Memoirs

of THE

Queensland Museum

Brisbane
30 June 2003

Volume 49
Part 1

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Volume 49
Part 1

Memoirs

of THE

Queensland Museum

Minister:

Director:

Managing Editor:
Editorial Assistant:

Hon. M.J. Foley, MLA
I.D. Galloway, PhD
P.A. Jell, PhD
P. Avem, BSc

PUBLISHED BY ORDER OF THE BOARD

30 JUNE 2003

<p> Queensland Museum
PO Box 3300, South Brisbane 4101, Australia
Phone 61 7 3840 7555
Fax 61 7 3846 1226

National Library of Australia card number
ISSN 0079-8835

NOTE

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A Queensland Government Project

Typeset at the Queensland Museum
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35 Hamilton Road, Moorooka, Queensland 4105

THREE NEW ENDEMIC GENERA OF THE ASTERON-C OMPLEX (ARANEAE:
ZODARIIDAE) FROM AUSTRALIA:

BASASTERON, EUASTERON AND SPINASTERON

BARBARA BAEHR

Baehr, B. 2003 06 30: Three new endemic genera of the Asteron-compXex (Araneae:
Zodariidae) from Australia: Basasterott , Euasteron and Spinasteron. Memoirs of the
Queensland Museum 49(1): 1-27. Brisbane. ISSN 0079-8835.

Three new genera are created for 19 new species in the -complex, a large group of

Australian Zodariidae. Basasteron gen. nov. contains only Basasterott leucosemum
(Rainbow', 1 920) and is characterised by the straight retrolateral cymbial flange and the very
short ventral and dorsal tibial apophysis. Euasteron gen. nov. includes the type species E.
enterprise sp. nov. (6,9) and E. atriceps sp. nov. ( 6 ), E. bartoni sp. nov. (d, 9), E.
Carnarvon sp. nov. (6 ,9). E. churchillae sp. nov. (6),E. gibsonae sp. nov. (6), E. harveyi
sp. nov. (6 ), E.johannae sp. nov. (6, 9 ), E.juliannae sp. nov. ( 6 , 9 ), E. krebsorum sp. nov.
( 6 , 9 ), E. lornc sp. nov. ( 6 ), E. milledgei sp. nov. (d , 9 ), E, monteithorum sp. nov. ( 6 , 9 ), E.
raveni sp. nov. (d), E. ulrichi sp. nov. (d), E. ursulae sp. nov. (d,9), E. willeroo sp. nov.
(d,9) and can be recognised by cymbial flange with rounded extension. Spinasteron gen.
nov. is identified by barely developed retrolateral cymbial flange and represented by S.
sanford sp. nov. (d ) and S. nigriceps sp. nov. (d ). CJ Basasteron, Euasteron, Spinasteron,
Australia , new genera.

Barbara Baehr, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia
(e-mail: BarbaraB@qm.qld.gov.au); 20 October 2002.

This is the fourth systematic paper, funded by
the Australian Biological Resources Study
Participatory Program, on the large Asteron-
complex (Baehr & Jocque, 1 996). Asteron Jocque
1991, has been revised (Jocque & Baehr, 2001 )
and contains 8 species. From the different characters
of all other species new genera were erected. So
far, Cavasteron, Leptasteron , Minasteron ,
Pentasteron , Phenasteron Pseudasteron and
Subasteron ( Baehr & Jocque, 2000, 200 1 ; Jocque
& Baehr, 2001) have been revised. This paper
contains another 3 new genera with 19 new
species. Basasteron shares the straight cymbial
flange with the basic genera Pentasteron ,
Phenasteron and Leptasteron of the
/L?/mw-complex and the semicircular DTA with
folded distal part with the more derived genera
Cavasteron , Minasteron , Euasteron and
Spinasteron.

Euasteron and Spinasteron are closely related
to Cavasteron and Minasteron sharing the special
shape of tibial apophyses with 2 apophyses
forming a deep concavity between them: the
dorsal one divided in two parts connected by a
ridge and the typical membranous semicircular
DTA with folded distal part. Euasteron can be
easily separated from both genera by the lack of a
large terminal claw-like spine in male palps.
Spinasteron is separated from these genera by the
barely developed retrolateral cymbial flange

(Fig. 4A) on the palpal cymbium in contrast to
flange with a well-developed extension (Fig. 4C).

Most species are named in honour of people
who supported this paper.

MATERIAL AND METHODS

Descriptions are generated w ith the aid of Intkey
(Dallwitz et al., 1 998) and shortened where possible.

Drawings were done from body, right palp,
epigyne and vulva. All measurements are in mm.

Detailed spination of legs were not done because
there are normally no characters to separate the
species. Obvious spination characters are men-
tioned w'hen they are important for separation of
species.

Colour pattern is described from preserved
spiders in alcohol. The white abdominal patches
can be yellow or orange in live spiders. If the
specimens are not well preserved they can loose
their colour pattern completely. That happened
mainly both in specimens stored in vials with
w'hite lids and in specimens that were collected in
long-term pitfall traps using ethylene glycol. In
these cases, the colour pattern is useless and the
species can be identified only by their genitalic
characters.

Abbreviations of characters: ALE, anterior
lateral eyes; AME, anterior median eyes; AS,
anterior spinnerets; CD, copulatory duct; CO,

2

MEMOIRS OF THE QUEENSLAND MUSEUM

copulatory opening; Ceph, cephalo-
thorax; DTA, dorsal tegular apophysis;

Dti A dorsal tibial apophysis; E, embolus;

EP, external prong on dorso-retrolateral
tibial apophysis; F, flange on cymbium;

FA, frontal appendage of embolus base =

VTA (ventral tegular apophysis); Fe,
femur; IP, internal prong on dorso-
retrolateral tibial apophysis; MS, median
spinnerets; MEG, median eye group;

ML. membranous lamella on retrolateral
part of DTA; MT, metatarsus; P, patella;

PE, prolateral extension of DTA; PR,
prong on retrolateral side of DTA; PLE,
posterior lateral eyes; PME, posterior
median eyes; PS, posterior spinnerets;

RE, rounded extension on flange; S,
spermatheca; sl/sw sternum length/
sternum width; SP, sperm duct; T, tibia;

Ta. tarsus; TBE, Trans basal area of
embolus; TF, transparent field on
embolus base; VtiA, ventral tibial
apophysis.

Abbreviations of institutions where
material was borrowed: AM, Australian
Museum, Sydney (M. Gray); CS1RO, (T.
Churchill); MAGNT, Museum & Art
Gallery of the Northern Territory, Darwin; QM,
Queensland Museum. Brisbane (R. Raven);
SAMA, South Australian Museum, Adelaide (D.
Hirst); VM, Melbourne Museum, Melbourne (C.
McPhee); WAM, Western Australian Museum,
Perth-(M. Harvey).

SYSTEMAT1CS

KEY TO BASASTERON , EUASTERON AND
SPINASTERON

1. Palp, cymbial flange, barely developed, AME big (Figs

2A-C, 3C,F) .2

Palp, cymbial flange, well developed either straight or
with rounded extension RE (Figs 3D,F, 4C) 3

2. Carapace dark brown, SP only partly crossing TF (Figs

3C,D) Spinas teron sanford

Carapace yellow and dark cephalic area. SP crossing TF
(Figs 2A-C, 3E,F) Spinas teron nigriceps

3. Cymbium with straight retrolateral flange, DTA short ,

AME small (Figs I. v\. Hi

Basasteron leucosemum ( Rainbow, 1 920)

Cymbial flange with rounded extension, DTA long, AME
small, big or equal ( Figs 4C.5A-C.6A-C). . Euasteron

Basasteron gen. nov.

TYPE SPECIES. Basasteron leucosemum (Rainbow, 1 920).

ETYMOLOGY. Basasteron refers to the fact that this
genus is a link between the basic genera Pentasteron,
Phentasteron , Leptasteron and Euasteron.

REMARKS. Jocquc ( 1 995) tentatively added this
species to Nostera. Because of the absence of the
males he had no clue to which genus the species
belonged. The newly described males from
recently collected material show that the species
belongs to the base of the Av/ero/i-complex.

DIAGNOSIS. AME smallest; recognised mainly
by characters from the male palp: tibia with short
ventrolateral and dorsolateral apophysis cymbial
flange straight, DTA short, semicircular, with
distal part folded containing embolus; prolateral
extension PE short, lying in cymbium;
retrolateral ly with membranous lamella; sperm
duct S-shaped, partly crossing transparent field.

DESCRIPTION. Medium-sized (4.0-5. 0), with
finely reticulated tegument. Carapace widest at
coxae II. Profile flat with highest point between
fovea and PME (Fig. 1 A).

Colour: carapace orange brown; sternum yellow;
chelicerae orange; maxillae and labium yellow,
distally white; abdomen dorsally sepia brown
with 2-3 pairs of white patches on anterior part
and 1 -2 in front of spinnerets; laterally with 2 to 3

NEW AUSTRALIAN SPIDER GENERA

3

white patches; ventrally dark brown,
mottled with white; legs yellow.

Eyes (Fig. 1A-C): in three rows (2-4-2).

Only ALE in first row, AME (in the
middle) and PLE in second, third only
PME. AME smallest: eye group width
about half of headwidth. MEG slightly
longer than wide. Clypeus straight, about
3 times diameter of ALE high. Chilum
single. Chelicerae as usual for family
with a few hairs in front and dense row on
distal promargin; no teeth. Maxillae and
labium triangular; sparsely haired;
maxillae with anteromesal scopula.
Sternum flat; heart-shaped with straight
anterior margin, shiny; with long setae
around margin.

Legs: formula 4123. Spination: few
spines on pairs I, II, more numerous on
III, IV. Paired tarsal claws with numerous
(ca. 10) teeth. Unpaired claw toothless,
on very small onychium. Metatarsal
preening brush on Mt II and III poorly
developed.

Abdomen: oval; without si gill a.
Spinnerets: AS short, conical, with very
short distal segment; MS and PS tiny. Colulus
represented by group of setae. Tracheal spiracle
slit-like covered by posterior sclerotised lip with
small stout spines.

Male palp (Fig. 3A,B): tibia with short
dorsolateral and ventrolateral apophysis shallow
concavity in between. Cymbium with simple
unmodified straight flange, several spines near
distal tip. DTA semicircular, distally folded with
short prolateral extension. Embolus semicircular
conducted by DTA.

Female lectotype described in Jocque, 1995.

Basastcron Icucosenium (Rainbow, 1920)
(Figs 1,3A,B, 18)

Storena leucosemum Rainbow, 1920: 237 (description of 0
Nostera leucosemum Jocqu6, 1995: 154 (comb, nov.)

MATERIAL. New South Wales: 36. Lord Howe I., North
Bay West End, lilt, 31°33’S, 159°05’E, 19 Dccl979, GM
(QM S47393); Lord Howe I., 31°33’S, 159°05'E (AM
KS 15672); d, Lord Howe I., southern end of Salmon
Beach in vicinity of Little Island, 31°34’08”S,
159°04’28”E, 19-28 Febr2001, pitfall (AM KS82348); 9,
Lord Howe I., W slope of Dawsons Point Ridge, closed
rainforest, 3 1 °3 T 1 2”S. 1 59°02’27”E, 20 Nov 2000 (AM
KS75769); 6. Lord Howe I.. Stephens Reserve, New
Settlement, 31°3ri2”S, 159°02’27"E 4-14 Dec 2000,
pitfall (AM KS76265); 6, Lord Howe I., behind beach at
Old Gulch, W footslopes, 3 1 °30’53 ”S, 1 59°02’36”E, 2- 1 1

Dec2 000, pitfall (AM KS75879); 2 9 , Lord Howe I., Stn 9,
SE slope of Kim’s Lookout, 3I°30 , 54 ,, S, 159°03’06”E,
31 Jan 1 97 1 , M. Gray (AM KS 1 56 1 8); 2 9 , Lord Howe L,
Stn 8 summit of Kim’s Lookout, 31°30’54”S,
159°03’00”E, M. Gray 31 Jan 1971 (AM KS15617).

DIAGNOSIS. AME small, carapace orange
brown, abdomen with 3 pairs of w hite patches on
anterior part and 1-2 in front of spinnerets, palp
with straight flange, DTA short circular, distally
part folded containing embolus. Dorsolateral and
ventrolateral tibial apophysis short.

DESCRIPTION. Male (holotype). Total length
4.56; ceph 2.48 long, 1.6 w r ide, 1.04 high; cl/cw
1.55; sternum 1.12 long, 0.96 wide; sl/sw 1.17;
abdomen 2.08 long, 1 .4 w ide.

Colour: carapace orange brown; sternum yellow;
chelicerae orange; maxillae and labium yellow,
distally white; abdomen sepia brown, dorsally
with 3 pairs of w'hite patches on anterior part and
1-2 in front of spinnerets; laterally with 2-3
elongate and broad white patches; ventrally
mottled with white; legs yellow to orange-bnwn.

Eyes: in 3 rows (2-4-2); AME smallest; eye group
wddth 0.53 of headw idth; AME 0.1; ALE 0.12;
PME 0.12; PLE 0.12; AML- AML 0.04;
AME-ALE 0.04; PME-PME 0.04; PME-PLE
0.08; ALE-PLE 0.04; eye spaces AME-PME 0.3;

4

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 3. Basasteron , Spinasteron spp. right male palps. A,B, B. leucosemum\ A, ventral; B, retrolateral; C,D, S.
sanford ; C, ventral; D, retrolateral; E,F, S. nigriceps ; E, ventral; F, retrolateral. Scale 0.5mm.

AME-AME 0.24; PME-PME 0.28. Clypeus
0.32mm high.

Abdomen: tracheal spiracle small, slit-like,
covered with small stout spines.

Legs: femur I-II with 1 row, III, IV with 2 rows of
long thin setae ventrally.

Male palp (Fig. 3A,B): tibia with short
dorsolateral and ventrolateral apophysis shallow
concavity in between. Cymbium F straight; DTA
semicircular, distal part folded, prolateral
extension short, lying in cymbium, retrolaterally
with membranous lamella; SP duct S-shaped,
partly crossing TF.

Female. Lectotype described in Jocque, 1995.

Variation: minor variation in the abdominal
pattern as well as in size. Specimens from Kims
Lookout and Old Gulch are slightly smaller and
have 2 pairs of white dots on top of the abdomen.
These specimens may belong to a second species
but only further material will solve this problem.

DISTRIBUTION. Known only from Lord Howe
Island (Fig. 18).

Spinasteron gen. nov.

TYPE SPECIES. Spinasteron nigriceps sp. nov.

ETYMOLOGY. For the very long, spinuous prong or end
on palpal DTA retrolaterally.

NEW AUSTRALIAN SPIDER GENERA

5

DIAGNOSIS. Apart from the characters typical
for the Asteron- complex, representatives of
Spinasteron are recognised by AM E biggest (Fig.
2A-C), but mainly by characters from the male
palp (Figs 3C-F, 4A): cymbial flange barely
developed with small knob; tibia, Dti A with short
IP and longer EP; VtiA normally longer than EP
of DtiA. DTA semicircular, with distal part
folded containing embolus; PE retrolateral with
long spinuous end, and normally with strong
spinuous prong; SP S-shaped, partly or totally
crossing TF. Base of embolus broad, TBE thin or
broadened.

DESCRIPTION. Medium-sized (4.00-5.00), with
finely reticulated tegument. Carapace at frontal
part broad, with edges widest at coxae II. Profile
flat, with highest point between fovea and PME
(Fig. 2A-C).

Colour: carapace yellow to sepia brown; sternum
yellow to orange brown; chelicerae medium
brown; maxillae and labium medium brown,
distally white; abdomen dorsally with scutum
sepia brown; with 1 -3 pairs of white patches on
anterior part and 1 in front of spinnerets or
chevrons; legs yellow to medium brown with or
without colour pattern.

Eyes (Fig. 1A-C) in three rows (2-4-2). Only
ALE in first row, AME (in the middle) and PLE in
second, third only PME. AME biggest; eye group
width more than half of head width. Clypeus
straight, between 2.5 and 3 times diameter of
ALE high. Chilum single. Chelicerae as usual for
family, with a few hairs in front and dense row on
distal promargin; no teeth. Maxillae and labium
triangular, sparsely haired; Maxillae with
anteromesal scopula. Sternum heart-shaped, with
straight anterior margin, shiny, with long setae
around margin.

Legs: formula 4123. Spination: few spines on
pairs I, II, more numerous on III, IV. Femur I, II
with 1 row of long thin setae ventrally, Femur III,
IV with 2 rows of long thin setae ventrally.
Metatarsal preening brush on Mt II and III poorly
developed. Paired tarsal claws with numerous
(ca. 10) teeth. Unpaired claw toothless, on very
small onychium.

Abdomen oval, with scutum, without sigilla.
Spinnerets: AS short, conical, with very short
distal segment; MS and PS tiny. Colulus
represented by group of setae. Tracheal spiracle
slit-like covered by posterior sclerotised lip.
Male palp (Fig. 3C-F): Cymbial flange barely
developed, with small knob; tibia, DtiA with
short IP and longer EP. DTA semicircular, with

distal part folded, containing embolus; PE long,
not lying in cymbium, sometimes reaching tibia;
retrolateral with long spinuous end, normally
with strong spinuous prong; SP S-shaped, partly
or totally crossing TF. Base of embolus broad,
TBE with or without keel.

Female unknown.

Spinasteron sanford sp. nov.

(Figs 3C,D, 21)

ETYMOLOGY. For the type locality, Mount Sanford
Station, Northern Territory. Noun in apposition.

MATERIAL. HOLOTYPE: 6 , Mt Sanford Station,
NATL Clay site, pitfall traps, 17°18’25”S, 130°46’28 M E,
3-9 Jul 96, T. Churchill (CSIRO A763ex). PARATYPES:
Northern Territory: d, as for holotype (CSIRO A763ex);
96. same locality, Loam site, 17°18’29 M S, 130°45’17 M E,
Apr 97 (CSIRO A3496); 6 , same locality, Oct 97 (CSIRO
A 1 057); 26, same locality, 3-9 Jul 96 (CSIRO A754); 6 ,
same locality, Jul 96 (CSIRO A764). 25m, Mt Sanford
Station, NATT, Sand site, 17°2r26"S, 130°49'36"E. Apr
97 (CSIRO A3499): 3 6 , same locality, 3-9 Jul 96 (CSIRO
A756); 3d, same locality, Oct 97 (CSIRO A3502); 3d,
same locality, 3-9 Jul 96 (CSIRO A754); d, Daly R.,
Quadrat G4, PWCNT, 17°2r26"S, 130°49’36"E, Jun 99
(CSIRO A2539); d , Kidman Springs, Grazing Gradient,
Site 3, pitfall traps, 16°07’29"S, 130°56’25"E, Apr 98
(CSIRO A 1320).

DIAGNOSIS. Differs from S. nigriceps by
carapace dark brown, abdomen dark browrn with
scutum and 1 pair of white spots on posterior part
and 1 in front of spinnerets, SP S-shaped,
crossing only partly TF.

DESCRIPTION. Male (holotype). Total length
4.76; ceph 2.36 long, 1 .68 wide, 1 .04 high; cl/cw
1.4; sternum 1.12 long, 1 wide; sl/sw 1.12;
abdomen 2.4 long, 1 .6 wide.

Colour: carapace sepia brown; sternum orange
brown; chelicerae medium brown; maxillae and
labium medium brown, distally white; abdomen
sepia brown; dorsally with scutum and 1 pair of
white spots on posterior part and 1 in front of
spinnerets; legs medium brown, with darker
brown lateral stripes; with indistinct colour
pattern; coxae I-IV pale; trochanter I-IV brown;
femur I dark brown, distally white, II-IV
proximal part white, distal part dark brown.

Eyes: AME largest; eye group width 0.71 of
headwidth; AME 0.2; ALE 0. 1 8; PME 0. 1 8; PLE
0.18: AME-AME 0; AME-ALE 0.04; PME-
PME 0.06; PME-PLE 0.08; ALE-PLE 0.04; eyes
group AME-PME 0.46; AME-AME 0.4;
PME-PME 0.44. Clypeus: 0.48mm high.

6

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Euasteron , Spinasteron spp. right male palps. A, S. sanford , retrolateral; B, E. Carnarvon ventral; C,D, E.
enterprise , C, retrolateral; D, ventral. Scale 0.5mm.

Male palp (Fig. 3C,D): DTA PE reaching tibia;
SP S-shaped, partly crossing TF; DtiA IP much
shorter than EP, both with sharp tip; VtiA long,
straight, with blunt tip.

Female unknown.

DISTRIBUTION. Northern Territory, Mount
Sanford Station (Fig. 21). Semi-arid grassland

500

NEW AUSTRALIAN SPIDER GENERA

7

Colour: carapace yellow with dark
cephalic area; sternum yellow; chelicerae
medium brown; maxillae and labium
medium brown, distally white; abdomen
sepia brown; dorsally with scutum, 3
pairs of white patches on anterior part and
1 in front of spinnerets; laterally,
ventral ly pale brown; legs yellow.

Eyes: AME largest; eye group width 0.66
of headwidth; AME 0.18; ALE 0.14;
PME 0.14; PLE 0.14; AME-AME 0.02;
AME-ALE 0.02; PME-PME 0.08;
PME-PLE 0.08; ALE-PLE 0.02; eyes
group AME-PME 0.4; AME-AME 0.4;
PME-PME 0.36. Clypeus: 0.48mm high.
Male palp (Fig. 3E,F): DTA, PE reaching
tibia; retrolaterally with strong prong; SP
S-shaped, crossing transparent field
totally; DtiA IP tiny, EP short blunt; VtiA
long, straight, with rounded tip.

Female unknown.

DISTRIBUTION. Northern Territory
(Fig. 21). Tropical savanna.

Euasteron gen. nov.

FIG. 5. Euasteron enterprise. A, body dorsal, B,C,
cephalothorax; B, lateral; C, frontal. Scale 1mm.

and tropical savanna, with barley mitchell grass
(Astrebla pectinata) and Vertisols soil.

Spinasteron nigriceps sp. nov.

(Figs 3E,F, 21)

ETYMOLOGY. For the dark cephalic area. Latin
nigriceps , black-headed.

TYPE SPECIES. Euasteron enterprise sp. nov.

ETYMOLOGY. Latin, Eu = real.

DIAGNOSIS. Cymbium with retrolateral
extension or lobe. Euasteron differs from
Cavasteron and Minasteron by the lack of the
large terminal palpal claw in males. Females of
Euasteron still have this huge terminal claw with
4-6 teeth. Epigyne copulatory ducts are more
elongate and coiled than in Cavasteron and
Minasteron.

MATERIAL. HOLOTYPE: <J, Daly river, NT, Quadrat
G4, PWCNT, pitfall traps, 17°2r26"S, 130°49’36 M E, Jun
99, T. Churchill (CSIRO A2539ex). PARATYPES:
Northern Territory: <3, L10, PWCNT pitfall traps,
14°58’61”S, 132°.0r60”E, 16 Jun 99, T. Churchill
(CSIRO A3429); 6 , Douglas Daly, Sand Site, pitfall traps,
13°53*36"S, 1 3 1 ° 1 5 ’55"E, 3-9 Jul 96 (CSIRO A0332); <J,
same locality. Grid 1 (CSIRO A 177).

DIAGNOSIS. Differs from S. sanford by:
carapace yellow with dark cephalic area,
abdomen with scutum and 3 pairs of white spots
on posterior part and 1 in front of spinnerets, SP
crossing TF totally.

DESCRIPTION. Male (holotype). Total length
4.2; ccph 2.2 long, 1.68 wide, 0.96 high; cl/cw
1.31; sternum 0.96 long, 0.88 wide; sl/sw 109;
abdomen 2 long, 1.28 wide.

DESCRIPTION. Medium-sized (3.4-6.2) with
finely reticulated tegument. Carapace widest at
coxae II. Profile flat with highest point between
fovea and PME (Figs 5,6).

Colour: carapace yellow, orange or sepia brown;
abdomen pale or sepia brown with or without
scutum, with 1-3 pairs of white patches on
anterior part and 1 -3 in front of spinnerets, or with
pale or dark chevrons; legs yellow to dark brown;
clearly annulated or without colour pattern.

Eyes (Figs 5A-C, 6A-C) in three rows (2-4-2).
Only ALE in first row, AME (in the middle) and
PLE in second, third only PME. AME smallest,
equal to others or largest; eye group width
between 0.47 and 0.66 of head width. MEG
slightly longer than wide, or as long as wide.
Clypeus straight, about 3 times diameter of ALE

8

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 6. Euasteron Carnarvon. A, body dorsal, B,C,
cephalothorax; B, lateral; C, frontal. Scale 1mm.

