ SMITH, KL GOWLETT-HOLMES

TABLE 1. Mean shell measurements (mm) for various living species of Notopala (*modified from Sheldon &

Walker 1993) compared with those for Notopala sp. (AMF13001, AMF122185-—122184), Albianopalin benkeari

(AMF122185) and A. lizsmithae (AMF122186). Because not all specimens of Notopala sp. were complete, the

number (n) of specimens used for each individual parameter is indicated separately. Standard deviation (SD) rows

were not included for A. benkeari and A. lizsmithae as only a single specimen has currently been recovered for each

of these taxa. Abbreviations: SHL, shell length; APL, apertural length; SHW, shell width; UMB, umbilical width;

APW, aperture width; SPL, spire length; ALL, aperture lip length.

Species Number SHL APL SHW UMB APW SPL ALL

*N. alisoni 20 21.59 11.91 16.3 02.59 09.77 14.78 05.35

(Brazier, 1979) SD 0.086 0.039 0.054 0.014 0.034 0.064 0.015

*N. essingionensis 20 21.70 = 12.00 16.58 1.96 10.10 14.47 6.11

(Frauenfeld, 1862) SD 0.082 0.038 0.067 0.016 0.038 0.067 0.025

*N. waterhousii 10 38.59 22.84 33.14 4.63 18.88 25.01 10.02

(Adams & Angus, 1864) SD 0.209 0.105 0.164 0.024 0.087 0.131 0.043

*N. hanleyi 42 16.67 10.50 13.83 1.29 8.52 11.00 5.57

(Frauenfeld, 1862) SD 0.520 0.026 0.036 0.00 0.025 0.037 0.013

*N. sublineata 28 18.32 10.84 14.53 2.05 9.21 12.04 5.24

(D) (Conrad 1850) SD 0.063 0.024 0.040 0.012 0.026 0.039 0.016

*N. sublineata 35 19.46 11.06 15.52 2.21 8.89 12.59 4.93

*‘Banded’ shells 15 22.15 13.18 18.10 2.06 10.69 14.6 6.42

SD 0.110 0.053 0.008 0.011 0.045 0.079 0.029

Notopala sp. 20 - 10.35 (19) 14.43 (20) 3.24 (10) 9.20(15) - 4.71 (9)

SD - 2.563 3.053 0.616 1.80 - 1.071

A. benkeari 1 17.7 10.85 16.4 3.35 9.85 - 6.2

A. lizsmithae 1 1521 9.95 12.9 3.4 8.15 - 4.4

DISCUSSION climatic conditions which characterised many

Albianopalin benkeari sp. nov., A. lizsmithae

sp. nov. and Notopala sp. share the distinction of

being the three oldest known definitively assigned

members of the Viviparidae in Australia and thus

serve to extend the range of the family in this

region back to at least the uppermost Early

Cretaceous. Since the family is confidently known

from the Jurassic to Recent in Europe, Viviparidae

therefore appears to be an ancient pre-Jurassic

group of probable Pangean origin. By the

Cretaceous, the family had diversified within the

Gondwanan region into a range of endemic genera

and species.

While there are numerous Cretaceous records

of freshwater bivalves from Australia (Dettman et

al 1992; Hocknull 1997; Jell & Duncan 1986;

Ludbrook 1985; McMichael 1957), there are very

few for non-marine gastropods from the same

period. The reasons for this apparent absence are

unknown, but could be related to preservational

biases (with shells rapidly breaking up or

dissolving after death). Another possibly reason

could be a lack of gastropod species due to poor

tolerance of the strongly seasonal near-freezing

Australian high latitude depositional environments

during the Early Cretaceous.

The implications of this study, combined with

subsequent investigations, could eventually

demonstrate a Gondwanan and _ possibly

Australian radiation for many of the species living

around the globe today.

ACKNOWLEDGMENTS

This paper would never have been initiated without

Ben Kear, who, since its inception, has more than

earned our sincere thanks. We wish also to thank Philip

Ryan for examining the statistical methods employed.

Greg Brown and George Koulis of the Royal Adelaide

Hospital (RAH) helped considerably by contributing

high-grade imaging. Elizabeth (Liz) Smith, Bob Jones,

Henk Godthelp, Tom Rich, Rolf Schmidt and Neville

Pledge supported this work and generously permitted

access to their collections. Philip Clarke of the South

Australian Museum, Adelaide (SAMA), gave financial

support and encouragement. Anne Hamilton-Bruce of

The Queen Elizabeth Hospital (TQEH) contributed

lateral thinking and assistance with editing. Chris Izzo

(zoology student at Adelaide University) assisted with

measuring and statistical data analysis. Origin Energy,

EARLY CRETACEOUS VIVIPARID SNAILS

The Advertiser newspaper, Coober Pedy Tourism

Association, the South Australian Museum and the

Waterhouse Club gave other support to this project.

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EARLY CRETACEOUS VIVIPARID SNAILS 203

APPENDIX

Measurements (mm) of all Lightning Ridge gastropod specimens used in this study (AMF13001,

AMF122166-AMF122186). Abbreviations: SHL, shell length; APL, apertural length; SHW, shell

width; UMB, umbilical width; APW, aperture width; SPL, spire length; ALL, aperture lip length. Spire

length measurements are omitted owing to incomplete apices in all specimens sampled.

Specimen SHL APL SHW UMB APW ALL

1. AMF122185 (type A. benkeari) 17.70 10.85 16.40 3.35 9.85 6.20

2. AMF122186 (type A. lizsmithae) 15.10 9.95 12.90 3.40 8.15 4.40

3. AMF13001 (Notopala sp.) 19.25 12.00 17.30 3.75 11.10 5.02

4. AMF122166 17.65 11.05 14.50 = 8.55 -

5. AMF122167 14.37 8.75 12.75 8.50 - -

6. AMF122168 11.60 - 11.85 2.30 7.80 3.35

7. AMF122169 15.62 8.80 12.45 3.00 - 5.00

8. AMF122170 12.60 8.80 11.65 3.00 7.65 4.35

9. AMF122171 19.00 13.25 19.90 _ 12.20 -

10. AMF122172 20.42 12.10 17.20 4.20 - 6.00

11. AMF122173 18.00 11.05 16.20 3.90 9.80 -

12. AMF122174 17.92 11.20 15.55 - 11.15 -

13. AMF122175 19.82 12.63 17.85 3.35 - a

14. AMF122176 19.35 11.10 16.75 - 10.50 -

15. AMF122177 19.57 12.65 17.50 - 11.70 -

16. AMF122178 18.55 10.05 15.15 ~ - -

17. AMF122179 17.95 11.30 15.20 3.80 9.60 4.80

18. AMF122180 10.22 6.50 9.00 - - 6.00

19. AMF122181 6.92 7.20 9.40 2.60 6.15 3.60

20. AMF122182 9.32 6.50 8.75 2:35, 5.60 3.15

21. AMF122183 17.80 11.70 16.00 - 9.80 -

. AMF122184 - 16.10 10.10 13.40 - 8.30