Euasteron willeroo sp. nov.
(Figs 12A,B, 16A,B, 19)

high. Chilum single. Chelicerae as usual
for family, with a few hairs in front and
dense row on distal promargin; no teeth.
Maxillae and labium triangular; sparsely
haired; maxillae with anteromesal
scopula. Sternum heart-shaped, with
straight anterior margin, shiny; with long
setae around margin.

Legs: formula 4123. Spination: few
spines on pairs I, II, more numerous on
III, IV. Paired tarsal claws with numerous
(ca. 1 2) teeth on inner side of paired claws
(Fig. 7B). Unpaired claw toothless, on
very small onychium. Metatarsal
preening brush on Mt II and III poorly
developed. Female palp with big claw and
4-6 teeth (Fig. 7D).

Abdomen: oval, with or without scutum.
Spinnerets: AS short, conical, with very
short distal segment; MS and PS tiny.
Colulus represented by group of setae.
Tracheal spiracle slit-like, covered by
posterior sclerotised lip, with or without
small stout spines.

Male palp (Figs 8-14): Tibia short, with 2
apophyses forming deep concavity between
them, dorsally divided in two parts and connected
by a ridge; internal prong (IP) long, short or
absent; external prong (EP) long or short; ventro-
lateral tibial apophyses hooked or bird-
head-shaped. Cymbial flange with rounded
extension. DTA, semicircular, distally folded
with prolateral extension (PE) not reaching or
reaching tibia, retrolaterally, with membranous
lamella or strong prong. Embolus thin,
semicircular, conducted by DTA.

Epigyne (Figs 15-17): with 1 or 2 copulatory
openings, with U- or V-shaped sclerotised tubes;
copulatory ducts elongate, coiled, ending in small or
bigger spermathecae.

KEY TO SPECIES GROUPS OF
EUASTERON

1 . All eyes equal in size; base of embolus with extremely

long semicircular FA; DTA with strong prong, epigyne
with 2 big oval CO, not attached to epigastric fold, CO
straight elongate coiled, ending in separated sperma-
thecae. (Figs 12A,B, 16A,B) E. willeroo

AME smaller, or bigger than other eyes, FA shorter not
semicircular, epigyne different (Figs 8-11,13,14, 1 5 A-F,
16C-F, 17A-H) 2

2. AME smaller than other eyes, eye group width > 0.55

(Fig. 5A-C) E. enterprise - group

AME bigger than other eyes, eye group width < 0.55 (Fig.
6A-C) E.juliannae-group

ETYMOLOGY. For the type locality.

MATERIAL. HOLOTYPE: 6 , Willeroo, NT, NAT, Clay
site, pitfall traps, 15°ir26"S, 131°36’42 ,r E, 3-9 Jul 96, T.
Churchill (CSIRO A0343). PARATYPES: Northern
Territory: 9 , as for holotype (CSIRO A0343); 4 9 , same
locality (CSrRO A3506); V , same locality (CSIRO A98 1 );
lid, same locality, Oct 97 (CSIRO A 1 173); 3d, same
locality, Apr 97 (CSIRO A983); 7d, same locality,
15°ir26 M S, 131°36’42"E, Oct 97 (CSIRO A 1 163); 2d,
Barkly Tablelands. Hayficld, Mitchel grassland, site 3,
pitfall traps, Dec 95 (CSIRO A2116); d, Mt Sanford
Station. Grazing, VRD, Site 2, pitfall traps, 17°18 , 25"S,
130°4678 H E, Apr 98 (CSIRO A1350); 7d, Mt Sanford
Station, NAT, Clay site, 17°18’25 M S, I30°46’28"E, Oct 97
(CSIRO A3501); 4d, same locality, Apr 97 (CSIRO
A35030); 12d, same locality, Jul 96 (CSIRO A762);

1 Od , 1 9, same locality, Jul 96 (CSIRO A763); 8d, same
locality, 3-9 Jul % (CSIRO A76I ), 3d. Kidman Springs,
NAT, Clay site, pitfall traps, 16°06’39 W S, 130°56’55"E,
Oct 97 (CSIRO A 1 1 2 1 ); d , same locality, Apr 97 (CSIRO
A3504); 3d, same locality, Oct 97 (CSIRO A3505); 7d,
same locality, Jul 96 (CSIRO A752); 7d, same locality.
Grid 3, 3-9 Jul 96 (CSIRO A755).

DIAGNOSIS. Eyes equal sized. Carapace sepia
brown, abdomen sepia brown with 1 pair of white
patches on posterior part and 1 in front of
spinnerets. Palp, base of embolus with extremely

NEW AUSTRALIAN SPIDER GENERA

9

FIG 7. Euasteroiu spp. A,B,D, E. Carnarvon ; A, tracheal spiracle; B, tarsal claw 1 right leg; D, female palpal claw
retrolateral; C, E. enterprise , tracheal spiracle.

long and bent FA. With 2 big oval CO, not
attached to epigastric fold.

DESCRIPTION. Mate (holotype). Total length
3.48; ceph 1 .76 long, 1 .28 wide, 0.72 high; cl/cw
1.37; sternum 0.92 long, 0.8 wide; sl/sw 1.15;
abdomen 1 .72 long, 1 wide.

Colour: carapace, sternum, chelicerae sepia to
medium brown; maxillae and labium medium
brown, distally white; abdomen sepia brown;
dorsally with 1 pair of white spots on posterior
part and 1 in front of spinnerets; legs pale brown,
darker brown lateral stripes; with indistinct
colour pattern; coxae, trochanter I-IV pale; femur
I dark brown, II-IV proximal part white, distal
part dark brown.

Eyes: equal sized; eye group width 0.58 of
headwidth; AME 0.1; ALE 0.1; PME 0.1: PLE
0.1; AME-AME 0.02; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.08; ALE-PLE

0.02; eyes group AME-PME 0.3; AME-AME
0.22; PME-PME 0.24. Clypeus: 0.36mm high.
Male palp (Fig. 12A.B): DTA, PE reaching tibia;
retrolaterally with strong prong; VTA =FA
elongate, sickleshaped; sperm duct S-shaped, not
crossing TF; DtiA IP 3times as long as wide, EP
long sticking out; VtiA bird-head-shaped.
Female (paratype) (Fig. 16A.B). Total length 4;
ceph 2.04 lone, 1 .28 wide, 0.72 high; cl/cw 1 .59;
sternum 0.84 long. 0.8 wide; sl/sw 1 .05; abdomen
1 .96 long, 1 .28 wide.

Colour: as in males, carapace sometimes orange
brown.

Eyes: equal sized; eye group width 0.51 of
headwidth; AME 0.1; ALE 0.1; PME 0.1; PLE
0.1; AME-AME 0.02; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.3; AME-AME
0.22; PME-PME 0.24.

Clypeus 0.36mm high.

10

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. Euasteron spp. right male palps. A,B, E. enterprise ; A, ventral: B, retrolateral; C,D, E. monteithorum ; C,
ventral; D, retrolateral; E,F, E. milledgei ; E, ventral; F, retrolateral. Scale 0.5mm. DTA = dorsal tegular
apophysis; E = embolus; EP = external prong on dorso-retrolateral tibial apophysis (DtiA); FA = frontal
appendage of embolus base = VTA (ventral tegular apophysis); IPML = internal prong on DtiA; membranous
lamella on retrolateral part of DTA; PE = prolateral extension of DTA; TF - transparent field on embolus base;
VtiA = ventral tibial apophysis.

Epigyne (Fig. 16A,B): with 2 big oval CO, not

attached to epigastric fold; CD elongate, coiled

about straight axis, ending in separated S.

DISTRIBUTION. Northern Territory (Fig. 19).

Semi-arid, desert.

KEY TO SPECIES OF EUASTERON
ENTERPRISE GROUP

Males

1. Carapace orange or orange brown 2

Carapace dark brown 4

2. Abdomen laterally with 2 pairs of white spots. Palp, tibial
apophyses dorsolaterally, with external prong long

sticking out (Fig. 8C,D) E. monteithorum

Abdomen laterally with 1 white spot. Palp, tibial
apophyses dorsolaterally, with external prong short not
sticking out (Figs 9A,B, 10A,B) 3

3. Palp; DTA retrolaterally with folded, retrolateral directed

prong (Fig. 9A,B) E.lorne

Palp; DTA retrolaterally with sharp basal directed prong
(Fig. 10A.B) E.hartoni

4. Palp without or with short FA directed prolatrally on base

ofembolus(Figs8A,B, I0C,D) 5

Palp with big FAdirected distally ( Tigs 8E,F,9C,D) . . 6

5. Abdomen with 1 big pair of white patches on anterior part

and 1 elongate white stripe in front of spinnerets; palp
without FA; DTA retrolaterally with big prong (Fig.
10C,D) . • E.gibsonae

Abdomen with scutum, 2 pairs of white spots on anterior
part and 2-3 white spots in front of spinnerets; palp with
short rounded FA directed prolaterally; DTA
retrolaterally with lamella (Fig. 8A.B) . . E. enterprise

6. Palp, DTA retrolaterally with lamella and small prong;

FA big, wider than long; dorsolateral tibial apophysis
internal prong short (Fig. 8E,F) E. milledgei

NEW AUSTRALIAN SPIDER GENERA

11

Palp, DTA retrolaterally with big prong; FA big longer
than wide; dorsolateral tibial apophysis internal prong 3

times long as wide (Fig. 9C,D) E.krebsorum

Females

1. Carapace orange or orange brown 2

Carapace dark brown 3

2. Epigyne with 2 small, semicircular CO (Fig. 15C.D);

abdomen laterally with 2 pairs of white spots

E. monteithorum

Epigyne with 1 broadly oval CO and long U-shaped
sclerotised tubes (Fig. 16C,D); abdomen laterally with 1
big white spot and pale booklungs E. bartoni

3. Epigyne SP touching; CO broadly oval internal (Fig.

16E,F) . ♦ E.krebsorum

Epigyne SP separated at least 1 diameter depart; CO very
small or inverted flask-shaped (Fig. 15 A,B,E,F) ... 4

4. Epigyne with 1 small oval CO and long thin V-shaped
sclerotised tubes; CD, inverted V-shaped, SP about 1 of

their diameter apart ( Fig. 1 5 A,B) E. enterprise

Epigyne, CO inverted flask-shaped, wide U-shaped
sclerotised tubes, CO straight; SP twice their diameter
apart (Fig. 1 5E,F) E. milledgei

500m, PYKN, 17°16’S, 145°51’E, 17-24 Oct 1981 (QM
S27615); d, NE Qld O’Donogue’s Falls, 150m on tree,
16°26'S, 145°20’E, 15 May 1995, G Monteith, Pyretmm
(QM S43294); d, ME Qld, Crediton, Complex Notophyl
vine forest, 950m, on tree trunk, 21°13'S, 148°34'E, 14-21
Sep 75, RKohout & V.Davies (QM S445 1 ); 6 , Millstrcam
Conservation Pk., PYKN, 17°32’S, I45°30’E, 6 Dec 98,
G& S. Monteith (QM S47574).

DIAGNOSIS. AME smallest. Carapace sepia
bown, abdomen sepia brown with scutum and 2
pairs of white patches on anterior part and 2-3 in
front of spinnerets. Palp DTA retrolaterally with
membranous lamella only. Epigyne, copulatory
ducts elongate coiled, inverted V-shaped.

DESCRIPTION. Male (holotype). Total length
4. 1 6; ceph 2.08 long, 1 .04 wide, 0.76 high; cl/cw
1.48; sternum 1.04 long, 0.92 wide; sl/sw 1.13;
abdomen 2.08 long, 1.28 wide.

Euasteron enterprise sp. nov.
(Figs 5, 4 C,D, 7C,D 8A,B 1 5 A,B, 1 8)

ETYMOLOGY. From type locality, a
proposed mine site on North Stradbroke
Island. Noun in apposition.

MATERIAL. HOLOTYPE: d, SE Qld,

N. Stradbroke 1. Enterprise, Blackbutt 2,

60m, night coll, 27°34'S, 153°27’E, 7-14
Jan 2002, R.Ravcn & B.Baehr (QM
S55531). PARATYPES: Queensland: 6
SE Qld, Ml Cotton, 200m, rainforest,
intercept trap G Monteith (QM S44277);

$ SE Qld, N. Stradbroke I, Enterprise,

Blackbutt 1, 90m, night coll, 27°33'S,

153°28’E, 7 -14 Jan 2002, R.Raven &

B.Baehr (QM S55596); d, Ml Cotton,
upper gully, Flighl/Int, 27°36*S,

153°13'E. 12 Dec 97-7 Mar 98, G
Monteith (QM S47571); d, Kenilworth
State Forest, Sunday Ck Rd, 9.8km W of
Giarlie Moreland Park, wet sclerophyU,

26°40*irS, 152°36’35”E, 7 May 1998,

G Milledge (AM KS55660); 9, same
locality. Junction with Gighcr Creek Rd,

26°4r54”S, 152°33 , 5”E, 7.5.1998, G
Milledge (AM KS55661); d, Little Yabba
Ck, rainforest, 26°37’S, !52°4rE, 30 Nov
91-7 Jan 92, DJ. Cook (QM S25198); d,

SE Qld, Ml Tamborine, 670m, rainforest.

UNLO, 27°56’S, 153°12'E, 10 Jul 74,

V.Davies, R.Raven, C.L. Wilton (QM
S4357); d, NE Qld, Maalan Rd, 2km S
Palmerston H'way, PYKN, 17°36'S,

145°42'E, 18 May 1995, G Monteith (QM FIG 9. Euasteron spp. right male palps. A,B, E. lorne\ A, ventral; B,
S41058); d, NE Qld, Bellcnden Ker retrolateral; C,D, E. krebsorum\ C, ventral; D, retrolateral. Scale
Ra,l/2km S Cabletowcr 7, Earthwatch 0.5mm. PR = prong on retrolateral side of DTA; RE - rounded
Expedition & Qld Museum, rainforest, extension on flange.

12

MEMOIRS OF THE QUEENSLAND MUSEUM

Colour: carapace, sternum and chelicerae sepia
brown; maxillae and labium medium brown,
distally white; abdomen sepia brown; dorsally
with scutum, with 2 pairs of white patches on
anterior part, with 2 in front of spinnerets;
laterally with 1 white patch; ventrally pale brown
to dark brown; legs medium brown, annulated;
coxae MV white or pale; trochanter I-IV brown;
femur 1 dark brown, with proximal white patch,

I I-IV proximal part white, distal part dark brown;
tibia 1 distally white.

Eyes (Fig. 5A-C): AME smallest; eye group
width 0.51 of headwidth; AME 0.06; ALE 0.1;
PME 0. 1 ; PLE 0. 1 ; AME-AME 0.04; AME-ALE
0.04; PME-PME 0.04; PME-PLE 0.08;
ALE-PLE 0.02; eyes group AME-PME 0.26;
AME-AME 0.16; PME-PME 0.24. Clypeus:
0.36mm high.

Abdomen: tracheal spiracle small slit-like, with
sclerotised cover.

Male palp (Fig. 8A,B): DTA: PE not reaching
tibia, retrolaterally with membranous lamella;
VTA =FA short, rounded, SP not crossing TF;
tibia short; IP tiny, EP short, blunt; VtiA bird-
head-shaped.

Female (paratype). Total length 4.24; ccph 2.08
long, 1.28 wide, 0.84 high; cl/cw 1.62; sternum
0.92 long, 0.84 wide; sl/sw 1.09; abdomen 2.16
long, 1.32 wide.

Colour: as in male but abdomen without scutum.

Eyes: In rows (2-4-2); AME smallest; eye group
width 0.5 of headwidth; AME 0.06; ALE 0.1;
PME 0. 1 ; PLE 0. 1 ; AME-AME 0.04; AME-ALE
0.04: PME-PME 0.04; PME-PLE 0.08;
ALE-PLE 0.02; eyes group AME-PME 0.26;
AME-AME 0.16; PME-PME 0.24. Clypeus:
0.36mm high.

Epigyne (Fig. 15A,B): With 1 small, oval CO,
with long, V-shaped, sclerotised tubes; CD
elongate coiled, inverted V-shaped, ending in
small, separated S. Female palpal claw strong
with more than 4 teeth.

Variation: Some specimens are slightly smaller
than HT. Variation in colour: Carapace with or
without dark fovea and bifurcate stripes,
abdomen with 1 or 2 white spots in front of
spinnerets and on femur I the proximal part can be
white.

DISTRIBUTION. Wet Tropics to Southern
Queensland (Fig. 18). Rainforest.

Euasteron bartoni sp. nov.

(Figs 10A,B, 16C,D, 21)

ETYMOLOGY. For N. Barton, the collector of the types.

MATERIAL. HOLOTYPE: d, Lucknow area, Vic., Oct
1979, 37°49’S, 147°39’E, 10.1979, N. Barton (AM
KS83927). PARATYPES: Victoria: 9, as for holotype
(AM KS84124).

DIAGNOSIS. Similar to E. krebsorum with AME
smallest; abdomen with 2 pairs of white patches
and 1 patch in front of spinnerets; booklungs
pale. Palp DTA retrolaterally with strong prong.
Epigyne with 1 broadly oval semicircular CO and
long, U-shaped, sclerotised tubes; CD elongate,
coiled, straight, ending in small, separated
spermathecae. Separated from E. krebsorum by
abdomen with only I spot in front of spinnerets
and FA short, not bent.

DESCRIPTION. Male (holotype). Total length
3.6; ceph 1.68 long, 1.2 wide, 0.68 high; cl/cw
1.4; sternum 0.8 long. 0.72 wide; sl/sw 1.11;
abdomen 1 .48 long, 0.92 wide.

Colour: carapace orange brown; with dark fovea
and dark bifurcate patches in front; sternum
orange brown; chelicerae medium brown;
maxillae and labium medium brown, distally
white; abdomen sepia brown, with 2 pairs of
white patches on anterior part and 2-3 in front of
spinnerets; laterally with 1 big white patch and
pale booklungs; ventrally dark brown; legs
medium brown, with darker brown lateral stripes;
clearly annulated; coxae I-IV white; trochanter
I-IV medium to dark brown; femur I, II dark
brown. III, IV proximal part white, distal part
dark brown.

Eyes: AME smallest; eye group width 0.54 of
headwidth; AME 0.06; ALE 0.08; PME 0.08;
PLE 0.08; AME-AME 0.04; AME-ALE 0.03;
PME-PME 0.04; PME-PLE 0.06; ALE-PLE
0.02; eyes group AME-PME 0.24; AME-AME
0.16; PME-PME 0.2. Clypeus: 0.28mm high.
Male palp (Fig. 1()A,B): DTA, PE not reaching
tibia; retrolaterally with strong prong; SP
S-shaped, not crossing TF; tibia, IP as long as EP,
blunt, VtiA short, hooked.

Female (paratype). Total length 3.8; ceph 1.88
long, 1.2 wide, 0.68 high; cl/cw 1.57; sternum
0.84 long, 0.76 wide; sl/sw 1.11; abdomen 1.92
long, 1.24 wide.

Colour: as in males.

Eyes: AME smallest; eye group width 0.52 of
headwidth; AME 0.06; ALE 0.08; PME 0.08;
PLE 0.08; AME-AME 0.04; AME-ALE 0.03;
PME-PME 0.04; PME-PLE 0.08; ALE-PLE

NEW AUSTRALIAN SPIDER GENERA

13

0.02; eyes group AME-PME 0.28; AME-AME
0.16; PME-PME 0.2. Clypeus 0.3mm high.
Epigyne (Fig. 16C,D): with 1 broadly oval CO
attached to epigastric fold and long, U-shaped,
sclerotised tubes; CD elongate, coiled about
straight axis, ending in small separated S.

DISTRIBUTION. Victoria (Fig. 21).

Euasteron gibsonae sp. nov.

(Figs 10C,D, 21)

ETYMOLOGY. For Leslie Gibson, the collector of the
types.

MATERIAL. HOLOTYPE. <3 , SW Qld, Astraleba Downs
NP., pitfall traps, 24°13’05"S, 140°34’48”E, Sep.1997,
L.Gibson (QM S55533). PARATYPES: Queensland: <J,
as for holotype (QM S55532). Northern Territory: 8d,
Barkly Tablelands, Connells Lagoon. Mitchell grassland.
Quadrat 1, pitfall traps, 18°52’S, 136°32’E, Mar 96, T.B.
Churchill, A. Fisher (CS1RO A643); 3d, same locality,
Helen Springs Stn., Quadrat 1 , 28°24’S, 1 33°52'E (CSIRO
A0431); d, same locality, Birrindudu, site 4, 18°24 S,
129°26'E, Apr 96, T.B. Churchill, A. Fisher (CSIRO
A 1449); d, same locality, Kirkimbie, site 4, 17°44'S,
129°14*E, Jun 95, T.B. Churchill, A. Fisher (CSIRO
A 1465); d, same locality (CSIRO A 1460).

DIAGNOSIS. AME small. Carapace sepia
bown, abdomen sepia brown with 1 big pair of
white patches on anterior part and 1 elongate in
front of spinnerets. Tibial apophysis, IP short EP
short blunt.

DESCRIPTION. Male (holotype). Total length
4.16; ceph 2.04 long, 1.56 wide, 0.8 high; cl/cw
1.31; sternum 0.92 long, 0.84 wide; sl/sw 1.09;
abdomen 2.12 long, 1.28 wide.

Colour: carapace, sternum and chelicerac sepia
brown; maxillae and labium medium brown,
distally white; abdomen sepia brown, dorsally
with 1 big pair of white patches on anterior part
and 1 elongate white stripe in front of spinnerets;
laterally sepia brown; ventrally dark brown; legs
medium brown.

Eyes: AME smallest; eye group width 0.47 of
headwidth; AME 0.06; ALE 0.1; PME 0.1; PLE
0.1; AME-AME 0.04; AME-ALE 0.02; PME-
PME 0.04; PME-PLE 0.08; ALE-PLE 0.04; eyes
group AME-PME 0.3; AME-AME 0.16; PME-
PME 0.24. Clypeus: 0.4mm high.

Male palp (Fig. 10CD): DTA, PE not reaching
tibia; retrolaterally with strong prong; SP not
crossing transparent field; tibia short; IP short, EP
short blunt; VtiA bird-head-shaped.

Female unknown.

DISTRIBUTION. SW Queensland to Barkly
Tableland (Fig. 21). Grassland.

Euasteron krebsorum sp. nov.

(Figs 9C,D, 16E,F 18)

ETYMOLOGY For K. & D. Krebs, collectors of the types.

MATERIAL. HOLOTYPE. d, Hay, NSW, 8km West of
irrigation channel, grassland, pitfall traps, 33°3'S,
144°45'E, 19 Dec 97-9 Jan 98, K. & D. Krebs (QM
S460984). PARATYPES: New South Wales: 2d, as for
holotype (QM S46736). South Australia: 50d,29,
Gawler, 16km. Open grassland, pitfall traps, 34°36’S,
138°55’E, 19 Dec 97-9 Jan 98, K. & D. Krebs (QM
S46710); <3,9, same locality (AM KS84125).

DIAGNOSIS. Carapace sepia brown, abdomen
sepia brown with 2 pairs of white spots on
anterior part and 2 in front of spinnerets. AME
small. DTA retrolaterally with prong. FA long
directed distally.

DESCRIPTION. Male (holotype). Total length
3.56; ceph 1 .84 long, 1.24 wide, 0.68 high; cl/cw
1.48; sternum 0.8 long, 0.72 wide; sl/sw 1.11;
abdomen 1.72 long, 1 wide.

Colour: carapace sepia brown; sternum orange
brown; chelicerae medium brown; maxillae and
labium medium brown, distally white; abdomen
sepia brown, dorsally with 2 pairs of white
patches on anterior part and 2-3 in front of
spinnerets; laterally with 1 white patch; ventrally
dark brown; legs medium brown; with indistinct
colour pattern; coxae I-IV white; femur I dark
brown, II-IV proximal part white, distal part dark
brown.

Eyes: AME smallest; eye group width 0.54 of
headwidth; AME 0.08; ALE 0. 1 ; PME 0. 1 ; PLE
0.1; AME-AME 0.02; AME-ALE 0.02;
PME-PME 0.04; PME-PLE 0.06; ALE-PLE
0.02; eyes group AME-PME 0.26; AME-AME
0.18; PME-PME 0.24. Clypeus: 0.28mm high.
Male palp (Fig. 9C,D): DTA, PE not reaching
tibia; retrolaterally with strong prong; VTA =FA
elongate directed distally, SP, not crossing TF;
tibia short; IP longer than EP, sharp, EP blunt;
VtiA hooked.

Female (paratype). Total length 3.56; ceph 1.84
long, 1.24 wide. 0.68 high; cl/cw 1.48; sternum
0.8 long, 0.72 wide; sl/sw 1.11; abdomen 1.72
long, 1 wide.

Colour: as in male.

Eyes: nearly equal sized, or AME smallest; eye
croup width 0.54 of headwidth; AME 0.08; ALE
0.1; PME 0.1; PLE 0.1; AME-AME 0.02;
AME-ALE 0.02; PME-PME 0.04; PME-PLE

14

MEMOIRS OF THE QUEENSLAND MUSEUM

0.06; ALE-PLE 0.02; eyes group
AME-PME 0.26; AME-AME 0.18;
PME-PME 0.24. Clypeus: 0.28mm
high.

Epigyne (Fig. 16EF): with 1 oval
semicircular CO attached to
epigastric fold and long V-shaped
sclerotised tubes, CD elongate
coiled, V-shaped. S touching.

DISTRIBUTION. South Australia,
New South Wales (Fig. 18). Open
forest and grassland.

Euasteron lorne sp. nov.
(Figs 9A,B, 18)

ETYMOLOGY. The species name is taken
from the type locality. A noun in apposition.

MATERIAL. HOLOTYPE: d, Lome State
Forest near Taree, NSW, site 86( 1 ) in litter,
29 Oct 78-10 Dec 79, 31°35’S, 150°23'E,

17.11.1978, D Milledge (AM KS15971).
PARATYPES: New South Wales: 2d,
Lome SF near Lome site 86 (5), in litter. 29
Oct 78-10 Dec 79, 31°35’S, 152°57’E,

12.11.1978, D Milledge (AM KS16059);
d, Lome SF near Lome site 86 (5), in litter.
29 Oct 78-10 Dec 79, 31°35 , S, 152°57’E,

12.11.1 978, D Milledge (QM S608 1 7).

FIG. 10. Euasteron spp. right male palps. A,B, E. bartoni ; A, ventral;
B, retrolateral; C,D, E.gibsonae ; C, ventral; D, retrolateral. Scale
0.5mm.

DIAGNOSIS. AME small.

Carapace orange, legs yellow
orange, abdomen sepia brown, with
2 pairs of white patches on anterior
part and 3 in front of spinnerets. Palp
DTA with folded prong.

DESCRIPTION. Male (holotype).

Total length 4. 1 6; ceph 2.12 long, 1 .6
wide, 1 high; cl/cw 1.32; sternum 1.0
long, 0.96 wide; sl/sw 1.04; abdomen 2.04 long,
1.32 wide.

Colour: carapace, sternum and chelicerae orange;
maxillae and labium orange, distally white;
abdomen sepia brown, dorsally with 2 pairs of
white patches on anterior part and 3 in front of
spinnerets, laterally with 1 white patch; ventrally
dark brown; legs yellow.

Eyes: AME smallest; eye group width 0.48 of
headwidth; AME 0.06; ALE 0.12; PME 0.12;
PLE 0.12; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0. 1 ; ALE-PLE 0.04;
eyes group AME-PME 0.3; AME-AME 0.16;
PME-PME 0.28. Clypeus: 0.4mm high.

Male palp (Fig. 9A,B): DTA PE not reaching
tibia; retrolaterally with folded prong; FA short.

directed prolaterally; SP S-shaped, not crossing
TF; tibia short; IP absent, EP short blunt; VtiA
hooked.

Female unknown.

DISTRIBUTION. Lome State Forest, New
South Wales (Fig. 18). Coastal forest.

Euasteron millcdgei sp. nov.

(Figs 8E,F, 15E,F, 18)

ETYMOLOGY. For D. Milledge, one of the collectors of
the types.

MATERIAL. HOLOTYPE: d , Gibraltar Ra, NSW, lower
rainforest, 450m, pitfall traps 29°35’S, 152°13’E, 30
Mar- 10 Nov 1980, G & S. Monteith (QM S4261).
PARATYPES: New South Wales: d, Bruxner Pk, via
Coffs Harbour, rainforest, 150m, pitfall traps, 30°14*S,

NEW AUSTRALIAN SPIDER GENERA

15

FIG 1 1 . Euasteron spp. right male palps. A,B. E raveni ; A, ventral; B retrolateral; C,D. E. han’eyi; C, ventral; D,
retrolateral; E,F, E. churchillae; E, ventral; F, retrolateral. Scales 0.5mm.

153°06’E, 22 Mar- 13 Nov 80, G & S. Monteitlt (QM
S3744); 6, Dorrigo Nat Pk, Never Never area. Sassafras
Ck, wet temperate rainforest, 30°22’S, 152°45’E, 7 Dec
92, D.J. Bickel, (AM K.S35052); 6, Lome SF near Lome
site 86(1), litter, 29 Oct 78-10 Dec 79, 31°35’S. 152°57’E,
4. 1 1 .1979, D. Millcdgc (AM K.S5607); 6 , 9 same locality,
4 Nov 79 (AM KS5633); d , same locality, 1 2 Nov 78 (AM
K.S 16060).

DIAGNOSIS. AME small. Carapace sepia
brown, abdomen sepia brown with 2 pairs of
white spots on anterior part and 3-4 in front ot
spinnerets. Palp, DTA retrolaterally with small
lamella and short prong. Epigyne, CO inverted
flask-shaped

DESCRIPTION. Male (holotype). Total length
4.28; ceph 2.24 long, 1 .68 wide, 1 .08 high; cl/cw

1.33; stemum 0.88 long, 0.84 wide; sl/sw 1.04,
abdomen 2.04 long, 1.4 wide.

Colour: carapace sepia brown; with dark fovea
and dark bifurcate patches in front; stemum orange
brown; chelicerac medium brown; maxillae and
labium orange, distally white; abdomen sepia
brown, dorsally with 2 pairs of white patches on
anterior part and 3 in front of spinnerets, laterally
with 3 elongate and broad white patches,
ventrally with 1 longitudinal white stripe from
epigastric fold to near tracheal spiracle; legs
yellow; clearly annulated; coxae I-tV white or
pale; trochanter 1-lV medium to dark brown,
femur I- IV proximal part white, distal part dark
brown.

16

MEMOIRS OF THE QUEENSLAND MUSEUM

Eyes: AME smallest; eye group width 0.53 of
headwidth; AME 0.08; ALE 0.12; PME 0.12;
PLE 0.12; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0. 1 ; ALE-PLE 0.04;
eyes group AME-PME 0.36; AME-AME 0.2;
PME-PME 0.28. Clypeus: 0.44mm high.

Male palp (Fig. 8EJF): DTA, PE not reaching
tibia; retrolaterally with membranous lamella
and short prong; FA big, flat, with rounded tip,
directed distally, SP not crossing TF; tibia short;
IP tiny, EP short, VtiA hooked.

Female (paratype). Total length 4.8; ceph 1.96
long, 1 .38 wide, 1 high; cl/cw 1 .44; sternum 0.84
long, 0.8 wide; sl/sw 1.05; abdomen 2.84 long, 2
wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.53 of
headwidth; AME 0.06; ALE 0.1; PME 0.1; PLE
0.1; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0. 1 ; ALE-PLE 0.04;
eyes group AME-PME 0.3; AME-AME 0.16;
PME-PME 0.26.

Epigyne (Fig. 15E,F): CO inverted flask-shaped,
with U-shaped sclerotised tubes, copulatory
ducts elongate coiled. Spermathecae twice their
diameter apart.

DISTRIBUTION. NE New South Wales (Fig.
18). 6

HABITAT. Rainforest.

Euasteron monteithorum sp. nov.

(Figs 8C,D, 15B,E, 18)

ETYMOLOGY. For Dr Geoff and Sybil Moneith of the
Queensland Museum, the collectors of the holotype.

MATERIAL. HOLOTYPE: <3, Poverty Point, NSW,
20kip SE Tenterficld, rainforest, 945m, 29°08’S, 1 52°07 , E,

3 Oct 78-21 Feb 79 G&S. Monteitli (QM SI2149).
PARATYPES: New South Wales: 6 , as for holotype (QM
S3772); 6 , Whian Whian SF, rainforest, 2 1 0m pitfall traps.
28°4rs. 153°19'E, 19 Aug- 16 Nov 74, G& S. Monteitli
(QM S 1 2 1 35); 6 . same locality but 28° 1 4'S, 1 53° 1 9’E, 1 9
Aug- 1 6 Nov 78. G& S. Monteith (QM S3774); 6 Victoria
Pk, via Alstonville, rainforest, 250m. pitfall traps, 28°53’S,
153°23’E, 16 Nov-26 Dec 74, G & S. Monteith (QM
s 12 142); d , same locality but 26 Dec 1974-23 Mar 75, G
& S. Monteith (QM S 12 1 41 ); d , 9 , same locality but (QM
S 12 144). Queensland: d,9, SE Qld, Kmombit Tops,
rainforest. 1000m, pitfall traps, 24°22’S, 15 POLE, 22-26
Feb 82, G Monteith, D. Yeates, G Thompson (QM S3730).

DIAGNOSIS. AME small. Carapace orange
brown, abdomen with 2 pairs of white patches on
anterior part and 2-3 in front of spinnerets. Palp,
DTA retrolaterally with lamella; tibia IP short.

sharp, EP blunt, sticking out. Epigyne with 2
small, semicircular CO.

DESCRIPTION. Male (holotype). Total length
3.76; ceph 1.96 long, 1 .48 wide, 0.96 high; cl/cw
1.32; sternum 1.0 long, 0.88 wide; sl/sw 1.13;
abdomen 1.8 long, 1.2 wide.

Colour: carapace orange brown; with dark fovea
and dark bifurcate patches in front; sternum
orange brown; chelicerae orange; maxillae and
labium pale brown, distally white; abdomen
sepia brown, dorsally with 2 pairs of big white
patches on anterior part and 2-4 in front of
spinnerets, laterally with 2 elongate white
patches, first one larger; ventrally with 1
longitudinal white stripe from epigastric fold to
near tracheal spiracle and 1 pair of white patches
near tracheal spiracle; legs pale brown; with
indistinct colour pattern; coxae I-IV white;
trochanter I-IV white; femur I-IV proximal part
white, distal part dark brown.

Eyes: AME smallest; eye group width 0.5 of
headwidth; AME 0.06: ALE 0.1; PME 0.1; PLE
0.1; AME-AME 0.04; AME-ALE 0.06;
PME-PME 0.08; PME-PLE 0.04; ALE-PLE
0.04; eyes group AME-PME 0.24; AME-AME
0.16; PME-PME 0.24. Clypeus: 0.36mm high.

Male palp (Fig. 8C,D): DTA PE not reaching
tibia; retrolaterally with membranous lamella;
FA short, straight, with rounded tip; SP not
crossing TF; tibia short; IP short, sharp; EP blunt
sticking out; VtiA short, hooked.

Female (paratype). Total length 3.72; ceph 1.8
long, 1.28 wide, 0.84 high; cl/cw 1.4; sternum
0.88 long, 0.8 wide; sl/sw 1.1; abdomen 1.92
long, 1 .48 wide.

Colour: as in males.

Eyes: AME smallest; eye group width 0.5 of
headwidth; AME 0.06; ALE 0. 1 ; PME 0. 1 ; PLE
0.1; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.08; PME-PLE 0.04; ALE-PLE
0.04; eyes group AME-PME 0.26; AME-AME

0. 16; PME-PME 0.24. Clypeus: 0.36mm high.

Epigyne (Fig. I5B,E): With 2 small, semi-
circular copulatory openings, copulatory ducts
elongate coiled, ending in small spermathecae.

DISTRIBUTION. Northeastern New South
Wales and southern Queensland (Fig. 18). Rain-
forest.

KEY TO SPECIES OF EUASTERON
JULIANNAE GROUP, MALES

1 . DTA retrolaterally with strong prong (Figs 11,1 2CD) . 2

NEW AUSTRALIAN SPIDER GENERA

17

DTA retrolaterally with lamella and without or only tiny
prong (Figs 13, 14) 5

2. Carapace dark brown, abdomen with scutum, palp, FA

long, straight, directed prolatero-distally (Fig. 1 1E,F)

E. churchillae

Carapace orange brown with dark fovea and bifurcate
patches, abdomen without scutum, palp, FA short or
long, bent and directed distally (Figs 1 1A-D, 12C,D) . 3

3. Abdomen laterally with I white patch, palp: DtiA EP long

(Fig. 12C,D) E.ulrichi

Abdomen laterally without any colour pattern, palp; DtiA
EP short (Fig. 1 I A-D) 4

4. Femur 1 dark brown, palp, SP partly crossing TF, FA short

(Fig. 1 1A,B)., E.raveni

Femur I proximal ly white, palp, SP partly crossing TF, FA

long bent (Fig. 1 1C,D) E. harveyi

5. Carapace yellow with dark margin and dark cephalic

area; abdomen pale with dark chevrons (Figs 6A-C,

13C,F) 6

Carapace sepia or orange brown, abdomen sepia brown
with 2 or 3 pairs of white spots on anterior part and 1 -3 in
front of spinnerets, or pale chevrons ,7

6. Palp, SP not crossing TF; FA wider than long; (Fig.

13C.D) E.atriceps

Palp, SP partly crossing TF, FA as long as wide (Fig.

13E,F) E. Carnarvon

7. Abdomen with pale chevrons; DtiA EP with deep
incision, palp, SP not crossing TF (Fig. 13 A,B) . . . E.

juliannae . . .

Abdomen with 2-3 pairs of white spots on anterior part

and 3 in front of spinnerets; DtiA EP without deep
incision , palp SP crossing TF completely (Fig. 4 A-D) 8

8. Abdomen with 3 pairs of white spots on anterior part and

3 in front of spinnerets, palp, SP broad (Fig. 14A,B)

E.johannae

Abdomen with 2 pairs of white spots on anterior part and
3 in front of spinnerets; palp, SP narrow (Fig. 14C,D)
E. ursulae

Euasteron churchillae sp. nov.

(Figs 1 1E,F, 19)

ETYMOLOGY. In honour of Dr Tracey Churchill, the
collector of the types and in esteem for her important
ecological work on Australian spiders.

MATERIAL. HOLOTYPE: d, Daly River, NT, Quadrat
C6, PWCNT, pitfall traps, 13°55’23”S, 131°24W’E,
21.6.99, T. Churchill (CSIRO A2545). PARATYPES:
Northern Territory: d, Daly R, Quadrat KIO, PWCNT,
14°43’6rS, 132°37’03”E, 30 Aug 98 (CSIRO A3431).

DIAGNOSIS. AME biggest, carapace sepia
brown, abdomen with scutum, with 2 pairs of
white patches on anterior part and 2 in front of
spinnerets. Palp, DTA with big prong, base of
embolus with big straight FA.

DESCRIPTION. Male (holotype). Total length
3.24; carapace 1.64 long, 1.2 wide, 0.8 high;
cl/cw 1.36; sternum 0.8 long, 0.72 wide; sl/sw
1.11; abdomen 1 .6 long, 0.88 wide.

Colour: carapace sepia brown; sternum orange
brown; chelicerae medium brown; maxillae and
labium medium brown, distally white; abdomen
sepia brown; dorsally with scutum and 2 pairs of
white patches on anterior part and 2 in front of
spinnerets; laterally with 1 white patch; legs
medium brown, with darker brown lateral stripes;
clearly annulated; coxae I-IV white; trochanter
I-IV brown; femur I-IV proximal part white,
distal part dark brown.

Eyes: AME largest; eye group width 0.6 of
headwidth; AME 0.13; ALE 0. 1 : PME 0. 1 ; PLE
0.1; AME-AME 0; AME-ALE 0.02; PME-PME
0.04; PME-PLE 0.08; ALE-PLE 0.02; eyes group
AME-PME 0.3; AME-AME 0.26; PME-PME

0. 24. Clypeus: 0.32mm high.

Male palp (Fig. 1 1E,F): DTA, PE not reaching
tibia; retrolaterally with strong prong; FA
elongate, straight, with rounded tip; SP not
crossing TF; DtiA IP shorter than EP, Ep with
blunt tip; VtiA birdhead-shaped.

Variation: the only other examined male (A343 1 )
is about 1/3 longer than the HT with body length
of 4.24mm and ceph length of 2.24mm. Body
shape colour and palp shape are the same.

Female unknown.

DISTRIBUTION. Daly River area, Northern
Territory (Fig. 19). Semi-arid area, desert.

Euasteron harveyi sp. nov.

(Figs 1 1C,D, 19)

ETYMOLOGY. In honour of Dr Mark Harvey of the
Western Australian Museum, in esteem for his important
work on Australian arachnids.

MATERIAL. HOLOTYPE: d, Marun CALM Site 8/4,
3.4km SW of Manning Ck, Prince Frederic Harbour 6.
WA, malaise trap. 15WS, 125°2rE, 125°2rE, 1 1 Jun 88,

1. D. Naumann (WAM T51851). PARATYPES: Western
Australia: d, Marun CALM Site 8/4, 3.4km SW of
Manning Ck, Prince Frederic Harbour 6, Malaise trap,
15°00’S, 125°2TE, l25 a 21’E v 11 Jun 88, I.D. Naumann
( WAM T51852).

DIAGNOSIS. AME biggest, carapace orange
brown, abdomen with with 2 pairs of white
patches on anterior part and 2-3 in front of
spinnerets, differs from E. ravetii in palp with
long bent FA, SP not crossing TF.

DESCRIPTION. Male (holotype). Total length
3.4; ceph 1.6 long, 1.28 wide, 0.72 high; cl/cw
1.25; sternum 0.76 long, 0.72 wide; sl/sw 1.05;
abdomen 1.8 long, 1.08 wide.

18

MEMOIRS OF THE QUEENSLAND MUSEUM

Colour: carapace orange brown;
with dark fovea and dark bifurcate
patches in front; sternum, chelicerae,
maxillae and labium white to
yellow; abdomen sepia brown;
dorsally with 2 pairs of white
patches on anterior part and 2-3 in
front of spinnerets; legs pale to
white; clearly annulated; coxae I-IV
white; trochanter I-IV white; femur
I-IV proximal part white, distal part
dark brown; tibia I-IV proximal and
distal part brown, medial part white.
Eyes: AME largest; eye group width
0.55 of headwidth; AME 0.12; ALE
0. 1 ; PME 0. 1 ; PLE 0. 1 ; AME-AME
0.02; AME-ALE 0.04; PME-PME
0.04; PME-PLE 0.08; ALE-PLE
0.04: eyes group AME-PME 0.32;
AME-AME 0.26; PME-PME 0.24.
Clypeus 0.32mm high.

Male palp (Fig. 11C,D): DTA, PE
not reaching tibia; retrolaterally with
strong prong; FA elongate,
sickleshaped, with rounded tip; SP
S-shaped, not crossing TF; DtiA IP
about 3 times longer than wide,
sharp, EP short, blunt; VtiA
bird-head-shaped.

Female unknown.

DISTRIBUTION. Known only from
type locality (Fig. 19).

FIG. 12. Euasteron spp. right male palps. A,B, E. willeroo; A, ventral;
B, retrolateral; C,D, E. ulrichi ; C, ventral; D, retrolateral. Scales
0.5mm.

Euasteron raveni sp. nov.

(Figs 1 1 A,B, 19)

ETYMOLOGY. In honour of Dr Robert
Raven of the Queensland Museum, in
esteem, for his important work on Australian
spiders^

MATERIAL. HOLOTYPE: 6, NW Qld, Mount Isa,
20°44 , S, 139°29’E, Jan 1981, W. Myors (AM KS6448).

DIAGNOSIS. AME biggest, carapace orange
brown, abdomen with with 2 pairs of white
patches on anterior part and 2-3 in front of
spinnerets, palp with strong prong and short FE,
SP partly crossing TF.

orange brown; chelicerae orange; maxillae and
labium orange, distally white; abdomen sepia
brown; dorsally with 2 pairs of white patches on
anterior part and 2-3 in front of spinnerets;
ventrally pale brown; legs medium brown, with
darker brown lateral stripes; clearly annulated;
coxae I-IV white; trochanter I-IV brown; femur I
dark brown, II-I V proximal part white, distal part
dark brown.

DESCRIPTION. Male (holotype). Total length
4.00; carapace 2.04 long, 1.52 wide, 0.8 high;
cl/cw 1.34; sternum 0.96 long, 0.84 wide; sl/sw
1.14; abdomen 1.96 long, 1.24 wide.

Colour: carapace orange brown; with dark fovea
and dark bifurcate patches in front; sternum

Eyes: AME largest; eye group width 0.61 of
headwidth; AME 0.14; ALE 0.12; PME 0.12;
PLE 0.12; AME-AME 0; AME-ALE 0.02;
PME-PME 0.04; PME-PLE 0.08; ALE-PLE
0.02; eyes group AME-PME 0.38; AME-AME
0.28; PME-PME 0.28. Clypeus: 0.4mm high.

NEW AUSTRALIAN SPIDER GENERA

19

FIG. 13. Euasteron spp. right male palps. A,B, E. juliannae: A, ventral; B retrolateral; C,D, E. atriceps ; C,
ventral; D. retrolateral; E,F, E. Carnarvon ; E, ventral; F, retrolateral. Scales 0.5mm.

Abdomen: tracheal spiracle small, slit-like,
covered with small stout spines.

Male palp (Fig. 1 1 A,B): DTA, PE not reaching
tibia; retrolaterally with strong prong; FA short,
straight; SP S-shaped, partly crossing TF; DtiA,
IP 3 times as long as wide, sharp, EP blunt; VtiA
bird-head-shaped.

Female unknown.

DISTRIBUTION. Known only from Mt Isa, NW
Queensland (Fig. 19).

Euasteron ulrichi sp. nov.

(Figs 12C,D, 19)

ETYMOLOGY. For Ulrich Bachr, the author’s son.

MATERIAL. HOLOTYPE: <J, 26km E Napier Downs,
WA, 17°15’S, 124°44’E, 24 Nov 1984, B. & M. Baehr
(QM S55907).

DIAGNOSIS. AME biggest, carapace orange
brown, abdomen with 2 pairs of white patches on
anterior part and 1 in front of spinnerets. Pulp
DTA retrolaterally with strong prong, base of
embolus with short straight FA directed distally.

DESCRIPTION. Male (holotype). Total length
3.6; ceph 1.84 long, 1.24 wide, 0.72 high; cl/cw
1.48; sternum 0.8 long, 0.8 wide; sl/sw 1;
abdomen 1.76 long, 1 .04 wide.

Colour; carapace orange brown: with dark tovea
and dark bifurcate patches in front; sternum
orange brown; chclicerae orange; maxillae and
labium yellow, distally white; abdomen sepia
brown; with dorsally 2 pairs of white patches on
anterior part and 1 in front of spinnerets; laterally
with 1 white patch; vcntrally pale brown; legs
yellow; clearly annulated; coxae I -IV white;

20

MEMOIRS OF THE QUEENSLAND MUSEUM

trochanter 1-IV brown; femur I-IV proximal part
white, distal part dark brown.

Eyes: AME largest; eye group width 0.62 of
headwidth; AME 0.14; ALE 0.12; PME 0.12;
PLE 0.12; AME-AME 0; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.32; AME-AME

0. 28. PME-PME 0.28. Clypeus: 0.36mm high.
Male palp (Fig. 12C,D): DTA, PE not reaching
tibia; retrolaterally with strong PR; FA short,
straight, directed distally; SP not crossing TF;
DtiA IP 3 times as long as wide, sharp, EP blunt
sticking out; VtiA bird-head-shaped.

Female unknown.

DISTRIBUTION. Known only from Napier
Downs (Fig. 19).

KEY TO SPECIES OF EUASTERON
JULIANNAE GROUP, FEMALES

1. Carapace yellow with dark margin and dark cephalic

area; abdomen pale with dark chevrons (Figs 6, 1 7E,F)
* E. Carnarvon

Carapace sepia or orange brown, abdomen sepia brown
with 2 or 3 pairs of white spots on anterior part and 1 -3 in
front of spinnerets, or pale chevrons . 2

2. Abdomen with pale chevrons; epigync, CD extremely

long straight (Fig. 1 7G,H) E. juliannae

Abdomen with 2-3 pairs of white spots on anterior part
and 3 in front of spinnerets; epigyne, CD long V-shaped
(Fig. 17A-D) 3

3. Abdomen with 3 pairs of white spots on anterior part and
3 in front of spinnerets; CO not attached to epigastric fold

(Fig. 17A,B) E.johannae

Abdomen with 2 pairs of white spots on anterior pari and
3 in front of spinnerets; CO attached to epigastric fold
(Fig. 1 7C,D) E. ursulae

Euasteron johannae sp. nov.

(Figs 14A,B, 17A,B, 20)

(WAM T44463); <3, same locality, site BR 1 2, 23°06’47 M S,
1 16°00’42"E (WAM T44464)'; 23,9, same locality,
quadrat 5, 23 0 24’41"S, 1I5°53’39"E, /9/1995, S. van
Leeuwen. B. Bromilow (WAM T44465); 3,9, same
locality, quadrat 10, 23°05 > 45"S 116°00’35 M E (WAM
T44466); 23,9, same locality, quadrat 11, 23°06’2rS,
115°59 , 52 M E (WAM T44467); 3, same locality, site
BR10,23°05’45"S, 116°00’35"E, 19/6/1994, RG &,GW.
Kendrick (WAM T44468); 3 (palp drawing) same
locality, site BR11, 23 o 06 t 2l"S, 115°59’52"E, 19-22 Jun
94 (WAM T44469).

DIAGNOSIS. Differs from E. atriceps and E.
Carnarvon by the sepia brown ceph, from E.
ursulae and E. juliannae by femur I, dark brown,
II, IV proximal part white, distal part dark brown
and by abdomen with 3 pairs of white patches on
anterior part and 3 in front of spinnerets.

DESCRIPTION. Male (holotype). Total length
4. 12; ceph 2.04 long, 1 .44 wide, 0.84 high; cl/cw
1.42; sternum 0.96 long, 0.92 wide; sl/sw 1.04;
abdomen 2.08 long, 1 .4 wide.

Colour: carapace sepia brown; sternum orange
brown; chelicerae medium brown; maxillae and
labium medium brown, distally white; abdomen
sepia brown; dorsally with 3 pairs of white
patches on anterior part and 3 in front of
spinnerets; legs medium brown, with darker
brown lateral stripes; clearly annulated; coxae
I-IV white; trochanter I-IV white; femur I dark
brown, II-I V proximal part white, distal part dark
brown.

Eyes: AME largest; eye group width 0.65 of
headwidth; AME 0. 1 4; ALE 0. 1 ; PME 0. 1 2; PLE
0.12; AME-AME 0.02; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.06; ALE-PLE
0.04; eyes group AME-PME 0.32; AME-AME
0.3; PME-PME 0.3. Clypeus: 0.32mm high.

ETYMOLOGY. For Johanna Baehr, the author’s elder
daughter.

MATERIAL. HOLOTYPE: 3, Barlee Range Nature
Reserve, quadrat 5, WA, dry pitfall traps, 23°24’4rS
1 15°53’39 M E, /6/1994, S.van Leeuwen, B. Bromilow
(WAM T44458). PARATYPES: WA: 3,9. same locality
(WAM T51853); 23,9, same locality, quadrat I,
23°02’39"S 1 15°48’44"E (WAM T44457) ; <3, same
locality, quadrat 10, 23°05’45"S, 116°00'35"E (WAM
144460); 18d,9, same locality, quadrat 1, 23°02'39"S,
1 15°48'44"E, /8/ 1993 (WAM T44453); 5(3,9, same
locality, quadrat 5, 23°24'4I"S, 115°53'39"E (WAM
T44454); <3, same locality, quadrat 9, 23°06’06"S,
116°00’28”E (WAM T44455); 6. same locality, quadrat
12, 23°06’47"S, 116°00'42"E (WAM T44456); !2c3,
same locality, site BR5. 23°24'4I"S, 1 15°53’39"E, 11-14
Jun 94, P.G &, GW. Kendrick (WAM T4446 1 ); 23 (3 , same
locality, site BR9, 23°06’06"S, 16°00’28"E, 19-22 Jun 94

Male palp (Fig. 14A,B): DTA, PE reaching tibia;
retrolaterally with membranous lamella; FA
short, rounded; SP broad, S-shaped, crossing TF
totally; DtiA IP as long as EP, sharp, EP blunt;
VtiA hooked.

Female (paratype). Total length 5.8; ceph 2.28
long, 1.52 wide, 0.84 high; cl/cw 1.5; sternum
1.04 long, 0.96 wide; sl/sw 1.08; abdomen 2.8
long, 2 wide.

Colour: as in male.

Eyes: AME largest; eye group width 0.66 of
headwidth; AME 0. 14; ALE 0.12; PME 0.14;
PLE 0.14; AME-AME 0.02; AME-ALE 0.02:
PME-PME 0.04; PME-PLE 0.04; ALE-PLE
0.08; eyes group AME-PME 0.42; AME-AME
0.3; PME-PME 0.34. Clypeus: 0.4mm high.

NEW AUSTRALIAN SPIDER GENERA

21

115°53'12"E, 11-14 Jun 94, P.G & GW.
Kendrick (WAM T44462). PARATYPES:
Western Australia: 4d , 9 , Barlee Range Nature
Reserve, site BR6, dry pitfall traps 23°23’2 1 "S,
115°53’12"E, 11-14 Jun 94, P.G & GW.
Kendrick (WAM T52337); 2 d, same locality,
(QM S60818); d , same locality, quadrat 7,
23°22’45 ,, S, 1 1 5°52’50"E, Jun 94, S. van
Leeuwen, B. Bromilow (WAM T44459); 8 ,
same locality, quadrat 7, 23°22’S 1 1 5°52’50"E,
/6/1994 (WAM 95/164); 28, Cape Range, site
TL- 1 , pitfall traps, 22 15'S. 1 1403'E. 16 May-5
June 90, J.M. Waldock (WAM 93/1030-3); 8 ,
same locality, site TL-7, 22 1 5'S, 1 14 04’E, 20
May-5 June 90 (WAM 93/1073-4); 8 , N-W
Cape Pcnin Site TL-10, 22°08’S, 1 14°05’E, 21
May-5 June 90, J.M. Waldock (WAM
93/1087-8); 8 . Cape Range, camp, 22°15’S,
1 14°03’E, 27 May 90, J.M. Waldock (WAM
93/1104).

FIG. 14. Euasteron spp. right male palps. A,B, E.johannae; A,
ventral; B, retrolateral; C,D, E. ursulae ; C, ventral; D,
retrolateral. Scale 0.5mm.

Epigyne (Fig. 17A,B): oval CO not attached to
epigastric fold, with short, V-shaped sclerotised
tubes, CD elongate coiled, axis in V-shaped
position. S touching.

DISTRIBUTION. Barlee Range, Western Australia
(Fig. 20).

Euasteron ursulae sp. nov.
(Figs 14C,D, 17C,D, 20)

DIAGNOSIS. Differs from E. atriceps
and E. Carnarvon by carapace sepia
brown, from E. johannae and E.
juliannae by femur I, II dark brown,
III, IV proximal part white, distal part
dark brown and by abdomen with 2
pairs of white patches on anterior part
and 3 in front of spinnerets.

DESCRIPTION. Male (holotype).
Total length 4.84; ceph 2.24 long, 1.56
wide, 1 high; cl/cw 1.43; sternum 1.12
long, 0.96 wide; sl/sw 1.16; abdomen
2.6 long, 1.44 wide.

Colour: carapace sepia brown; sternum
orange brown; chelicerae medium
brown; maxillae and labium medium
brown, distal ly white; abdomen sepia
brown; dorsally with 2 pairs of white
patches on anterior part and 3 in front of
spinnerets; legs medium brown, with
darker brown lateral stripes; clearly
annulated; coxae I-IV white; trochanter
I-IV brown; femur I, II dark brown, III, IV
proximal part white, distal part dark brown.

Eyes: AME largest; eye group width 0.64 of
headwidth; AME 0.16; ALE 0.12; PME 0.14;
PLE 0.14; AME-AME 0; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.04; AME-AME
0.32; PME-PME 0.32. Clypeus: 0.4mm high.

ETYMOLOGY. For Ursula Baehr, the author’s younger
daughter.

MATERIAL. HOLOTYPE: d, Barlee Range Nature
Reserve, WA, site BR6, dry pitfall traps 23°23’2rS,

Male palp (Fig. 14C,D): DTA, PE reaching tibia;
retrolaterally with membranous lamella; FA
short, rounded; SP S-shaped, narrow, crossing TF
totally; DtiA IP as long as EP, sharp, EP blunt;
VtiA bird-head-shaped.

22

MEMOIRS OF THE QUEENSLAND MUSEUM

Female (paratype). Total length 5.8;
ceph 2.6 long, 1.68 wide, 1.04 high;
cl/cw 1.54; sternum 1.12 long, 1 wide;
sl/sw 1.12; abdomen 3.2 long, 2.2 wide.
Colour: same as males.

Eyes: AME largest; eye group width
0.66 of headwidth: AME 0.16; ALE
0. 1 4; PME 0. 1 5; PLE 0. 1 5; AME-AME
0.04; AME-ALE 0.04; PME-PME 0.06;
PME-PLE 0.08; ALE-PLE 0.04; eyes
group AME-PME 0.04; AME-AME
0.36; PME-PME 0.36. Clypeus:
0.52mm high.

Epigyne (Fig. 17C,D): Oval CO
attached to the epigastric fold, with
V-shaped sclerotised tubes, CD
elongate coiled, axis in V-shaped
position. S touching.

DISTRIBUTION. Western Australia
(Fig. 20).

Euasteron juliannae sp. nov.

(Figs 13A,B, 17GH, 20)

ETYMOLOGY. For Ms Julianne Waldock of
the Western Australian Museum, the collector
of some of the types.

MATERIAL. HOLOTYPE: <3, Nerren Nerren
Station, WA, site NE 4, dry pitfall traps,
27°07’22"S, 1 14°47’58"E, 15-20 Oct 94, J.M.

Waldock et al. (WAM T44500). PARATYPES:

Western Australia: 3, Nerren Nerren Station,
site NE 3, wet pitfall traps, 27°07 , 24 M S,

1 1 4°46’4 1 "E, 25 Aug- 1 7 Oct 94, J.M. Waldock
et al. (WAM T44499); 3,9, near Wicherina
Dam. vehicle vibration, 28°43’49"S, 1 15°00M7 M E, 2 Nov
98, J.M. Waldock (WAM T44432); 3,9, same locality
(QM S60815); 63, Nanga Station, site NA 2, wet pitfall
traps 26 0 29’23 M S, 1 14°03 , 21 ,, E,23 Aug-160ct94, P. West
et al. (WAM T44488); 33, same locality, dry pitfall traps,
1 5-20 Oct 94 (WAM ex T44490); 9 , R.GC. Mine, 1 0km S
of Eneabba, site 3, 29°56’S, 115°17’E, 31 Oct 97, R.P.
McMillan, pitfall traps (WAM T44413).

DIAGNOSIS. Differs from E. atriceps and E.
Carnarvon by sepia brown ceph, from E.
johannae and E. ursulae by proximal part white,
distal part dark brown femur 1-IV. Abdomen dark
with pale chevrons. Palp, SP not crossing
transparent field. DTA retrolaterally with
lamellae and tiny prong. Epigyne copulatory
ducts extremely elongate, coiled about straight
axis.

DESC RIPTION. Male (holotype). Total length
4.68; ceph 2.36 long, 1.8 wide, 1.04 high; cl/cw

FIG 15. Euasteron spp. epigynes. A,B, E. enterprise ; A, ventral;
B, dorsal (cleared). C,D, E. monteithorum; C, ventral; D, dorsal.
E,F, E. milledgei ; E, ventral; F, dorsal. Scales 0.1mm. CO =
copulatory opening; CD = copulatory duct; S = spermatheca.

1.31; sternum 1.08 long, 1 wide; sl/sw 1.08;
abdomen 2.32 long, 1 .68 wide.

Colour: carapace sepia brown; with dark fovea
and dark bifurcate patches in front; sternum
orange brown; chelicerae medium brown;
maxillae and labium medium brown, distally
white; abdomen sepia brown; dorsal ly with pale
chevrons; laterally with 1 white patch; legs
medium brown, with darker brown lateral stripes;
clearly annulated; coxae I-IV white; trochanter
I-IV white; femur I-IV proximal part white, distal
part dark brown.

Eyes: AME largest; eye group width 0.57 of
headwidth; AME 0.16; ALE 0.14; PME 0.14;
PLE 0.14: AME-AME 0.02; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.4; AME-AME
0.34; PME-PME 0.34. Clypeus: 0.48mm high.

Male palp (Fig. 13A,B): DTA, PE not reaching
tibia; retrolaterally with membranous lamella

NEW AUSTRALIAN SPIDER GENERA

23

FIG. 16. Euasteron spp. epigynes. A,B, E. willeroo ; A,
ventral; B, dorsal (cleared). C,D, E.bartonr, C,
ventral; D, dorsal. E,F, E. krebsorum\ E, ventral; F,
dorsal. Scales 0.1 mm.

and tiny prong; FA short, with rounded tip; SP not
crossing TF; tibia short; IP short, EP blunt with
incision; VtiA hooked.

Female (paratype). Total length 6.2; ceph 2.6
long, 1.8 wide, 1.04 high; cl/cw 1.44; sternum
1.16 long, 1 wide; sl/sw 1.16; abdomen 3.6 long,
2.2 wide.

Colour: as in males.

Eyes: AME largest; eye group width 0.6 of
headwidth; AME 0.16; ALE 0.14; PME 0.14;
PLE 0.14: AME-AME 0.02; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.42; AME-AME
0.34; PME-PME 0.34. Clypeus; 0.52mm high.
Epigyne (Fig. 17G,H): Oval CO, attached to
epigastric fold U-shaped sclerotised tubes, CD
extremely elongate, about 4 X as long as wide,
coiled, axis straight. S touching.

DISTRIBUTION. Western Australia (Fig. 20).

Euasteron atriceps sp. nov.

(Figs 13C,D, 20)

ETYMOLOGY. Latin atriceps meaning dark-headed;
refers to the dark cephalic area.

MATERIAL. HOLOTYPE: < 5, Nerren Nerren Station,
WA, site NE2, 27°03'24"S, 114°35’21"E, 25 Aug-16 Oct
94, J.M. Waldock et al. t wet pitfall traps (WAM T44495).
PARATYPES: Western Australia: 1 8 <3 , same as holotype
(WAM T52336); 42 d, Nanga Station, site NA4,
26°32 , 47"S, 113°57 , 49"E, 16 Oct 94-19 Jan 95, N.
McKenzie, J. Rolfe, wet pitfall traps (WAM T44592); 2Y,
same locality (QM S60814); I5d, Nanga Station, site
NA5, 26°35’34 M S, 113°53 , 23 M E, 16 Oct 94-19 Jan 95, N.
McKenzie, J. Rolfe, wet pitfall traps (WAM T44593).

DIAGNOSIS. Differs from E. johannae , E.
ursulae and E.juliannae in carapace yellow with
dark brown cephalic area, abdomen with dark
margin and chevrons on top; from E. Carnarvon
in smaller size and SP not crossing TF.

DESCRIPTION. Male (holotype). Total length
3.48; ceph 1.8 long, 1.44 wide, 0.84 high; cl/cw
1.25; sternum 0.84 long, 0.72 wide; sl/sw 1.17;
abdomen 1 .68 long, 1.16 wide.

Colour: carapace yellow; with dark brown
cephalic area, with dark margin and chevrons on
top; sternum yellow; chelicerae medium brown;
maxillae and labium yellow, distally white;
abdomen pale; dorsally with dark chevrons;
ventrally mottled with white; legs pale brown;
with indistinct colour pattern; coxae I-IV pale;
trochanter I-IV pale; femur I-IV pale with brown
ring distally; tibia I-IV proximally and distally
part with brown spot.

Eyes: AME largest; eye group width 0.62 of
headwidth; AME 0.13; ALE 0. 1 ; PME 0.11; PLh
0 12; AME-AME 0.02; AME-ALE 0.02;
PME-PME 0.04; PME-PLE 0.06; ALE-PLE
0.04; eyes group AME-PME 0.3; AME-AME
0.28; PME-PME 0.26. Clypeus: 0.36mm high.
Male palp (Fig. 13C,D): DTA, PE not reaching
tibia; retrolaterally with ML; FA wider than long,
rounded; SP S-shaped, not crossing TF; DtiA IP
as long as EP sharp, EP blunt, VtiA
bird-head-shaped.

Female unknown.

DISTRIBUTION. Yamarna Station, WA (Fig. 20).

Euasteron Carnarvon sp. nov.

(Figs 6, 13E,F, 17E,F, 20)

ETYMOLOGY. From the type locality. A noun in
apposition.

24

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. Yl.Euasteron spp. epigynes. A,B, E.johannae ; A,
ventral; B, dorsal (cleared). C,D, E. ursulae ; C,
ventral; D, dorsal. E,F, E. Carnarvon ; E, ventral; F,
dorsal. QH, E.juliannae\ G, ventral; H, dorsal. Scale
0.1mm.

MATERIAL. HOLOTYPE: d , Nanga Station, WA, site
NA 1, diy pitfall traps, 26°28 , 40"S, 1 14°04’34"E, 15-20
Oct 94, P. West et al. (WAM T44575). PARATYPES:
Western Australia: 2d,l 9, Nanga Station, site NA 1, dry
pitfall traps, 26°28’40"S, 114°04’34"E, 15-20 Oct 94, P.
West et al. (WAM T5 1 854); d , 9 , same locality, site NA 1 ,

FIG 18. Records of Basasteron and Euasteron species
in Australia.

1 1 May-3 Aug 95, N. Hall (QM S6083 1 ); d , same locality,
site NA 5, 26°35'34"S, 1 1 3°53'23" E. P. West et al. (WAM
T44577); d, Zuytdorp, site ZU 1 wet pitfall traps,
27°15 , 42"S, 1 14°01 W'E, 26 Aug- 15 Oct 94, A. Sampey
et al. (WAM T445 1 1 ); 2d , Zuytdorp, site ZU 1 , wet pitfall
traps, 27°15 , 42"S, 1 14°01 WE, 15 Oct 94-1 1 Jan 95, N.
McKenzie, J. Rolfe (WAM T44512); 2d, Zuytdorp, site
ZU 2, wet pitfall traps, 27°15M1 M S, 114°0r48"E, 26
Aug- 15 Oct 94, A. Sampey et al. (WAM T44514); 2d,
same locality, 1 5 Oct 94-1 1 Jan 95, N. McKenzie, J. Rolfe
(WAM T445 1 5); d , same locality, site ZU 3, 27° 1 5’34”S,

1 14°04’03"E ) 26 Aug-17 Oct 94, A. Sampey et al. (WAM
T44518); d, same locality, site ZU 4, dry pitfall traps,
27°15’28 M S, 1 14°09 , 02 M E, 14-19 Oct 94, A. Sampey et al.
(WAM T44520); d,9, Barlee Range Nature Reserve,
quadrat 2, wet pitfall traps, 23°03 , 41”S, 1 1 5°47’ 1 5"E, Jun
94, S. van Lceuwcn, B. Bromilo (QM S60831).

DIAGNOSIS. Differs from E. johannae , E.
ursulae and E.juliannae in carapace yellow, dark
brown cephalic area and dark margin and
chevrons on top. From E . atriceps in bigger size
and SP partly crossing TF.

DESCRIPTION. Male (holotype). Total length
4.04; ceph 2.12 long, 1.64 wide, 0.92 high; cl/cw
1.29; sternum 1 long, 0.96 wide; sl/sw 1.04;
abdomen 1 .92 long, 1 .4 wide.

Colour: carapace yellow; with dark brown cephalic
area, with dark margin and chevrons on top;
sternum yellow; chelicerae medium brown;
maxillae; labium yellow, distally white; abdomen
pale; dorsally with dark chevrons; ventrally
mottled with white; legs pale brown; with

NEW AUSTRALIAN SPIDER GENERA

25

FIG. 19. Records of Euasteron species in Australia.

indistinct colour pattern; coxae 1-1V pale;
trochanter I-IV pale; femur I-IV pale with brown
ring distally; tibia I-IV proximally and distally
with brown spot.

Eyes: AME largest; eye group width 0.62 of
headwidth; AME 0.15; ALE 0.12; PME 0.13;
PLE 0.13; AME-AME 0.02; AME-ALE 0.02;
PME-PME 0.06; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.36; AME-AME
0.32; PME-PME 0.32. Clypeus: 0.4mm high.

Male palp (Fig. 13E,F): DTA, PE not reaching
tibia; retrolaterally with ML and short prong; FA
short, rounded tip; SP S-shaped, partly crossing
TF; tibia short; DtiA IP as long as EP, tapered, EP
blunt, pale brown; VtiA bird-head-shaped.

Female (paratype). Total length 4.04; ceph 2.12
long, 1 .64 wide, 0.92 high; cl/cw 1 .29; sternum 1
long, 0.96 wide; sl/'sw 1.04; abdomen 1.92 long,
1 .4 wide.

Colour: as in males.

Eyes: AME largest; eye group width 0.62 of
headwidth; AME 0.15; ALE 0.12; PME 0.13;
PLE 0.13; AME-AME 0.02; AME-ALE 0.02;
PME-PME 0.06; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.36; AME-AME
0.32; PME-PME 0.32. Clypeus: 0.4mm high.

Epigyne (Fig. 17E,F): Oval CO attached to the
epigastric fold, with U-shaped sclerotised tubes,
CD elongate 3 times as long as wide, coiled, axis
more or less straight. S touching.

DISTRIBUTION. Western Australia (Fig. 20).

DISCUSSION

The Asteron-oompitx now contains 1 1 genera
with 57 species. Asteron originally included only
A. reticulatum and A. mas Jocque, 1991.
However, A. mas is still not correctly placed in
any of the described genera. The first overview of
Euasteron indicated that A. mas should be a
member of this genus. A more careful exam-
ination showed that A. mas is more derived in
palpal structure and in the epigyne than any
Euasteron species.

In Basasteron, Euasteron and Spinasteron , the
abdominal pattern is an excellent character to
separate species if the spiders are well preserved.
The most common pattern is a dark abdomen
with 2-5 pairs of white patches on top, and 1 -3 in
front of spinnerets. It occurs also in all other
described genera in the ,4.ste/r>/?-complex as well
as in Habronestes and Storena (Jocque & Baehr,
1992).

As in Habronestes , the relative size of AME is
a very stable character to separate species-groups
in Euasteron. Using the eye pattern, Euasteron
can be divided into the E. enterprise-group (7
species) the AME are the smallest; E. willeroo
has equal-sized eyes; and the E. juliannae-gxoup
(7 species) have AME the largest.

Nevertheless, the definition of genera in
Australian zodariids has been mainly based on
genital characters (Baehr & Jocque, 1996, 2000,
2001; Jocque & Baehr, 2001).

During this work it became obvious that the
retrolateral structure of the cymbium is a good
synapomorphic character to separate taxa of the
Asteron-complex.

Basasteron shares the straight retrolateral
cymbial flange with (Fig. 3B) with more basic
Pentastem/u Phenasteron and Leptasteron. But
the general bulb structure of Basasteron — a
circular shape of the DTA and the shape of the
embolus — is basically the same as in
Cavasteron , Minasteron , Euasteron and
Spinasteron. Whereas in Basasteron , the
embolus originates prolaterally, in Cavasteron ,
Minasteron , Euasteron and Spinasteron , the
origin is basal.

The special type of tibial apophysis — with 2
apophyses forming a deep concavity in between,
dorsal one divided in two parts, the internal prong
(IP) and the external prong (EP) connected by a
ridge; ventro-lateral tibial apophysis hooked or
bird-head-shaped — is obviously a synapomorphy
of Cavasteron , Euasteron , Minasteron and

26

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 20. Records of Euasteron species in Western

Australia.

Spinasteron. It is not developed in Basasteron ,
which has only 2 small apophyses and a
beginning of a ridge. All these preliminary data
indicate that Basasteron is the sister genus of
Cavasteron , Euasteron, Minasteron , and
Spinasteron.

Euasteron can be easily separated from
Cavasteron and Minasteron by the lack of a large
terminal claw in male palps and by the more
elongate and curled copulatory ducts in the
epigynes.

Spinasteron shares the special tibial apophysis
structures and the special bulb characters with
Cavasteron , Euasteron and Minasteron. The
barely developed retrolateral cymbial flange with
only a tiny knob seems to be a reduction of the
flange with a well-developed rounded extension
(RE) (Fig. 4B,C) occurring in Cavasteron,
Euasteron, Minasteron. Spinasteron could be a
more derived sister genus to Euasteron. Spinasteron
is not fully revised yet. All these species have
very complicated palps.

ACKNOWLEDGEMENTS

n e rrace Y Churchill (formerly of the

C.S.I.R.O.), Michael Gray, Graham Milledge,
Australian Museum (Sydney), Mark Harvey,
Julianne Waldock, Western Australian Museum

FIG. 2 1 . Records of Spinasteron and Euasteron species
in Australia.

(Perth), David Hirst, South Australian Museum
(Adelaide) and Robert Raven, Queensland
Museum (Brisbane) for loan of the material and
support of the work. The following people of the
Northern Territory 1 would like to thank for
providing access to research sites: Dave Moller,
Manager of Willeroo Station; Don Cherry,
Manager of Kidman Springs Station; Paul Stone,
Manager of Mount Sanford Station, Heytesbury
Pastoral. For an excellent working atmosphere I
thank Valerie Davies, for being patient, my
children Johanna and Ursula Baehr, and for
preparing the maps and assisting with the SEM
Robert Raven.

LITERATURE CITED

BAEHR, B&JOCQUE, R. 1996. A revision of Asteron,
starring male palpal morphology (Araneae,
Zodariidae). Proceedings of the XIII International
Congress of Arachnology, Geneva, 3-8
September 1995. Revue suisse Zoology, hors
seriel: 15-28.

2000. Revisions of the genera in the Asteron -
complex (Araneae, Zodariidae). The new genera
Cavasteron and Minasteron. Records of the
Western Australian Museum 20: 1-30.

NEW AUSTRALIAN SPIDER GENERA

27

2001. Revisions of the genera in the Asteron-
complex (Araneae, Zodariidae). The new genera
Pentasteron, Phenasteron , Leptasteron and
Subasteron. Memoirs of the Queensland
Museum 46(2): 359-385.

DALLW1TZ, M.J., PAINE, T.A. & ZURCHER, E.J.
1998. Interactive keys. Pp. 201-212. In Bridge, P.,
Jeffries, P., Morse, D.R. and Scott, P.R. (eds)
Information technology, plant pathology and
biodiversity. (CAB International: Wallingford).

JOCQUfe, R., 1991. A generic revision of the spider
family Zodariidae (Araneae). Bulletin of the
American Museum of Natural History 20 1 : 1 - 1 60.

1995. Notes on Australian Zodariidae (Araneae), II.
Redescriptions and New Records. Records of the
Australian Museum 47: 141-160.

JOCQUE, R. & BAEHR, B. 1992. A revision of the
Australian spider genus Storena (Araneae,
Zodariidae). Invertebrate Taxonomy 6: 953- 1 004
2001. Revisions of the genera in the Asteron-
complex (Araneae, Zodariidae). A steron Jocque
and the New Genus Pseudasteron. Records of the
Australian Museum 53: 21-36.

RAINBOW, W.J. 1920. Arachnida from Lord Howe
and Norfolk Islands. Records of the South
Australian Museum 1: 229-272.

28

MEMOIRS OF THE QUEENSLAND MUSEUM

BLASTOBASINE COLEOPHORID MOTHS AS PREY
FOR THE AUSTRALIAN A RAN El I) SPIDER
CELAENIA CALOTOIDES RAINBOW. Memoirs of the
Queensland Museum 49(1): 28. 2003:- The Australasian
araneid Celaenia Thorell, and species of Ordgarius Keyscr-
ling, Cladomelea Simon and Mastophora Holmberg (‘bolas*
spiders), have unique life-history strategies. As older juvenile
and adult females, all are thought to be obligate predators of
Lcpidoptera, using complex chemical mimicry of female sex
pheromones to attract male moths of only a few species
(Eberhard, 1977; Stowe et al., 1987; Yeargan, 1988, 1994;
Haynes et al., 1 996). Young juveniles and males of some taxa
also attract male nematoceran Diptera (Eberhard, 1980;
Yeargan & Quatc, 1996, 1997). Stowe et al. (1987) showed
that 3 molecules released by M. comigera (Hentz) ([Z]-9-
tetradecenyl, [Z]-9-tetradecenal & [Z]- 1 1 -hexadecenal) are
identical to sex pheromone components of some prey moth
species. Gemeno et al. (2000) showed that M. hutehinsoni
Gertsch females produce an allomone blend that mimics, in
both composition and blend ratio, the sex pheromone of the
noctuid moth Lacinipolia renigera (Stephens) Web
reduction is characteristic of this group with a single line of
sticky silk used to capture flying moths. In Celaenia no web
snare is made, and moths are grasped directly with the
enlarged, spined legs 1 and II (pers. obs.).

Of 1 1 species of Celaenia (7 from Australia), few specific
prey records are available (Table 1 ). Notes on C. kinbergi are
listed under C. excavata L. Koch, the senior synonym.
Roberts (1937) collected several moths dropped by a female
C. excavata , and suggested ‘olfactory attraction* may be
involved, but did not identify any taxa. Mascord (1980)
reported that a female of C. excavata captured 43 male moths
‘of one species’ in one month.

Observations of two juvenile and one adult female C. calot-
oidcs Rainbow, 1908 were maintained over five weeks during
the Spring of 2002, in Brisbane. Cardboard trays and paper
bags were set up as receptacles under the spiders to catch
dropped prey items. The female (5mm long; with egg sacs)
was observed for 39 days (5 Oct.- 1 3 Nov.). Similarly, an older
juvenile (2.5mm long) was observed for 30 days ( 1 4 Oct.- 1 3
Nov.) along with a small juvenile ( 1 .5mm long, after having
hatched out of the female’s first egg sac on 20 Oct.), the latter
observed for 12 days between November 1 - 1 3, All specimens
were observed as they were found, in-situ in a suburban
garden (27 o 30’53’*S I53°04’06”E).

After laying its third egg sac on 7-8 Oct., the adult female C.
calotoides captured three male Blastobasis Zeller (Gelech-
ioidea, Coleophoridae, Blastobasinae) (Table 2). The larger
juvenile spider captured four Blastobasis moths during the
study period. The small ncwly-hatched juvenile spider caught
one moth during the study period, and this too was a
Blastobasis species. All specimens appear to be conspecific

TABLE 1 . Published prey records for spiders in Celaenia.

Spider

Loc.

Prey

Reference i

Adult 9 C.
excavata

Qld,

Aus.

Spodoptera mauritia
(Lepidoptera, Noctuoidea,
Noctuidae, Amphipyrinae)

Zillman,

1988

Adult 9
Celaenia sp.

NZ

‘Tortrix moths’ (Lcpidoptera.
Tortricoidea, Tortricidae)

Forster &
Forster, 1999

Juvenile
Celaenia sp.

NZ

‘Moth flies’ (Diptera,
Ncmatoccra, Psychodoidca,
Psychodidae)

Forster &
Forster, 1999

Adult 9C.
distincta

Tas..

Aus.

‘Night-flying moths’

Hickman,

1970

Adult 9C.

|| atkinsoni

Tas.,

Aus.

‘Small moths’

Hickman,

1970

(although the genitalia of some were damaged by the spiders
whilst feeding), and further observations of moths attracted to
house lights in the area revealed the presence of a single, very
common species (based on the uniform morphology of the
males' eenitalia). These data provide the first evidence of a
gelechioid moth being targeted by a species of Celaenia , and
the first record of a newly-hatched juvenile feeding on a
lepidopteran (as opposed to a dipteran) after emergence from

the egg sac. , , _ . .

Moths referred to herein are lodged at the Queensland
Museum, with the adult female C. calotoides (QM S60739).

Literature Cited

EBERHARD, W.G. 1977. Aggressive chemical mimicry by a bolas
spider. Science 198: 1 173-1 175.

1980. The natural history and behavior of the bolas spider
Mastophora JizruJeant sp. n. ( Araneidac). Psyche 87:143-1 69.
FORSTER RR.& F ORSTER, L.M. 1 999. Spiders of New Zealand
and their worldwide kin. (University of Otago Press: Dunedin).
GEMENO, C , YEARGAN, K.V. & HAYNES, K.F. 2000. Aggressive
chemical mimicry by thebolas spider Mastophora hutehinsoni.
identification and quantification of a major prey’s sex
pheromone components in the spider s volatile emissions.
Journal of Chemical Ecology 26(5): 1235-1243.

MASCORD. R. 1980. Spiders of Australia, a field guide. Reed

Publishers, Singapore. t

HAYNES, K.F.. YEARGAN, K.V.. MILLAR, J.G. & CHASTA N,
B.B. 1996. Identification of the sex pheromone of Tetanolita
mvnesalis (Lcpidoptera: Noctuidae), a prey species for the
bolas spider Mastophora hutehinsoni. Journal of Chemical
Ecology 22: 75-89. r\

HICKMAN, V.V. 1970. Three Tasmanian spiders of the genus C elaema
There! I (Araneida) with notes on their biology. Papers and
Proceedings of the Royal Society of Tasmania 105: 75-K2.
RAINBOW, W.J. 1908. Studies in Australian Araneidae. Records of
the Australian Museum 7(1): 44-46.

ROBERTS. N.L. 1937. Some notes on the bird-dung spider ( Celaenia
excavata) Proceedings of the Royal Zoological Society of
New South Wales: 23-28.

STOWE, M.K.. TUMLINSON, J.H. & HEATH, R.R. 1987. Chemical
mimicry: bolas spiders emit components of moth prey species
sex pheromones. Science 236: 964-967.

YEARGAN, K.V. 1988. Ecology of a bolas spider, Mastophora
hutehinsoni : phenology, hunting tactics, and evidence for
aggressive chemical mimicry. Oecologia 74: 524-530.

1 994. Biology of bolas spiders. Annual Review of Entomology 39:
81-99.

YEARGAN, K.V. & QUATE, L.W. 1996. Juvenile bolas spiders
attract psychodid flies, Oecologia 106: 266-271.

1997. Adult male bolas spiders retain juvenile hunting tactics.
Oecologia 112: 572-276.

ZILLMAN, E. 1988. Observations on the bird-dropping spider.
Queensland Naturalist 28(5-6): 28-31.

Michael G. Rix, Queensland Museum. PO Box 3300, South
Brisbane 4101. Australia ; 8 May 2003.

TABLE 2. Moth prey ( Blastobasis sp.) records for adull and
juvenile C. calotoides in Brisbane. Length is taken
longitudinally from head to wing tips, after specimens had
been wrapped with silk by spiders.

Prey

moth ref.

Spider

Length

Date captured

Al

6 adult

8.5mm

11-12/10/02

A2

3 adult

7.0mm

9-10/11/02

A3

6 adult

7.5mm

12-13/11/02

J1

6 older juvenile

5.5mm

22/10/02

12

6 older juvenile

5.5mm

24/10/02

J3

6 older juvenile

5.0mm

1-2/11/02

J4

6 older juvenile

6.5mm

5/11/02

B1

6 small juvenile

6.0mm

1 1/1 1/02

TROPASTERON GEN. NOV. OF THE TS7£/?6W-COMPLEX (ARANEAE:
ZODARI1DAE) FROM TROPICAL QUEENSLAND

BARBARA BAEHR

Baehr, B. 2003 06 30: Tropasteron gen. nov. of ihe Asteron-comp\ex (Araneae: Zodariidae)
from tropical Queensland. Memoirs of the Queensland Museum 49(1): 29-64. Brisbane.
ISSN 0079-8835.

Tropasteron contains 22 new species of the Asteron-complex, a large group of endemic
Australian Zodariidae. Tropasteron consists of two species-groups which are distinguished
by their size, slightly different palps and epigynes, and their different abdominal colour
pattern. The T. cleveland-group with 18 species includes the type species Tropasteron
Cleveland sp. nov. ( 6 , 9 ) and T. catxlwell sp. nov. ( 6 , 9 ), T. cooki sp. nov. ( 6 , 9 ), T. daviesae
sp. nov. ( d , 9 ), T. eacham sp. nov. ( 6 , 9 ), T. fox sp. nov. ( 6 , 9 ), T. Halifax sp. nov. ( 6 , 9 ), T.
heatherae sp. nov. ( 8 , 9 ), T. julatten sp. nov. ( 6 , 9 ), T. luteipes sp. nov. ( 6 , V ), T. malbon sp.
nov. ( 6 , 9 ), T. monteithi sp. nov. ( 6 , 9 ), T palmerston sp. nov. ( 6 ). T. raveni sp. nov. ( 6 , 9 ),
T. splendens sp. nov. (<J), T thompsoni sp. nov. (d,9), T. tribulation sp. nov. (d,9), T.
yeatesi sp. nov. (d,9). The larger and much more rare species of the Tropasteron
andreae - group arc represented by Tropasteron andreae sp. nov. (<3,9), T. magnum sp. nov.
( 6 ), T. pseudomagnum sp. nov. ( ) and T. robertsi sp. nov. ( 6 ). All species are found only in
the tropical rainforests of Queensland. □ Spiders, rainforest, Zodariidae , Tropastemn.

Barbara Baehr, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia
(e-mail: BarbaraB@qm.qld.gov.au) ; 20 October 2002.

This is the fifth systematic paper on the
Asteron-con\p\ex (Baehr & Jocque, 1996) of
zodariid spiders, that now contains 12 genera
with 79 species.

Like the rest of the /tomw-complex (Baehr, in
press; Baehr & Jocque, 2000, 2001; Jocque &
Baehr 2001 ) Tropasteron is endemic to Australia
and occurs almost exclusively in the rainforest
mountain systems of north Queensland. Most of
the 766 specimens were collected in wet tropic
surveys undertaken by G. Monteith, V. Davies, R.
Raven and other Queensland Museum staff, to
examine the broad patterns of species geography
across the area (Monteith, 1995). Target taxa
were insects with low vagility, e.g. Scarabaeidae
(Reid & Storey, 2000; Storey & Monteith, 2000),
Tenebrionidae (Bouchard, 2002), wingless
Carabidae (Baehr, 1995) and Aradidae
(Monteith, 1997). In addition to the
mygalomorph spiders (e.g. Raven 1979, 2000)
and some arancomorph spider groups of
Amphinectidae and Kababinae (e.g. Davies,
1995, 1999, 2000a, 2000b), Tropasteron
becomes the sixth Wet Tropics spider group to be
used for low vagility studies.

Tropasteron gen. nov., mentioned in Baehr &
Jocque (1996) as Asteron queenslandicum-
group, can be easily separated from all other
genera by a number of synapomorphies
mentioned in the genus description.

MATERIAL AND METHODS

Descriptions are generated with the aid of
DELTA and Intkey (Dallwitz et al., 1998) and
shortened where possible.

Drawings were done from body, right palp,
epigyne and vulva. All measurements are in mm.

Spination and colour patterns are dealt with as
in the revision of Euasteron (Baehr, this volume).

Divisions of the Wet Tropics used in the discussion
of distribution are the Rainforest Mountain Blocks
proposed by Monteith (1995) and modified in
Bouchard (2002).

Abbreviations of characters: ALE, anterior
lateral eyes; AME, anterior median eyes; AS,
anterior spinnerets; CD, copulatory duct; CE,
rounded extension on cymbium; Ceph, cephalo-
thorax; CO, copulatory opening; CR, chitinous
rim; DTA, dorsal tegular apophysis; DtiA,
dorsolateral tibial apophysis; E, embolus; EP,
external prong on dorso-retrolateral tibial
apophysis; F, flange on cymbium; Fe, femur; IP,
internal prong on dorso-retrolateral tibial
apophysis; MS, median spinnerets; MEG,
median eye group; MT, metatarsus; P, patella; PE,
prolateral extension of DTA; PLE, posterior
lateral eyes; PME, posterior median eyes; PS,
posterior spinnerets; S, spermatheca; sl/sw,
sternum length/width; SP, sperm duct; T, tibia;
Ta, tarsus; TF, transparent field on embolus base;
VtiA, ventral tibial apophysis.

30

MEMOIRS OF THE QUEENSLAND MUSEUM

Abbreviations of institutions where material is
deposited: AM, Australian Museum, Sydney (M.
Gray); QM, Queensland Museum, Brisbane (R.
Raven).

Abbreviations of common collector names: DC,
D.J. Cook; DY, D. Yeates; GM, G.B. Monteith;
GT, G.I. Thompson; HJ, H. Janetzki; JG, J.
Gallon; MS, M. Shaw; RR, R.J. Raven; PL, P.
Lawless; SH, S. Hamlet; VD, V. Davies.

SYSTEMAT1CS

KEY TO SPECIES GROUPS OF

TROPASTERON

1 . Clypeus at least 5.5 times diameter of ALE high. Palp,
(EP) distally flattened, cymbial flange short straight,
with distal hook, only about 10% of cymbium long,
epigync CO lyriform (Fig. 6G-J) . . . T. andreae-group
Clypeus less than 5 x diameter of ALE high. Palp, (EP)
with sharp tip, cymbial flange straight, without distal
hook, about 15% of cymbium long, epigyne CO
semicircular or oval (Figs 5A-L, 6A-F) . . T.
Cleveland- group

KEY TO SPECIES OF T
CLEVELAND-G ROUP

Males

1. Palp without LTA spine (Figs 9B) 2

Palp with LTA spine (Fig. 1C) 3

2. Palp, DtiA, EP short, about as long as wide at base;

abdomen with 3 pairs of big white patches dorsally (Fig.
5D,9A-F) . T.daviesae

Palp, DtiA, EP long, EP at least twice as long as wide at
base; abdominal pattern not as above (Fig. 6A; 1 8A-D)
T.palmerston

3. Abdomen frontally without any white or pale spots (Fig.

1A) 4

Abdomen frontally with pairs of white spots (Fig. 22C) . 1 0

4. Palp DtiA, EP short, about as long as wide at base (Fig.

5A,D,F,6B) .5

Palp DtiA, EP long, EP at least twice as long as wide at
base (Fig. 5B,C,E,G-L) 7

5. Palp, with short LTA spine, about as long as wide; sperm
duct semicircular; abdomen with 2 pairs of white dots

dorsally (Fig. 19A-D) T.raveni

Palp, with long LTA spine, at least 3 times as long as wide;
sperm dupt inverted U-shaped (Fig. 1C, 1 IB) 6

6. Abdomen with 2 pairs of white dots dorsally, second pair

forming a transverse stripe (Fig. 1 1A-D) T.fox

Abdomen with 2 pairs of white dots dorsally, without
transverse stripe (Fig. 1 A-C) T. Cleveland

7. Abdomen ventrally without any colour pattern, dorsally

with 2 pairs of white dots (Fig. 12 A-D) . . . . T. Halifax
Abdomen ventrally otherwise (Figs 8D, 17D.23D, F) . 8

8. Abdomen ventrally with 1 pair of white stripes (Fig.

17A-D) T.monteithi

Abdomen ventrally with white chevrons (Fig. 1 A-C) . 9

9. Abdomen with light patches arranged as in Fig. 8C, D

T.cooki

Abdomen with light patches arranged as in Fig. 23C-F
T.yeatesi

10. Abdomen frontally with 1 additional pair of fronto-lateral

white spots (Fig. 23C-F) 1 1

Abdomen frontally without additional pair ol
fronto-lateral white spots (Fig. 2 1 C-D) 14

1 1 . Palp, with short LTA spine (Fig. 14B) .... T.julatten
Palp, with long LTA spine, at least 3 x as long as wide

(Fig. 10B,15B,2IB) 12

12. Abdomen frontally with 1 pair of white spots additional to

1 fronto-lateral pair (Fig. 21 A-D) I thompsoni

Abdomen frontally with more than 1 pair of white spots
additional to 1 fronto-lateral pair (Figs 10C, 15C) . . 13

1 3. Abdomen frontally with 2 pairs of white spots additional

to 1 fronto-lateral pair (Fig. 10C) T.eacham

Abdomen frontally with 3 pair of white spots additional to
1 fronto-lateral pair (Fig. 15C) T. luteipes

14. Palp with short LTA spine (Fig. 16B) T.malbon

Palp with long LTA spine, at least 3 times longer than

wide (Figs 7B,13B,20B,22B) 15

15. Abdomen frontally with 1 pair of white spots (Fig. 7A-D)

T.cardwell

Abdomen frontally with more than 1 pair of white spots
(Figs 13C, 12C,22C) 16

1 6. Abdomen frontally with 3 pairs of white spots (Fig. 22A)

T. tribulation

Abdomen frontally with 2 pairs of white spots (Figs 20C,
13C) * 17

17. Abdomen dorsally with 2 pairs of white spots and 2 in

front of spinnerets, additional to frontal spots (Fig.

20A-D) . T.splendens

Abdomen frontally with 4 pairs of white spots, dorsally
with 2 pairs and 2 in front of spinnerets, additional to
frontal spots (Fig. 13 A-D) T. heatherae

Females

1. Abdomen frontally without any white spots (Fig. 1 A) . 2

Abdomen frontally with pairs of white spots (Fig. 22C) . 9

2. Abdomen dorsally with 2 pairs of white dots (Fig. 1 A) . 3

Abdomen dorsally otherwise 5

3. Femur I dark brown, II-I V proximal pari white, distal part

brown epigyne (Fig. 3B,C) T. Cleveland

Femur I-IV same colour pattern 4

4. Femur I-IV proximal margin brown, medium part white,

distal part brown, epigync longer than wide, CO

semicircular(Fig. 19C-G) T.raveni

Femur I-IV proximal part white, distal part brown, epigyne
wider than long, CO oval (Fig. 12C-G) .... T. Halifax

5. Abdomen ventrally without any colour pattern, epigyne,

CO flattened oval (Fig. 1 1 C-F) T.fox

Abdomen ventrally with colour pattern 6

6. Abdomen ventrally with 1 pair of white stripes; epigyne

(Fig. 17C-F) T.monteithi

Abdomen ventrally with chevrons (Figs 8D, 9D, 23 D,F) . 7

7. Legs pale without colour pattern, small about 3.5mm;

epigyne (Fig. 9C-F) T. daviesae

Legs with indistinct colour pattern or clearly annulated . 8

8. Abdomen laterally with 1 long white stripe, epigyne CO

flat oval (Fig. 23C-H) T.yeatesi

Abdomen laterally with white chevrons, epigyne CO
semicircular (Fig. 8C-F) T.cooki

9. Abdomen frontal with 1 additional pair of fronto-lateral

white spots (Figs 1 0C, 1 4C, 1 5C, 2 1 C) 10

NEW QUEENSLAND SPIDER GENUS

31

Abdomen frontal without fronto-lateral white spots (Figs
7C, 13C, 16C,22C) ... 13

10. Abdomen ventrally without any colour pattern (Figs 14D,

11

Abdomen ventrally with white stripes or patches (Fig.

21D ) 12

1 1 . Abdomen frontally with 2 pairs of small white spots,

epigyne as in Fig. 1 4C-F T. julatten

Abdomen frontally with 3 pairs of small white spots,
epigyne as in Fig. 1 5C-F 7! luteipcs

12. Abdomen ventrally with 3 white stripes, epigyne as in

Fig. 21C-F T.thompsoni

Abdomen ventrally with 1 pair of white patches near

tracheal spiracle, epigyne as in Fig. 10C-E. . T.eacham

13. Abdomen frontally with 1 pair of small white spots (Figs

7C-F, 16C-F) 14

Abdomen frontally with 2-3 pairs of small white spots,
epigyne as in Figs 1 3C-F, 22C-F 15

14. Epigyne, CO semicircular (Fig. 16C-F). . . . T.malbon
Epigyne, CO flattened oval (Fig. 7C-F) . . . T. cardwell

15. Abdomen frontally with 2 pairs of small white spots,

epigyne ( Fig. 13C-F) T.heatherae

Abdomen frontally with 3 pairs of small white spots,
epigyne (Fig. 22C-F) T. tribulation

Tropasteron gen. nov.

TYPE SPECIES. Tropasteron Cleveland sp. nov.

ETYMOLOGY. Occurs only in tropical eastern Queens-
land, from Iron Range (12°40’S) to Carmila (21°52’S).

DIAGNOSIS. Tropasteron gen. nov., mentioned
in Baehr & Jocque (1996) as Asteron queens -
landicum- group, can be easily separated from all
other genera by the synapomorphies:

1 ) the very short, straight cymbial flange, with
only 10-15% length of cymbium and the small
rounded extension CE (Fig. IB);

2) the long spine-like LTA (Fig. 3C), the more or
less cylindrical base of the embolus (Fig. 3B);

3) the prolateral extension (PE) of semicircular,
distally-folded, dorsal tegular apophysis (DTA),
short not longer than base of DTA (Fig. 3B);

4) the ventro-lateral tibial apophysis with
chitinous rim (CR) (Fig. IB);

5) the bifurcate dorsolateral palpal tibial
apophysis, internal prong (IP) about 1/3 of
external prong (EP) (Figs 5A-L, 6A-J);

6) the more or less square epigyne with semi-
circular to flat oval copulatory opening (CO), not
attached to the epigastric fold, copulatory duct
(CD) elongate with big semicircular distal loop,
curled and ending in not touching spemathecae
(S) (Fig. 2B,C);

7) the strong female palpal claw with more than 6
teeth (Fig. 4B).

Although Tropasteron is well separated by
these synapomorphies, all species are similar in
their palpal and epigynal structure. Therefore,
unfortunately, the abdominal and leg colour
pattern are often the only characters to easily
separate species.

DESCRIPTION. Medium-sized spiders (3.00-
7.50) with finely reticulate tegument. Carapace
widest at coxae II. Profile flat with highest point
between fovea and PME (Fig. 1 A,D).

Colour: carapace, sternum yellow to sepia
brown; chelicerae yellow to medium brown;
maxillae and labium yellow to medium brown,
distally white; abdomen dorsally sepia brown
with or without weak scutum with 2-4 pairs of
white patches on anterior part and 1 -3 in front of
spinnerets; frontally with or without 1-3 pairs of
small white spots, frontolaterally with or without
1 pair of small white spots; laterally with 2-4
white stripes or mottled with white; ventrally
paler brown to dark brown, mottled with white;
with or without with 1 pair of elongate white
stripes and or 1 white stripe ifom epigastric fold
to near tracheal spiracle, legs yellow, pale to
medium brown, indistinct or clearly annulated.

Carapace: broadly oval, finely reticulated.
Sternum: shining; heart-shaped anteriorly straight;
with short and longer setae.

Eyes (Fig. 1A,D,E): in 3 rows (2-4-2). AME
smallest; eye group width 0.46-0.69 of
headwidth. MEG slightly longer than wide.
Clypeus straight, about 5.5-6 times (T andreae-
group) and 3-4.4 times ( T. Cleveland- group)
diameter of ALE high. Chilum single or divided.
Chelicerae as usual for family with few hairs in
front and dense row on distal promargin; no teeth.
Maxillae and labium triangular; sparsely haired;
maxillae with anteromesal scopula. Sternum flat;
heart-shaped with straight anterior margin,
shining; with long setae around margin.
Mouthparts: chelicerae, basal segment laterally
with condyle; maxillae triangular, with
promarginal scopula; labium triangular.
Abdomen: PLS small; widely spaced with PMS
in one row; PMS tiny; colulus only some hairs;
tracheal spiracle small slit-like, with a tiny pit
(Fig. 3A).

Legs: Formula 4123. Spination: few spines on
pairs I, II, more numerous on III, IV. Paired tarsal
claws with numerous (±14) teeth on inner side of
claw. Unpaired claw toothless, on very small
onychium. Metatarsal preening brush on Mt II
and III poorly developed.

32

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 1. Tropasteron Cleveland male. A, body dorsal; B,C, plap; B, lateral, C, ventral, scale 0.5mm, D,E,
cephalothorax; D, lateral; E, frontal, scale 1 mm. ALE = anterior lateral eyes. AME - anterior median eyes, CE =
rounded extension on cymbium, CH - hook on cymbial flange, C-R - chitinous rim, DTA - dorsal tegular
apophysis, DtiA = dorsolateral tibial apopysis, E = embolus, F = flange on cymbium, LTA = lateral tegular
apopysis (spine), MEG = median eye group, PE = prolateral extension of DTA, PLE = posterior lateral eyes,
PME = posterior median eyes, SP = sperm duct, TF = transparent field on embolus base, VtiA = ventral tibial
apophysis.

Male palp (Fig. 1B,C): tibia short with dorso-
lateral and ventrolateral apophysis. Dorsolateral
tibial apophysis bifid with sharp internal (IP) and
external prong (EP). In T. andreae-gxowp EP
distally flattened. Cymbial flange short straight,
only 10-15% of cymbium long, distally followed
by small rounded extension, incision in between.

DTA, semicircular, distally folded with short
PE, not longer then base of DTA; retrolaterally
with retrolaterally with sharp tip and without any
additional extension; sperm duct inverted
U-shaped; embolus thin, semicircular, embolus
base cylindrical; tibia short; ventrolateral tibial
apophysis with chitinous rim dorsally, connected
with dorsolateral apophysis. Femur with 2 strong
spines dorsally. Embolus semicircular conducted
by DTA.

Females are often bigger than males but body
shape and colour pattern are the same. In contrast
the female ceph is higher and clypeus is bowed,
visible from dorsal view (Fig. 2A,D,E). Epigynes
(Fig. 2B, C) of all Tropasteron species are very
similar: all are as long as wide, with semicircular
oval or lyriform CO, not attached to the epigastric

fold, CD elongate with big semicircular distal
loop, curled and ending in not touching S.

Tropasteron Cleveland sp. nov.

(Figs I, 2, 3, 4, 5 A. 29)

ETYMOLOGY. The species name is taken from the type
locality. A noun in apposition.

MATERIAL. HOLOTYPE: d, Mt Cleveland, NE Qld,
500m, blady grass, open forest. 19 0 15’23”S, 147°0r40”E,
22 Jan-21 Mar 1991, A. Graham, pitfall (QM SI 7982).
PARATYPES: NE Qld: 1 1 f, 19m, same data as holotype
(QM S57097); 2m, same locality, 20 Jan- 12 Mar 1991
(QM SI 7972); 1 f, as previous, 19°16’S, I47°03’E, Jan-12
Mar 1991 (QM S46780); 1m, as previous, summit, 560m,
open forest. 19°15’23"S, !47°01’40”E, 22-24 Mar 1991,
GM, DC (QM SI 7928); 2f, 6m, as previous, 500m,
rainforest, 19°I5'23”S, 147°0L40”E, 20 Jan-12 Mar
1 99 1 , A. Graham (QM S 1 7902).

DIAGNOSIS. Differs from most other species by
short EP (Fig. 5D). Similar to T raveni in the
abdomen lacking white spots frontally, but
differs by femur I dark brown, II-IV proximal
part white, distal part dark brown.

NEW QUEENSLAND SPIDER GENUS

33

FIG 2. Tropasteron Cleveland female. A, carapace; B,C, epigyne; B, ventral, C, dorsal, scale 0.25mm; D,E,
cephalothorax; D, lateral; E, frontal, scale 1mm. CD = copulatory duct, CO - copulatory opening, S
spermatheca.

DESCRIPTION. Male (holotype). Total length

4. 1 2. Ceph 2.08 long, 1 .56 wide, 0.84 high; cl/cw

1.33; sternum 1.04 long, 0.96 wide; sl/sw 1.08;

abdomen 2.04 long, 1 .44 wide.

Colour; Carapace orange brown. With dark fovea
and dark bifurcate patches in front; sternum
orange brown; chelicerae orange; maxillae and
labium medium brown, distally white. Abdomen
(Fig. 1A) sepia brown; dorsally with 2 pairs of
white patches on top and 2-3 in front of the
spinnerets; lateral with 1 white patch; ventrally
dark brown. Legs medium brown, with darker
brown lateral stripes; coxae I -IV white;
trochanter I-IV brown; femur I dark brown, 11-IV
proximal part white, distal part dark brown; tibia
I distally white.

Eyes: in 2 rows with 4 eyes, both rows procurved;
AME smallest. PLE largest; eye group width 0.69
of headwidth; AME 0.10; ALE 0.12; PME 0.13;
PLE 0.14; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.34; AME-AME
0.24; PME-PME 0.30. Clypeus 0.48 high; chilum
undivided.

Male palp (Figs 1 B,C, 3C): tegulum (LTA) a long
spine, at least twice as long as wide; EP about as
long as wide at base (Fig. 5A).

Female (paratype). Total length 4.92. Ceph 2.44
long, 1.68 wide, 0.96 high; cl/cw 1.45; sternum
1.08 long, 1.00 wide; sl/sw 1.08; abdomen 2.48
long, 1 .92 wide.

Colour: As in male.

Eyes: AME smallest; PLE largest; eye group
width 0.53 of headwidth; AME 0.10; ALE 0.12;
PME 0.14; PLE 0.14; AME-AME 0.04;
AME-ALE 0.04; PME-PME 0.06; PME-PLE
0.12; ALE-PLE 0.04; eyes group AME-PME
0.36; AME-AME 0.24; PME-PME 0.34. Clypeus
0.60 high.

Epigyne: (Fig. 2B,C) with oval CO.

DISTRIBUTION. Queensland (Fig. 29). Known
only from the summit of Mt Cleveland, a small,
isolated mountain on the coast 25km E of
Townsville. Open forest.

Tropasteron cardwell sp. nov.

(Figs 5B, 7, 29)

ETYMOLOGY. The type locality, a noun in apposition.

34

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 3. Tropasteron Cleveland male. A, pitted shield covers tracheal spiracle; B, palp ventral; C, LTA spine; D, tip
ofeymbiurn; E, tarsal claw 1st right leg.

MATERIAL. HOLOTYPE; d, Cardwell Ra. NE Qld,
Upper Broadwater Ck Valley, 700m, rainforest. 18°I8‘S,
145°56’E, 18 Dec 1986-14 Jan 1987, GM. GT. SH. pitfall
(QM S 1 2235). PARATYPES: NE Qld: 29,1 <3 , same data
as holotype (QM S57093); 1 9,1 d, same dam as holotype
(AM KS83917); 2d, Cardwell Ra, Mt Macalister, 700m,
rainforest, 18°19'S, 145°5 , E,20Dec 1 986, GM. GT, moss
on trees and rocks (QM S25692); 9d, same locality ,
18°18;S, I45°56*E, 18 Dec 1986-14 Jan 1987, GM. GT,
SH, llight intercept trap(QM S41929); 2d, Upper Boulder
Ck, 8km N ofTuIly, 250m, rainforest, 17°05’S, I45°54’S,
4-7 Dec 1989, GM, GT, HJ, pitfall (QM S46826); Id,

Boulder Ck. Walter Hill Ra, rainforest, 250-600m,
17.05’S, 145.54’E, 24-27 Oct 1983, GM, DY, GT (QM
S3808); 1 d , Broadwater Pk. 35km NW Ingham, rainforest,
18°22'S, 145°57’E, 21-22 Dec 1986, SH (QM S12239);
1 d, as previous, 18°22’S, I45°57 T E, 22 Dec 1986-3 Jan
1987, SH. pitfall (QM SI 2238); 7d, Douglas Ck Rd,
Kirrama SF, 800m, rainforest, 18°12’S. MS^’E, 10 Dec
1986-1 1 Jan 1987. GM. GT, SH, flight intercept trap (QM
SI 2240); 7d, as previous, 26-30 Aug 1987, RR, pitfall
(QM S 12242); 9d, Mt Smoko Rd. Kirrama Ra, 700m,
rainforest, 18 o 09'S, 145°37 , E, 10 Dec 1986-11 Jan 1987,
GM, GT, SH, pitfall (QM SI 8 179); 2d, Kirrama Ra (Mt

NEW QUEENSLAND SPIDER GENUS

35

I 1G. 4. Tropasteron Cleveland female. A, tarsal claw 1 st right leg B, palpal claw with teeth; C, pitted shield covers
tracheal spiracle; D, CO of epigyne.

Hosie), 930m, rainforest, 1 8° IPS, 145°45’E, ll-Dec-86,
GM, GT, Berlcsate (QM S34852); 4c5, Kirrama Ra, Main
Rd, west side. 700m, rainforest, 18°12'S, 145°45’E, 10
Dec 1 986- 1 1 Jan 1 987, GM. GT, SH, pitfall (QM S 1 2263 );
29,2(5, Kirrama Ra, 700m, 18°06’S, 145°42'E, 9 Dec
1986, GM. GT, sieved litter (QM S25683); 26, Kirrama
Ra, Yuccabine Ck, 700m, I8 0 12'S, 145°45’E, 10 Dec-
1986, GM, GT. sieved litter (QM S25694): 26, Kirrama
Ra. Douglas Ck Rd, 800m, 18°12'S, 145°45’E, 10 Dec
1986-11 Jan l987,GM,GT,SH,pitfall(QMS18177);2c5,
Gayundah Ck, Hinchinbrook Is. 10m, rainforest, 18°22 , S,
146°15 , E, 8-17 Nov 1984. VD, JG, pitfall (QM S25791);
19,9d, as previous (QM S25721): 1(5, as previous,
18°22’S, 146°13’E, 8-18 Nov 1984, GM, GT, DC, pitfall
(QM S3 800).

DIAGNOSIS. Very similar in colour pattern and
body shape to T. splendens and T. malbon , differs
in colour pattern by: abdomen without weak
scutum and ventrally with 1 pair of small white
spots. T. cardwell has shorter legs and different
shape of palp with long spine, at least twice as
long as wide (LTA).

DESCRIPTION. Male (holotype). Total length
4.24. Ceph 2.20 long, 1 .60 wide, 1 .00 high; cl/cw
1.37; sternum 1.08 long, 0.96 wide; sl/sw 1.12;
abdomen 2.04 long, 1 .32 wide.

Colour: carapace orange brown. With dark fovea
and dark bifurcate patches in front; sternum
orange brown; chelicerae orange; maxillae and

36

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. A-L, tibia (dorsal view) ofTropasteron spp; A,
T. Cleveland ; B, T. cardwell ; C, I cooki ; D. T.
daviesae ; E, T. cacham ; F, I fox; G, T. Halifax ; H, T
heatherae ; I, T. julatten ; J. T. luteipes; K, T. rnalbon ;
L, T. monteithi. Scale 0.25mm. EP = external prong
on dorso-retrolateral apopysis; IP = internal prong on
dorso-retrolateral apopysis.

labium medium brown, distally white. Abdomen
(Fig. 7C, D) sepia brown; frontally with 1 pair of
small white dots; dorsally with 2 pairs of white
patches on top and 2-3 in front of the spinnerets;
lateral with 1 white patch; ventral ly dark brown;
with 1 pair of white patches near tracheal
spiracle. Legs pale brown, with darker brown
lateral stripes; coxae I-IV w'hite or pale;
trochanter I-IV brown; femur I-IV proximal part
white becoming continuously dark brown
distally; tibia I distally white.

Eyes: AME smallest, PLE largest; eye group
width 0.60 of headwidth; AME 0.08; ALE 0.12;
PME 0.13; PLE 0.14; AME-AME 0.04;
AME-ALE 0.04; PME-PME 0.04; PME-PLE
0.10; ALE-PLE 0.04; eyes group AME-PME

FIG. 6. A-J, tibia (dorsal view) of Tropasteron spp; A,
T. palmerston ; B, T. raveni ; C, T splendens ; D, T.
thompsoni ; E, T. tribulation ; F, T. yeatesi; G, T.
andreae ; H, T magnum; I, T. pseudomagnum; J, T.
robertsi. Scale 0.25mm.

0.36; AME-AME 0.20; PME-PME 0.30.
Clypeus 0.52 high.

Male palp (Fig. 7A, B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct
semicircular; EP at least twice as long as wide at
base (Fig. 5B).

Female (paratype). Total length 4.20. Ceph 2.20
long, 1.44 wide, 0.96 high; cl/cw r 1.52; sternum
0.92 long, 0.88 wide; sl/sw 1.04; abdomen 2.00
long, 1.44 wide.

Eyes: AME smallest; eye group width 0.57 of
headwidth; AME 0.08; ALE 0.12; PME 0.13;
PLE 0.13; AME-AME 0.04; AMI '-ALL 0.04;
PME-PME 0.04; PME-PLE 0.12; ALE-PLE
0.04; eyes group AME-PME 0.38; AME-AME
0.20; PME-PME 0.30. Clypeus 0.52 high.
Epigyne (Fig. 7E, F): Wider than long, w ith very
flat, broadly oval CO (Fig. 7E, F).
DISTRIBUTION. Queensland, Wet Tropics
blocks 1 2, 1 3 and 1 5 (Fig. 29). Occurs widely on
the Kirrama Range, on the eastern rims of the
Cardwell and Walter Hill Ranges, and on nearby
Hinchinbrook Island. Rainforest.

NEW QUEENSLAND SPIDER GENUS

37

FIG. 7. Tropasteron cardwell. A,B, palp; A, lateral; B, ventral; scale 0.25mm; C,D, abdomen; C, dorsal; D,
ventral; scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

FIG. 8. Tropasteron cooki. A,B, palp; A, lateral; B, ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral;
scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

Tropasteron cooki sp. nov.

(Figs 5C, 8, 3 IB)

ETYMOLOGY. For Douglas Cook of the Queensland
Museum, one of the collectors.

MATERIAL. HOLOTYPE: 6, North Bell Peak, Malbon
Thompson Ra, NE Qld, 17°07’S, 145°53’E, 20-22 Nov
1990, GM, GT, pitfall (QM S57007). PARATYPES: NE
Qld: 1 9 ,7 6 , as for holotype (QM S25696); 1 9 , 1 6 , Bell

Peak North, 10km E Gordonvale, 850- 1000m, rainforest,
17°06'S, I45°53’E, 13 Oct 1982, GM, DY. GT, sieved
litter (QM S4235); 2 9 , 1 6 , Mt Edith Rd. Lamb Ra., 900m,
rainforest, I7°06’S, 145°37’E, 12 Oct 1982, GM, DY, GT.
sieved litter (QM S3798); 39,1 <3, North Bell Peak, via
Gordonvale. 900m, rainforest, 17 o 05'S, 145 0 53'E, 16 Sep
81, GM, DC (QM S3766); 56, North-South Bell Pk
saddle, Malbon Thompson Ra, 17°07’S, 145°54’E, 20-21
Nov 1990, GM, GT, pitfall (QM S25788).

38

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 9. Tropasteron daviesae. A,B, palp; A, lateral; B, ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral;
scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

DIAGNOSIS. T. cooki is similar to T. yeatesi but
differs from all other species by abdomen without
frontal spots and with 4 pairs of white patches on
top. Differs from I yeatesi in only 2 spots in front
of the spinnerets. Male palp with short spine
(LTA), about as wide as long.

DESCRIPTION. Male (holotype). Total length
4.08. Ceph 2.00 long, 1 .60 wide, 1 .04 high; cl/cw
1.25; sternum 1.04 long, 0.88 wide; sl/sw 1.18;
abdomen 2.08 long, 1 .36 wide.

Colour: carapace orange brown; sternum orange
brown; chelicerae medium brown; maxillae and
labium medium brown, distallv white. Abdomen
(Fig. 8C,D) sepia brown; dorsally with 4 pairs of
white patches on top and 2 in front of the
spinnerets; lateral with chevrons; ventrally pale
brown; mottled with brown. Legs pale brown to
medium brown; coxae I-IV white; trochanter
I-IV brown; femur I-IV proximal part white,
distal part dark brown.

Eyes: AME smallest, PLE largest; eye group
width 0.58 of headwidth; AME 0.09; ALE 0. 14;
PME 0.13; PLE 0.16; AME-AME 0.04: AME-
ALE 0.03; PME-PME 0.04; PME-PLE 0.10;
ALE-PLE 0.02; eyes group AME-PME 0.34;
AME-AME 0.20; PME-PME 0.30. Clypeus 0.44
high; chilum divided.

Male palp (Fig. 8A,B): tegulum (LTA) a short
spine, about as wide as long; sperm duct inverted
U-shaped; EP at least twice as long as wide at
base (Fig. 5C).

Female (paratype). Total length 6.12. Ceph 2.76
long, 1 .96 wide, 1 .36 high; cl/cw 1 .40; sternum
1.00 long, 1.00 wide; sl/sw 1.00; abdomen 3.36
long, 2.56 wide.

Colour as in male.

Eyes: AME smallest; eye group width 0.54 of
headwidth; AME 0.09; ALE 0.14; PME 0.14;
PLE 0.14; AME-AME 0.02; AME-ALE 0.06;
PME-PME 0.06; PME-PLE 0.16; ALE-PLE
0.06; eyes group AME-PME 0.40; AME-AME
0.20; PME-PME 0.34. Clypeus 0.68 high.
Epigyne: wider than long, anteriorly wider, with
semicircular CO (Fig. 8E,F).

Variation: Females are bigger than males and
have more chevrons.

DISTRIBUTION. Queensland, Wet Tropics
blocks 7 and 11 (Fig. 3 IB). Known only from
high elevations on the Lamb Range and the Bell
Peak region of the Malbon Thompson Range.
Rainforest.

Tropasteron daviesae sp. nov.

(Figs 5D, 9, 30B)

ETYMOLOGY. For Dr Valerie Davies, the main founder
of the spider collection of the Queensland Museum.

MATERIAL. HOLOTYPE: 6 , Upper Boulder Ck, Walter
Hill Ra, NE Qld, 850-1000m, rainforest, 17°05’S,
145°54’E, 17-18 Nov 1984, VD, GM, JQ DC, GT, pitfall
(QM S57092). PARATYPES: NE Qld: 1 same data as
holotype but, 900m, 27 Oct 1983, GM, DY, GT, sieved
litter (QM S3806); 1 <3, as previous, 200-650m, 24-27 Oct

NEW QUEENSLAND SPIDER GENUS

39

FIG. 10. Tropasteron eacham. A,B, palp; A, lateral; B, ventral; scale
0.5mm; C, abdomen dorsal; scale 1 mm; D,E, epigyne; D, ventral; E,
dorsal; scale 0.25mm.

1983 (QM S3787); 2d, as previous, 850- 1000m, 17-18
Nov 1984, VD, GM, DC, C.T, pitfall (QM S3785); 1 d, as
previous (QM S4266); 1 d, Maalan SF (NQ 12), 17°35’S,
145°36’E, 26 Nov 1 992-15 Apr 1993. R., J. & S. Raven, P.
& E. Lawless, pitfall (QM S24368); 1 d. U Boulder Ck,
10km N Tully, 800m, 17°05'E, 145°54’S, 4-7 Dec 1989,
GM, GT, HJ, pitfall (QM S41523); 1 9,12d, Maalan SF
(NQ 12), 17°35’30”S, 145°36'45"E,25 Jul 1992-26 Nov
1992, RR, P. & E. Lawless, MS, pitfall (QM S24486); 1 d,
as previous (AM KS 83918)..

DIAGNOSIS. One of the smallest species, it
differs from all others by Ep very short (Fig. 5D),
no palpal spine (LTA) and abdomen with 3 pairs
of big pale dorsal spots but without frontal spots.

DESCRIPTION. Male (holotype). Total length
3.64. Ceph 1 .88 long, 1.32 wide, 0.88 high; cl/cw
1.42; sternum 0.88 long, 0.80 wide; sl/sw 1.10;
abdomen 1.76 long, 1.28 wide.

Colour: carapace yellow orange; sternum,
chelicerae, maxillae and labium yellow, distally
white. Abdomen sepia brown (Fig. 9C,D);
frontally without or with 1 pair of weak small
white dots; dorsally with 3 pairs of big white
patches on anterior part and 2 to 3 in front of spin-
nerets; lateral with chevrons; ventrally white;
mottled with brown. Legs pale to white.

Eyes: AME smallest; eye group width 0.59 of
headwidth; AME 0.09; ALE 0.11; PME 0.11;
PLE 0.11; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.30; AME-AME
0.22; PME-PME 0.26. Clypeus 0.32 high.

Male palp (Fig. 9A,B): tegulum (LTA) absent;
sperm duct inverted U-shaped; EP about as long
as wide at base (Fig. 5D).

Female (paratype). Total length 3.52. Ceph 1.92
long, 1.20 wide. 0.92 high; cl/cw 1.60; sternum
0.84 long, 0.84 wide; sl/sw 1 .00; abdomen 1 .60
long, 1 .08 wide.

Colour; as in male.

Eyes: AME smallest; eye group width 0.5 of
headwidth; AME 0.06; ALE 0.10; PME 0.08;
PLE 0.10: AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.10; ALE-PLE
0 03' eyes group AME-PME 0.28; AME-AME
o' 16; PME-PME 0.22. Clypeus 0.40 high.
Epigyne: wider than long, anteriorly wider, with
semicircular CO (Fig. 9E, F).

Variation; in some specimens, the abdominal dots
can be expanded.

DISTRIBUTION. Queensland, Wet Tropics block
12 (Fig. 30B). Occurs only at high elevations
along the Walter Hill Range. Rainforest.

Tropasteron eacham sp. nov.

(Figs 5E, 10A-E, 30 A)

ETYMOLOGY. The type locality is near Lake Eacham; a
noun in apposition.

MATERIAL. HOLOTYPE; <3, Mt Hypipamec NP, The
Crater, NE Qld, 1 7°25’29”S, 145°29'00”E, 2-May-98, G
Milledge, pitfalls open, 25 Apr - 2 May 1998 (AM KS
77352). PARATYPES: NE Qld: 1 9 ,12c?, same data as
holotype (AM KS 55635); 3d, Lake Eacham, 750m,
1 7° 1 7 ’ S, 1 45°38’E, 9 Dec 1 989- 1 4 Jan 1 990, GM, GT, H J,

40

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 11. Tropasteron fox . A,B, palp; A, lateral; B, ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral;
scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

pitfall (QM S25961 ); 26 , Mt Father Clancy, 9km S Millaa
Millaa, 800m, 17°35 , S, 145°38’E, 6-14 Dec 1988, GM,
GT. flight intercept trap (QM S23056); l <5, Wongabe!
State Forest (NQ 26), 17°19’S, 145°29*E, 5 Nov 1991-23
Jul 1992, RR. PL, MS, pitfall (QM S24208); If, 26, as
previous, 26 Nov 1992-15 Apr 1993, R., J. & S. Raven. P.
& E. Lawless, pitfall (QM SI 9748); 4 6, Maalan SF,
1000m, rainforest, 17°35'S, 145°35’E, 20-24 Apr 1978,
VD, RR (QM S3851).

DIAGNOSIS. Similar to T. julatten and T.
luteipes with 2 frontal and 1 small frontal lateral
pair of small white dots but T. eacham as the
largest species, differs from T. julatten by long
palpal spine (LTA) and from T. luteipes by legs
with indistinct colour pattern.

DESCRIPTION. Male (holotype). Total length
4.76. Ceph 2.44 long, 1 .72 wide, 1 .08 high; cl/cw
1.42; sternum 1.08 long, 0.96 wide; sl/sw 1.12;
abdomen 2.32 long, 1.60 wide.

Colour: carapace orange brown. Sternum orange
brown; chelicerae medium brown; maxillae and
labium medium brown, distally white. Abdomen
(Fig. 10C) sepia brown; frontally with 2 pairs of
small white dots, and 1 small lateral pair; dorsally
with 3 pairs of white patches on top and 1 -2 in
front of the spinnerets; lateral with 3-4 white
patches, first circular, others elongate; ventrally
dark brown; with 1 pair of white patches near
tracheal spiracle. Legs medium brown; coxae
1-1 V white or pale; trochanter I-IV brown; femur
I-IV proximal part white, distal part pale brown,
or I-IV dark brown; tibia I distally white.

Eyes: AME smallest; eye group width 0.60 of
headwidth; AME 0.12; ALE 0.12; PME 0.16;
PLE 0.16; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.42; AME-AME
0.28; PME-PME 0.36. Clypeus 0.56 high.

Male palp (Fig. 10A.B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct
inverted U-shaped.

Female (paratype). Total length 5.76. Ceph 2.72
long, 1.84 wide, 1,28 high; cl/cw 1.48; sternum
1.12 long, 1.04 wide; sl/sw 1.08; abdomen 3.04
long, 2.40 wide.

Colour: as in males.

Eyes: AME smallest; eye group width 0.57 of
headwidth; AME 0.12; ALE 0.14; PME 0.16:
PLE 0.16; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.16; ALE-PLE
0.06; eves group AME-PME 0.42; AME-AME
0.28; PME-PME 0.36. Clypeus 0.68 high.
Epigyne: wider than long, CO inverted flask-
shaped (Fig. 10D,E).

DISTRIBUTION. Queensland, Wet Tropics blocks
8, 9 and 12. (Fig. 30A). Occurs on the Atherton
Tableland and on mountains immediately to the
west (Hugh Nelson Range) and south (Mt Father
Clancy). Rainforest.

Tropasteron fox sp. nov.

(Figs 5F, 11, 29B)

ETYMOLOGY. For the type locality, a noun in apposition.

NEW QUEENSLAND SPIDER GENUS

41

MATERIAL. HOLOTYPE: d, Seaview Ra, NE Qld, Mt
Fox Rd, 600m, rainforest, 18°50'S, 145°50’E, 15 Dec
1986-2 Jan 1987, GM, GT, SH, pitfall (QM S56866).
PARATYPES: NE Qld: 59,17(5, as for holotype (QM
S 1 8 1 78); 1 9 2 6 , as previous, 1 5-Dec-86, GM, GT, (QM
S25690); Id, Cardwell Gap (NQ 5), 18°3r53”S,

1 46° 1 1’ E, 26 Nov 1 992- 1 6 Apr 1 993, RR, PL, pitfall (QM
S24565); 1 9,2d, Wallaman Falls Rd, sieved litter, 500m,
rainforest, 18°35 , S, 145°5rE, 1 Jan 87, SH, pitfall (QM
S25704); 29 6,96, Wallaman Falls, via Ingham, sieved
litter, 500m, rainforest, 18°36’S, 145°48'E, 14 Dec 1986-2
Jan 1987, GM, GT, SH, pitfall (QM SI 81 76); l9,ld,as
previous (AM KS83919); Id, as previous, 1 Oct 1980,
GM, pitfall (QM S3761); 12d, Wallaman Falls Rd,
junction, 650m, rainforest, 18°39'S, 145°52'E, 5-12 Feb
1996, GM, pitfall (QM S41434); 3d, as previous (QM
S38223); Id, Cardwell Gap, open forest, 18°32'S,

1 46° 1 L E, 4- 1 2 Feb 1 998, GM, DC, pitfall (QM S4 1 923).

DIAGNOSIS. One of the smallest species, it
differs from all others by abdomen with 2 pairs of
white dots on the anterior part, second pair
forming a horizontal stripe, 3 spots in front of the
spinnerets and short EP.

DESCRIPTION. Male (holotype). Total length
3.44. Ceph 1 .76 long, l .24 wide, 0.84 high; cl/cw
1.42; sternum 0.84 long, 0.80 wide; sl/sw 1.05;
abdomen 1 .68 long, 1 .24 wide.

Colour: carapace orange brown. Sternum orange
brown; chelicerae medium brown; maxillae and
labium pale brown, distally white. Abdomen
(Fig. 1 1C, D) sepia brown; dorsally with 2 pairs
of white patches on the anterior part, second pair
forming a horizontal stripe, and 3 spots in front of
the spinnerets; lateral with 2 elongate white
patches, first one broad; ventrally pale brown.
Legs pale to white; coxae I-IV white or pale;
trochanter I-IV brown; femur MV proximal part
white, distal part dark brown; tibia I distally
white.

Eyes: AME smallest, PLE largest; eye group
width 0.57 of headwidth; AME 0.08; ALE 0.10;
PME 0.10; PLE 0.12; AME-AME 0.10;
AME-ALE 0.03; PME-PME 0.04; PME-PLE
0.06; ALE-PLE 0.02; eyes group AME-PME
0.26; AME-AME 0.18; PME-PME 0.24.
Clypeus 0.38 high.

Male palp (Fig. 1 1 A,B): tegulum (LTA) a short
spine, about as wide as long; sperm duct inverted
U-shaped; EP about as long as wide at base (Fig.
5F).

Female (paratype). Total length 3.76. Ceph 1.88
long, 1.12 wide, 0.64 high; cl/cw 1.68; sternum
0.80 long, 0.76 wide; sl/sw 1.05; abdomen 1.88
long, 1 .36 wide.

Colour: as in male.

Eyes: AME smallest, or PLE largest; eye group
width 0.56 of headwidth; AME 0.07; ALE 0.10;
PME 0.10; PLE 0.12; AME-AME 0.02;
AME-ALE 0.03; PME-PME 0.04; PME-PLE
0.08; ALE-PLE 0.04; eyes group AME-PME
0.30; AME-AME 0.16; PME-PME 0.24.

Clypeus 0.40 high.

Epigyne: wider than long, with very Hat, broadly
oval CO (Tig. 1 1 E,F).

Variation: the lateral white stripes can be
connected with the pale ventral side of the
abdomen.

DISTRIBUTION. Queensland, Wet Tropics
blocks 13 and 14 (Fig. 29B). Occurs on the
Seaview Range and the southern end of the
Cardwell Range, both adjacent to the valley of the
Herbert River. Rainforest and open forest.

Tropasteron halifax sp. nov.

(Figs 5G 12, 29 A)

ETYMOLOGY. For the type locality, a noun in apposition.

MATERIAL. HOLOTYPE: 6, Mt Halifax, NE Qld,
summit, 1050m, 19°07’S, 146°23’E, 1 Jan-20 Mar 1991,
A. Graham, pit (QM S56919). PARATYPES: NE Qld:
l 9 ,46 , as for holotype (QM S25689); 1 6 , 7km WNW of
Paluma (Site 32), 19°05'S. 146°06‘E, 16 Dee 1988-13 Jan

1989, R. Storey, G Dickinson, flight intercept trap (QM
S25749); 1 9,1 d, Paluma, Mt Spec, rainforest, 18°57’S,

1 46° 1 T E, 20 Jan- 1 0 Feb 1 983, N PWS per K. Smith, pitfall
(QM S444S); 56, as previous, 20 Dee 1982-20 Jan 1983,
K McDonald (QM S4385); 26, Paluma Dam Rd. Site 2,
720m, rainforest, 19°14’S, 146°13’E, 17Nov 1990-8, Dec

1990, GM, J. Seymour, flight intercept trap (QM S25958);
19,3d, Paluma Ra 75km NW Townsville, 18°47’S,
146°19’E, 18°1 .1982, B. & M. Baehr (QM S56920); 6 6 ,
12km WSW Paluma (Site 33), Mareeba DPI, 19°05'S,
146°06'E, 16 Dec 1988-13 Jan 1989. R. Storey, G
Dickinson, flight intercept trap (QM S25747); 12d,
Wallaman Falls, via Ingham, 500m, rainforest 18°36 S,
145°48'E. 14 Dec 1986^2 Jan 1987, GM. GT, SH. pitfall
(OM SI 2262); Id, Paluma Dam Rd. Site 5, 850m,
18°57'S, 146°09'E, l7Nov-8 Dec 1990,GM,J. Seymour,
pitfall (QM S46855); lid, Paluma Dam Rd, Site 5, 850m,
19°0rS 146°13'E, 8 Dec 1990-5 Feb 1991, GM, J.
Seymour, pitiall (QM S46850); I 9 ,12d , Paluma Dam Rd,
Site 3, 800m, 19°0TS, I46°13 , E, 08 Dec 1990-05 Feb

1991, GM, Seymour, pitfall (QM S3 1890); Id, Paluma
Dam Rd, Site 5,850m, 19°0rS, 1 46° 1 3’ E, 08 Dec 1990-5
Feb 1991, GM, J. Seymour, ilight intercept trap (QM
S4 1921): 3d, Paluma Dam Rd, Site 2, 720m, rainforest
19°14'S, 146°13’E, 17 Nov-8 Dec 1990, GM,J. Seymour,
pitfall (QM S46891); 15d, Mt Halifax, 19°07’S,
146°23’E, 1st wk Dec 90-8/1/91, A. Graham, pitfall (QM
S33735).

42

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 12. Tropasteron halifax. A,B, palp; A, lateral; B, ventral; scale
0.5mm; C,D, abdomen; C, dorsal; D, ventral; scale I mm; E,F,
epigyne; E, ventral; F, dorsal; scale 0.25mm. G, 1st right leg male.
Fe = femur; MT = metatarsus; P = patella; T = tibia; Ta = tarsus.

DIAGNOSIS. T. halifax , a small species with
very long palpal spine (LTA), differs from all
other species by abdomen without frontal spots
and with 2 pairs of big white dots on the anterior
part and 3-4 in front of the spinnerets.

DESCRIPTION. Male (holotype). Total length
3.72. Ceph 1 .92 long, 1 .28 wide, 0.80 high; cl/cw
1.50; sternum 0.84 long, 0.80 wide; sl/sw 1.05;
abdomen 1.80 long, 1.20 wide.

Colour: carapace orange brown; sternum orange
brown; chelicerae medium brown; maxillae and
labium pale brow r n, distally white. Abdomen
(Fig. 1 2C,D) sepia brown; dorsally with 2 pairs of
big white patches on the anterior part and 3-4 in
front of the spinnerets; lateral with 1-2 white
stripes, first one broad; ventrally pale brown.
Legs yellow, with darker brown lateral stripes;
coxae I-IV white; trochanter l-IV brown; femur
I-1V proximal part white, distal part dark brown;
tibia I distally white.

Eyes: AME smallest, PLE largest; eye group
width 0.62 ofheadwidth; AME 0.08; ALE 0.10;
PME 0.12; PLE 0.16; AME-AME 0.04; AME-
ALE 0.03; PME-PME 0.04; PME-PLE 0.06;

ALE-PLE 0.02; eyes group AME-PME 0.32;
AME-AME 0.20; PME-PME 0.28. Clypeus 0.40
high.

Male palp (Fig. 12A,B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct
inverted U-shaped; EP at least twice as long as
wide at base (Fig. 5G).

Female (paratype). Total length 4.00. Ceph 2.16
long, 1.28 wide, 0.92 high; cl/cw 1.68; sternum
0.84 long, 0.84 wide; sl/sw 1.00; abdomen 1.84
long, 1.32 wide.

Colour: as in male.

Eyes: AME smallest, or PLE largest; eye group
width 0.66 ofheadwidth; AME 0.08; ALE 0. 10;
PME 0.12; PLE 0.14; AME-AME 0.02;
AME-ALE 0.04; PME-PME 0.04; PME-PLE
0.08; ALE-PLE 0.03; eyes group AME-PME
0.03; AME-AME 0.18; PME-PME 0.28.
Clypeus 0.48 high.

Epievne: wider than long, with broadly oval CO
(Fig. 12E,F).

DISTRIBUTION. Queensland, Wet Tropics
blocks 14 and 16 (Fig. 29A). Occurs along the

NEW QUEENSLAND SPIDER GENUS

43

FIG. 13. Tropasteron heatherae. A,B, palp; A, lateral; B, ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D,
ventral; scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

summit ol the subcoastal ranges from Wallaman
Falls south to Paluma. Rainforest.

Tropasteron heatherae sp. nov.

(Figs 5H. 13, 3 IB)

ETYMOLOGY. For Heather Janetzki of the Queensland
Museum, a collector of the types.

MATERIAL. HOLOTYPE: <3, Stewart Ck, NE Qld, 4km
NNE Mt Spurgeon, Camp 1, 1250-1300m, 16°24’S,
145°13’E, 1 5-20 Oct 1991,GM,DC, HJ. L. Roberts, pitfall
(QM S57008). PARATYPES: NE Qld: 7(3, as for holotype
(QM S25799); 2d, Mt Spurgeon, 7km N of (camp 2),
1250m, 15°28T8”S, I45°13'18”E, 17-190ct 1991, GM.
DC, L. Roberts, pitfall (QM S57009); 1 d , Mossman Bluff
Track, 5- 10km W Mossman, Site 9, 1260m, rainforest,
16°39’S, 145 0 34'E, 17-31 Dec 1 988, GM, GT, ANZSES
Expedition, flight intercept trap (QM S 1 8 1 70); 1 9 ,1 <3 , as
previous, 1-19 Jan 1989 (QM SI 8164); 1 d, as previous,
Site 7, 1000m, 16°28’S, 145°22'E, 20 Dec 1989-15 Jan
1990, pitfall (QM S25763); Id, as previous, 1300m, 21
Dec 1 989, GM, GT (QM S25800); 1 d , Carbine Tableland,
plane crash site. 1330m, 16°24'27”S, 145°16'20”E, 27-28
Nov 1990. GM, GT, DC, R. Sheridan, HJ (QM S21478);
Id, as previous, pitfall (QM S21479); Id, as previous
(QM S26109); Id, Carbine Tableland, Mossman Bluff
camp, 1000m, 16°27 , 52' , S, 1 45°17’ 12^E, 30 Nov 1990,
GM, HJ (QM S25687): Id, Carbine Tblnd, above fem
patch. Devils Thumb, 16°23’S, 145°17'E. 26-27. Nov
1990, GM, HJ, pitfall (QM S25808); 3d, Cape
Tribulation, 4.5km W (Site 9), 760m, rainforest, 16°05’S,
145°26’E, 23 Scp-7 Oct 1982, GM, DY, GT, pitfall (QM
S4290); Id, Mt Demi, 7km SW Mossman, 1100m,
rainforest, 16°03’S, 145°19’E,29 0ct 1983, DY, GT (QM
S3804); 1 d. Devils Thumb, 10km NW Mossman, 1 150m,

rainforest, 16°23'S, I45°17’E, 9 Oct 1982, GM, DY, GT
(QM S4255); 4d, Kamak-Devils Thumb, 8- 12km NW
Mossman, Site 10, 1080m. 16°23*S, 1 45° 1 7’E, 26 Dec
1989-15 Jan 1990, ANZSES expedition, pitfall (QM
S18175); 1 d. Mt Pieter Botte, 950m, 16°04'S, 145°24’E,
21 Nov- 8 Dec 1993, GM, HJ, Roberts, pitfall (QM
S47060); 3d, Devils Thumb, 12km NW Mossman,
1000m, rainforest, 16°23’S, 145° 1TE, 26-27 Dec 1989,
ANZSES expedition, pitfall (QM SI 8388); 3d,
Kamak-Devils Thumb, 8- 12km NW Mossman. Site 5,
440m. 16°23'S, 145°I7'E, 26 Dec 1989-15 Jan 1990,
ANZSES expedition (QM S25130); 1 d, as previous (QM
S25124); 2d, as previous. Site 6, 900m, 16 0 23’S,
1 45° 1 7*E, 26 Dec 1 989- 1 5 Jan 1 990, ANZSES expedition
(QM S29967); Id, Mossman Bluff Track, 5- 10km W
Mossman, Site 7, 1000m, rainforest, 16°28'S, 145°22’E,
20 Dec 1989-15 Jan 1990. GM, GT, ANZSES Expedition
(QM S41543): 4d, as previous, 16-30 Dec 1988 (QM
S21701); 3d, as previous, Site 9, 1260m, rainforest. 20
Dec 1 989- 1 5 Jan 1 990 (QM S3395 1 ); 1 d , as previous. Site
9, 1260m, 16°25’S, 145°2’E, 17-31 Dec 1988 (QM
S40865); 2d, as previous. Site 10, 10km W Mossman,
1 300m, rainforest 16°25’S, 145°2’E, 20 Dec 1989-15 Jan
1990 (QM S34065); 2d. Windsor Tableland, site 4,
1270m. 16.14’33”S, 145.0 1*03 ”E, 27 Dec 1988-9 Jan
1989, E. Schmidt. ANZSES Expedition, FIT (QM S4939).

DIAGNOSIS. Differs from all other species by
abdomen with 1 or 2 pairs of white spots frontally
and with 4 pairs on top and 2 in front of the
spinnerets, second transverse can form 1 line.

DESCRIPTION. Male (holotype). Total length
3.84. Ceph 2.04 long, 1 .48 wide, 0.88 high; cl/cw

44

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 14. Tropasteron julatten. A,B, palp; A, lateral; B, ventral; scalc0.5mm; C,D, abdomen; C, dorsal; D, ventral;
scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

1.38; sternum 0.92 long. 0.88 wide; sl/sw 1.04;
abdomen 1.80 long, 1.20 wide.

Colour: carapace orange brown; sternum orange
brown; chelicerae medium brown; maxillae and
labium medium brown, distally white. Abdomen
(Fig. 13C,D) sepia brown; frontally with 1-2
pairs of smal l white dots; dorsally with 4 pairs of
white patches on top and 2 in front of the
spinnerets, second transverse can form 1 line;
lateral with chevrons; ventrally pale brown;
mottled with brown. Legs pale brown, medium
browm; coxae I-IV white; trochanter I-IV brown;
femur I-IV proximal part w'hite, distal part dark
brown; tibia I distally white.

Eyes: AME smallest; eye group width 0.57 of
headwidth; AME 0.10; ALE 0.14; PME 0.14;
PLE 0.14; AME-AME 0.02; AME-ALE 0.03;
PME-PME 0.02; PME-PLE 0.08; ALE-PLE
0.03; eyes group AME-PME 0.34; AME-AME
0.22; PME-PME 0.30. Clypeus 0.40 high; chilum
divided.

Male palp (Fig. 13A,B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct
semicircular; EP at least twice as long as wide at
base (Fig. 5H).

Female (paratype). Total length 5.56. Ceph 2.76
long, 1.80 wide, 1.36 high; cl/cw 1.53; sternum
1.20 long, 1.08 wide; sl/sw 1.11; abdomen 2.80
long, 1 .96 wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.52 of
headwidth; AME 0.10; ALE 0.14; PME 0.14;
PLE 0.14; AME-AME 0.04; AME-ALE 0.06;
PME-PME 0.06; PME-PLE 0.16; ALE-PLE
0.06; eyes group AME-PME 0.46; AME-AME
0.24; PME-PME 0.36. Clypeus 0.72 high.
Epigyne: wider than long, with broadly oval CO
(Fig. 13E,F).

Variation: females bigger than males; colour
pattern of the females are sometimes with more
chevrons.

DISTRIBUTION. Queensland, Wet Tropics
blocks 2 and 4 (Fig. 3 IB). Known from many
locations on the western and eastern edges of the
Carbine Tableland and from the mountains west
of Cape Tribulation. Rainforest.

Tropasteron julatten sp. nov.

(Figs 51, 14, 30B)

ETYMOLOGY. For the type locality, a noun in apposition.

MATERIAL. HOLOTYPE: d, Black Mt, NE Qld, 17km
ESE Julatten. 800- 1000m, rainforest, 16°39’S, 145°29’E,
29-30 Apr 1982, GM. M. Yeates, DC, (QM S4287).
PARATYPES: NE Qld: Id, Baldy Mt Rd. SW Atherton,
1150m, rainforest, 17°16’S, 145°25’E, 9 Dec 1988, GM,
GT, sieved litter (QM S25813); 1 d, Lake Eacham, 750m,
17°17’S, 145°38’E,9Dec 1989-14Jan 1990,GM,GT, HJ.
pitfall (QM S25960); 29,1 d, Bones Knob, 3km W,
1100m, 17°13’S, 145°25’E, 10 Dec 1995, GM, GT, DC
(QM S38213); 17m, Yungaburra, 17°16’S, 145°35’E, 19
Sep 76, R.Mascord (AM KS55636).

NEW QUEENSLAND SPIDER GENUS

45

FIG 15. Tropasteron luteipes. A,B, palp; A, lateral; B, ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral;
scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

DIAGNOSIS. Similar to T. eacham and T.
luteipes with 2 frontal and 1 small frontal lateral
pair of small white dots, but differs from T.
eacham and T. luteipes by short palpal spine
(LTA) and pale legs.

DESCRIPTION. Male (holotype). Total length
3.76. Ceph 1 .76 long, 1 .44 wide, 0.72 high; cl/cw
1.22; sternum 0.88 long, 0.84 wide; sl/sw 1.04;
abdomen 2.00 long, 1 .40 wide.

Colour: carapace yellow; sternum pale to orange
brown; cheiicerae pale brown, distally dark
brown; maxillae and labium pale brown, distally
white. Abdomen (Fig. 14C,D) sepia brown;
frontally with 2-3 pairs of white dots, and 1 small
lateral pair; dorsally with 3 pairs of white patches
on top and 1 -2 in front of the spinnerets; lateral
with 1 elongate horizontal white stripe reaching
front; ventral ly dark brown. Legs pale to white,
with darker brown lateral stripes; coxae I-IV
white; trochanter I-IV pale; femur 1-IV pale with
slightly darker distally.

Eyes: AME smallest; eye group width 0.56 of
headwidth; AME 0.07; ALE 0.10; PME 0.10;
PLE 0.10; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.30; AME-AME
0.18; PME-PME 0.26. Clypeus 0.40 high.

Male palp (Fig. 14A,B): tegulum (LTA) a short
spine, about as wide as long; EP at least twice as
long as wide at base (Fig. 51).

Female (paratype). Total length 3.76. Ceph 1.76
long, 1 .44 wide, 0.72 high; cl/cw 1 .22; sternum
0.88 long, 0.84 wide; sl/sw 1.04; abdomen 2.00
long, 1 .40 wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.56 of
headwidth; AME 0.07; ALE 0,10; PME 0.10;
PLE 0.10; AME-AME 0.04: AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.30; AME-AME
0.18; PME-PME 0.26. Clypeus 0.40 high.
Epigyne: wider than long, anteriorly wider, with
broadly oval CO (Fig. 14E,F).

Variation: white spots in front may come together
to a stripe.

DISTRIBUTION. Queensland, Wet Tropics
blocks 6, 8 and 9 (Fig. 30B). Known from several
areas at the northern end of the Atherton
Tableland and from a little further north at Black
Mountain. Rainforest.

Tropasteron luteipes sp. nov.

(Figs 5J, 15, 31 A)

ETYMOLOGY. Latin luteipes , bright yellow legs.

MATERIAL. HOLOTYPE: d, Mt Spurgeon, NE Qld,
2.5km S, 1100m, open forest, 16°28’S, 145°12'E, 13-21
Oct 1991, GM. HJ, pitfall (QM S57094). PARATYPES:
NE Qld: 1 9,3d, same data as holotype (QM S25793);
1 9,3d, as previous (trap 2). 1080m, 1 6°27 , S. 145°irE,
19 Nov 1997-8 Feb 1998, GM, DC, pitfall (QM S41933);
Id, as previous (trap 3), 1100m (QM S41931); 29,1 d,

46

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 16. T ropasteron malbon. A,B, palp; A, lateral; B, ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral;
scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

Windsor Tbld, NW, 1180m, open forest, 16°13’S,
144°59’E, 24 Nov 1997-9 Feb 1998, GM, DC, pitfall (QM
S41914).

DIAGNOSIS. Similar to T. eacham and T.
julatten with 2 frontal and 1 small frontal lateral
pair of small white dots but differs from both by
clearly annulated legs and from T. julatten by
long palpal spine (LTA).

DESCRIPTION. Male (holotype). Total length
4.48. Ceph 2.20 long, 1 .64 wide, 1.16 high; cl/cw
1.34; sternum 1.0 long, 0.92 wide; sl/sw 1.08;
abdomen 2.28 long, 1.60 wide.

Colour: carapace sepia brown; sternum sepia
brown; chelicerae medium brown; maxillae and
labium medium brown, distally white. Abdomen
(Fig. 15C,D) sepia brown; frontally with 3 pairs
of small white dots, and 1 small lateral pair;
dorsally with’ 3 pairs of white patches on top and
1-2 in front of the spinnerets; lateral with 3 white
patches, first small circular, others long; ventrally
dark brown. Legs yellow; coxae I-IV white;
trochanter I-IV brown; femur I-IV proximal part
white, distal part dark brown.

Eyes: AME smallest; eye group width 0.55 of
headwidth; AME 0.09; ALE 0.14; PME 0.14;
PLE 0.14; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.34; AME-AME
0.22; PME-PME 0.32, Clypeus 0.52 high.

Male palp (Fig. 15A,B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct

inverted U-shaped; EP at least twice as long as
wide at base (Fig. 5J).

Female (paratype). Total length 4.36. Ceph 2.28
long, 1.44 wide, 0.96 high; cl/cw 1.58; sternum
0.96 long, 0.92 wide; sl/sw 1 .04; abdomen 2.08
long, 1.28 wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.56 of
headwidth; AME 0.07; ALE 0.13; PME 0.12;
PLE 0.14; AME-AME 0.04; AME-ALE 0.05;
PME-PME 0.06; PME-PLE 0.12; ALE-PLE
0.05; eyes group AME-PME 0.36; AME-AME
0.18; PME-PME 0.30. Clypeus 0.52 high.
Epigyne: about as w ide as long, with broadly oval
CO (Fig. 15E,F).

DISTRIBUTION. Queensland, Wet Tropics
blocks 3 and 4 (Fig. 3 1 A). Known only from high
altitude eucalypt forests on Windsor Tableland
and the adjacent western edge of the Carbine
Tableland. Open forest.

Tropasteron malbon sp. nov.

(Figs 5K, 16, 30A)

ETYMOLOGY. For the type locality, a noun in apposition.

MATERIAL. HOLOTYPE: d, North Bell Peak, via
Gordonvale, NE Qld, 900m, rainforest, 17°05’S,
145°53’E, 16 Sept 1981, GM, DC (QM S57084).
PARATYPES: NE Qld: Id, as for holotype (QM S3777);

1 $,3d, North-South Bell Pk. saddle, Malbon Thompson
Ra, 17°07’S, 145°54’E, 20-21 Nov 1990, GM, GT, pitfall
(QM S57086); 2d, North Bell Peak, Malbon Thompson

NEW QUEENSLAND SPIDER GENUS

47

Ra, 17°07’S, 145°53’E, 20-22 Nov 1990, GM, GT, pitfall
(QM S57085); 4d, as previous, 20 Nov 1990 (QM
S258 10); Id, Bellenden K.er Ra, 0.5km S Cable Tower 7,
500m, rainforest, 17°16’S, 145°51’E, 17-24 Oct 1981,
Earthwatch Expedition & Qld Museum (QM S27616);

3 d , as previous, pitfall (QM S276 11); Id, as previous, 1 -7
Nov 1 98 1 (QM S276 1 2); 2 d , as previous (QM S27609).

DIAGNOSIS. With weak scutum but similar in
colour pattern and body shape to T. splendens ,
differs in colour pattern by: abdomen frontally
with 1 pair of small white spots and different
shape of palp, with a very short membranous
spine (LTA).

DESCRIPTION. Male (holotype). Total length
4. Ceph 2.20 long, 1 .60 wide, 1 high; cl/cw 1 .37;
sternum 1.08 long, 0.88 wide; sl/sw 1.22;
abdomen 1 .80 long, 1 .28 wide.

Colour: ceph, sternum orange brown; chelicerae
orange; maxillae and labium pale brown, distally
white. Abdomen (Fig. 16C,D) sepia brown;
frontally with 1 pair of small white dots; dorsally
with scutum and 2 pairs of white patches on
anterior part and with 2 in front of the spinnerets;
lateral with 1 white stripe; ventrally pale brown.
Legs pale brown; coxae I-IV white; trochanter
1-IV brown; femur I-IV proximal part white
becoming continuously dark brown distally.

Eyes: AME smallest; eye group width 0.61 of
headwidth; AME 0.10; ALE 0.12; PME 0.14;
PLE 0.14; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.02; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.36; AME-AME
0.24; PME-PME 0.30. Clypeus 0.48 high.

Male palp (Fig. 16A,B): tegulum (LTA) a short
spine, about as wide as long; sperm duct inverted
U-shaped; EP at least twice as long as wide at
base (Fig. 5K).

Female (paratype). Total length 5.08. Ceph 2.52
long, 1 .68 wide, 1.16 high; cl/cw 1 .50; sternum 1
long, 0.96 wide; sl/sw 1.04; abdomen 2.56 long,
1.84 wide.

Colour: as in male.

Eyes: AME smallest; AME 0. 1 0; ALE 0. 1 2; PME
0.14; PLE 0.14: AME-AME 0.02; AME-ALE
0.02; PME-PME 0.06; PME-PLE 0.10; ALE-
PLE 0.04; eyes group AME-PME 0.38; AME-
AME 0.24; PME-PME 0.34. Clypeus 0.56 high.

Epigyne: about as wide as long, with semicircular
CO (Fig. 16E,F).

Variation: additional to the lateral white stripe
there may be a small white dot. Some males also
have a dorsal weak abdominal scutum.

DISTRIBUTION. Queensland, Wet Tropics
blocks 10 and 1 1 (Fig. 30A). ICnown from mid-
altitude on the eastern side of the Bellenden Ker
Range and from the highest part of the nearby
Malbon Thompson Range. Rainforest.

Tropasteron montcithi sp. nov.

(Figs 5L, 17, 31 A)

ETYMOLOGY. For Dr Geoff Montcith of the Queensland
Museum, collector of the holotype.

MATERIAL. HOLOTYPE: d, Mossman Bluff Track, NE
Qld. 5- 10km W Mossman. Site 4, 800-1 100m, rainforest,
16°25’S, 145°2’E, 20 Decl989-15 Jan 1990, GM, GT,
ANZSES Expedition, pitfall (QM S56838).
PARATYPES: NE Qld: 5 d , as for holotype (QM S 1 8 1 6 1 );
2d, as previous (QM S18188); 4 d, as previous (Site 5),
760m, 16°28’S, 145°22’E, 16-30 Dec 1988. pitfall (QM
SI 6646); 1 d, as previous. Site 6 A, 810m (QM S 18 154);
2d, as previous, 600m, 16°39’S, I45°34’E, 16-30 Dec
1988 (QM SI 81 73); 2d, as previous, 600m, 1-16 Jan
1 989, flight intercept trap (QM S 1 8 1 82); 2d , as previous,
Site 5A, 650m, 20 Dec 1989-15 Jan 1990, pitfall (QM
S 181 55); Id, as previous. Site 5, 660m, 1-16 Jan 1989,
flight intercept trap (QM S 1 8 1 7 1 ); 2 d , as previous, 20 Dec
1989-15 Jan 1990, pitfall (QM SI 81 56); I9,4d, as
previous, Site 6, 660m. flight intercept trap (QM S 1 8 1 80);
Id, Mt Lewis Rd, 22km from FT way (Site 3), 1000m,
rainforest, 16°3S'S, 145°I7'E, 18 Dec 1989-13 Jan 1990,
GM, GT ANZSES Expedition, pitfall (QM S 18 181); 1 d,
as previous (QM S 18160); 1 d, Mt Lewis Rd. 16km from
H’way (Site 20), 960m, rainforest, 16°35'S. 145°17'E, 18
Dec 1989-13 Jan 1990, GM, GT ANZSES Expedition,
pitfall (QM SI 8183); Id, Mt Demi, 7km SW Mossman,
1100m, 16°30*S, 145°19E , 29 Oct 1983, DY, GT,
Berlesate (QM S56839); 29,29 d, as previous, 17 Dec
1995-25 Jan 1996, GM, GT, Ford, pitfall (QM S41359);
l d, as previous (AM KS83921).

DIAGNOSIS. Differs from all other species by
abdomen without white spots frontally and with 3
to 4 pairs of white patches on top and 2 to 3 in
front of the spinnerets, ventrally with 1 pair of
white stripes.

DESCRIPTION. Male (holotype). Total length
3.92. Ceph 2.00 long, 1 .52 wide, 0.88 high; cl/cw
1.31; sternum 0.96 long, 0.92 wide; sl/sw 1.04;
abdomen 1 .92 long, 1 .24 wide.

Colour: carapace, sternum orange brown; chelicerae
medium brown; maxillae and labium medium
brown, distally white. Abdomen (Fig. 17C,D)
sepia brown; dorsally with 3 or 4 pairs of white
patches on top and 2 or 3 in front of the
spinnerets; lateral with 2-3 white stripes;
ventrally pale brown; with 1 pair of white stripes
and with 1 pair of white patches near ; coxae I-IV

48

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 17. Tropasteron monteithi. A,B, palp; A, lateral; B, ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D,
ventral; scale 1mm; E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

white; trochanter I-IV brown; femur I-IV
proximal part white, distal part dark brown.

Eyes: AME smallest; eye group width 0.50 of
headwidth: AME 0.09; ALE 0.14; PME 0.14;
PLE 0.14; AME-AME 0.04; AME-ALE 0.04:
PME-PME 0.04; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.36; AME-AME
0.22: PME-PME 0.32. Clypeus 0.52 high; chilum
undivided.

Male palp (Fig. 17A,B): tegulum (LTA) a short
spine, about as wide as long; EP at least twice as
long as wide at base (Fig. 5L).

Female (paratype). Total length 5.04. Ceph 2.24
long, 1.52 wide, 0.84 high; cl/cw 1.47; sternum
0.92 long, 0.92 wide; sl/sw 1 ; abdomen 2.80 long,
2.08 wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.53 of
headwidth; AME 0.08; ALE 0.14; PME 0.14;
PLE 0.14; AME-AME 0.04; AME-ALE 0.06;
PME-PME 0.04; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.36; AME-AME
0.20; PME-PME 0.32. Clypeus 0.60 high.

Epigyne: wider than long, with oval CO (Fig.
17E,F).

Variation: lateral white patches on abdomen
sometimes fused with ventral white spots.

DISTRIBUTION. Queensland, Wet Tropics block
4 (Fig. 3 1 A). Restricted to high elevations on the

central and eastern Carbine Tableland.
Rainforest.

Tropasteron palmerston sp. nov.

(Figs 6A, 18, 30B)

ETYMOLOGY. For the type locality, a noun in apposition.

MATERIAL. HOLOTYPE: d, Palmerston NP, NE Qld
(NQ 11), 670m, rainforest, I7°35’S, 145°42’E, 30 Nov
1992-15 Apr 1993, RR, PL, pitfall (QM S57091).
PARATYPES: NE Qld: 1 d , as for holotype (QM S57090);
1 d, as previous, 30 Oct 1991-24 Jul 1992, RR, PL, MS
(QM S247 17); Id, as previous, 30 Oct 1 99 1 -24 Jul 1 992,
RR, PL, MS (QM S24726); Id, as previous, 30 Nov
1992-15 Apr 1993, RR, PL(QM S22947); 10d, as previous,
25 Jul-30 Nov 1992, RR, PL. MS, pitfall (QM S21932).

DIAGNOSIS. Similar to T. monteithi in
abdominal colour pattern, but differs in abdomen
with 3-4 pairs of white patches on anterior part
and 3-4 in front of the spinnerets and male palp
without spine (LTA).

DESCRIPTION. Male (holotype). Total length
3. 1 6. Ceph 1 .64 long, 1 .20 wide, 0.72 high; cl/cw
1.36; sternum 0.76 long, 0.76 wide; sl/sw 1;
abdomen 1 .52 long, 1 wide.

Colour: carapace, sternum orange brown;
chcliccrae orange; maxillae and labium pale
brown, distally white. Abdomen (Fig. 18C,D)
sepia brown; dorsally with 3-4 pairs of white
patches on anterior part and 3-4 in front of the
spinnerets; lateral with 1 white stripe; ventrally
dark brown; with 1 pair of long white patches.

NEW QUEENSLAND SPIDER GENUS

49

Tropasteron raveni sp. nov.

(Figs 6B, 19, 29B)

ETYMOLOGY. For Dr Robert Raven of the
Queensland Museum, in esteem for his
important work on Australian spiders.

MATERIAL. HOLOTYPE: 8, Eungella NP,
ME Qld. Finch Hatton Gorge (NQ 37),
rainforest 2I°04'E, 148°38’S, 3 Dec 1992-23
Apr 1993, RR, PL. pitfall (QM S56855).
PARATYPES: ME Qld: I9,17d, as for
holotype (QM S24632); Id, as for holotype
(AM KS83922); 1 d, as previous, RR, PL, MS
(QM S24769); l9,9d, as previous (QM
S24780); Id, as previous, 9 Nov 1991-28 Jul
1992 (QM SI 9896); 29, Finch Hatton NP,
21°09’S, 148°38’E, 14-Feb-86, RR. JG (QM
S9936); 1 9,1 d, Eungella area, eucalypt forest,
21°0r4PS, 148°3r4r*E, 20 Apr 1998, RR,
JG pitfall (QM S55633); 1 9 ,4d , Upper Hall Ck
via Carmila, 21°52’S, 149°18’E, 4 Dec 1996-6
Apr 1997, GM, E. Mulder, pitfall (QM S40602);
19, 2d, Mt Hayward, 20°20'S, 148°45’E, 20
Nov 1992-mid Apr 1993, GM, DC, pitfall (QM
S41052).

DIAGNOSIS. Differs from most other
species by short EP. Abdomen similar to
T. Cleveland with no white spots
frontally, but differs by femur I-IV
proximal margin brown, medium part
white, distal part dark brown.

FIG. 18. Tropasteron palmerston . A,B, palp; A, lateral; B,
ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral; scale
1mm.

Legs pale brown; coxae I-IV white; trochanter
I-IV brown; femur I-IV proximal part white
becoming continuously dark brown distally; tibia
I distally white.

Eyes: AME smallest, PLE largest; eye group
width 0.62 of headwidth; AME 0.08; ALE 0.10;
PME 0.10; PLE 0.12; AME-AME 0.02;
AME-ALE 0.04; PME-PME 0.04; PME-PLE
0.08; ALE-PLE 0.04; eyes group AME-PME
0.28; AME-AME 0.18; PME-PME 0.24.
Clypeus 0.30 high.

DESCRIPTION. Male (holotype). Total
length 3.8. Ceph 2.00 long, 1.40 wide,
0.88 high; cl/cw 1.43; sternum 0.92
long, 0.88 wide; sl/sw 1.05; abdomen
1.80 long, 1.36 wide.

Colour: carapace, sternum orange brown;
chelicerae orange; maxillae and labium medium
brown, distally^ white. Abdomen (Fig. 19C,D)
sepia brown; dorsally with 2 pairs of white
patches on top and 2-3 in front ol the spinnerets,
lateral with 1 white stripe; ventrally pale brown.
Legs yellow, with darker brown lateral stripes,
coxae 1-IV white; trochanter I-IV brown; femur
1-1 V proximal margin brown, medium part white,
distal part dark brown; tibia 1 distally white.

Male palp (Fig. 18A,B): tegulum (LTA) absent;
sperm duct semicircular; EP at least 2 times as
long as wide at base (Fig. 6A).

Female unknown.

DISTRIBUTION. Queensland Wet Tropics
block 12 (Fig. 30B). Known only from the
Palmerston River valley at the southern edge of
the Atherton Tableland. Rainforest.

lyes: AME smallest, PLE largest; eye group
vidth 0.60 of headwidth; AME 0.10; ALE 0.13;
)ME 0.13; PLE 0.14; AME-AME 0.02;
\ME-ALE 0.02: PME-PME 0.04; PME-PLE
).08; ALE-PLE 0.01; eyes group AME-PME
i T.n- amf-AME 0.22: PME-PME 0.30.

Clypeus 0.44 high.

Male palp (Fig. 19A,B): tegulum (LTA) a short
spine, about as wide as long; sperm duct

50

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 19. Tropasteron raveni . A,B, palp; A, lateral; B,
ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D,
ventral; scale I mm; E,F, epigyne; E, ventral; F,
dorsal; scale 0.25mm; G, 1st right leg male; scale
1mm.

semicircular; EP about as long as wide at base
(Fig. 6B).

Female (paratype). Total length 3.76. Ceph 1.96
long, 1.32 wide, 0.84 high; cl/cw 1.48; sternum
0.84 long, 0.84 wide; sl/sw 1 ; abdomen 1 .80 long,
1.28 wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.56 of
headwidth; AME 0.08; ALE 0.12; PME 0.12;
PLE 0.12; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.10; ALE-PLE
0.04; eyes group AME-PME 0.32; AME-AME
0.20; PME-PME 0.30. Clypeus 0.44 high.
Epigyne: about as long as wide, with semicircular
CO (Fig. 19E,F).

DISTRIBUTION. Widespread in the central
Queensland rainforest system from near
Proserpine south through the Eungella Range to
the mountains west of Carmila. (Fig. 29B).
Rainforest and eucalypt forest.

Tropasteron splendens sp. nov.

(Figs 6C, 20, 3 IB)

ETYMOLOGY. Latin splendens , shining; surface of
abdomen and carapace is iridescent.

MATERIAL. HOLOTYPE: d, Big Tableland, NE Qld,
700m, 15°43’S, 145°17’E, 20-21 Dec 1990, ANZSES
expedition, pitfall (QM S57095). PARATYPES: NE Qld:
10d, as for holotype (QM S25681); 4 d. Big Tableland,
740m, I5°43'S. 145°17’E. 20 Dec 1990-8 Jan 1991,
ANZSES expedition, flight intercept trap (QM S25688);
2d. Lambs Head, 10km W Edmonton, 1200m. rainforest,
17°02 , S. 145°38'E, 13 Dec 1988, GM. GT, Moss (QM
S25684); 3 <3, Mt Finnigan summit, via Helenvale,
850-950m, 15°49’S. l45°17'E,3-5 Dec 1990, DC, GT, L.
Roberts, pitfall (QM S25686); 1 9,3d, Mt Finnigan
summit, via Helenvale, 15°49’S, 145°17'E,3-5 Dec 1990,
pitfall (QM S25693); Davies Creek Road, via Mareeba
750m. I7°02’33 M S, 145°36'5rE, 4-13 Dec 1988,
GM.GT, pitfall (QM S49365).

DIAGNOSIS. Similar in colour pattern and body
shape to T. malbon , differs in colour pattern by:
abdomen frontally with 2 pair of small white
spots; and different shape of palp, with a very
long spine (LTA).

DESCRIPTION. Male (holotype). Total length
4.32. Ceph 2.24 long, 1 .68 wide, 0.88 high; cl/cw
1.33; sternum 1.08 long, 1 wide; sl/sw 1.08;
abdomen 2.08 long, 1 .32 wide.

Colour: carapace, sternum orange brown;
chelicerae medium brown; maxillae and labium

NEW QUEENSLAND SPIDER GENUS

51

FIG. 20. Tropasteron splendens. A,B, palp; A, lateral; EL ventral;
scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral; scale 1mm.

medium brown, distal ly white. Abdomen (Fig.
20C,D) sepia brown; frontally with 2 pairs of
small white dots; dorsal ly with or without weak
scutum and 2 pairs of white patches on anterior
part and with 2 in front of the spinnerets; laterally
with 1-2 white stripe a first small, second longer;
ventral ly dark brown. Legs medium brown;
coxae I-IV white; trochanter I-IV brown; femur
I-IV proximal part white becoming continuously
dark brown distally; tibia 1 distally white.

Eyes: AME smallest, PLE largest; eye group
width 0.49 of hcadwidth; AME 0.08; ALE 0.13;
PME 0.13; PLE 0.14; AME-AME 0.02;
AME-ALE 0.04; PME-PME 0.04; PME-PLE
0.10; ALE-PLE 0.04; eyes group AME-PME
0.32; AME-AME 0.18; PME-PME 0.30.
Clypeus 0.56 high.

Male palp (Fig. 20A,B): tegulum (LTA)
a long spine, at least twice as long as
wide; sperm duct inverted U-shaped;
EP at least twice as long as wide at base
(Fig. 6C).

Female unknown.

DISTRIBUTION. Queensland, Wet
Tropics blocks 1 and 7 (Fig. 3 IB).
Distribution shows a disjunction of
150km between the mountains at the
northern end of the Wet Tropics (Mt
Finnigan and Big Tableland) and the
Lamb Range, west of Cairns.
Rainforest.

Tropasteron thompsoni sp. nov.
(Figs 6D, 21 A-F, 30B)

ETYMOLOGY. For Geoff Thompson of the
Queensland Museum, a collector of the types.

MATERIAL. HOLOTYPE: <3, Lambs Head,
NE Qld, 10km W Edmonton, 1200m,
rainforest, I7°2’S, 145°39’E, 10 Dec 1989-8
Jan 1990, GM, GT, HJ, pitfall (QM S56850).
PARATYPES: NE Qld: 19,2<3, as for
holotype (QM S25962); 1 <3 1 Mt Edith, Lamb
Ra, 1140m, rainforest, 17°I0*S, 145°37'E, II
Oct 1 982, GM, DY, GT (QM S3780); 1 9 , 1 2 <3 ,
Mt Formatinc South, 700m, 16, 433’S,
145°37’E, 23-24 Nov 1990, GM, GT (QM
S2569 1 ); 1 Y. as previous (AM KS83923); 5 c 3 ,
Davies Ck Rd, 20km ESE Mareeba, 750m,
17°02’33”S, l45°36'5rE, 4-13 Dec 1988,
GM, GT, pitfall (QM S3 1 586); 1 c 3, Isley Hills,
1050m, 17°03’S, 145°42'E, 30 Nov 93, GM,
HJ (QM S25959); lc3, Mareeba, 22km SE,
900m, rainforest, 17°05’S, 145°36’E, 4 Nov
1983, DY, GT, sieved litter (QM S3802); 1 <3,
Atherton Tableland, Tinaroo Falls Dam,
17°1 1 ’S, 145°34’E. 30 Dec 81, B. & M. Baehr,
pitfall (QM S57088); 1 6 , Mt Hypipamee NP, The Crater,
15km S Atherton, 17°25 , 29”S, 145°29W’E, 29 Dec 81,
B.&M. Baehr (QM S57087).

DIAGNOSIS. Differs from all other species by
abdomen ventrally with 1 pair of white stripes
and 1 white stripe from epigastric fold to near
tracheal spiracle. Male palp with long strong
spine (LTA).

DESCRIPTION. Male (holotype). Total length
4.08. Ceph 2.08 long, 1.56 wide, 1 high; cl/cw
1.33; sternum 1 long, 0.92 wide; sl/sw 1.08;
abdomen 2.00 long, 1.48 wide.

Colour: carapace, sternum sepia brown; chelicerae
medium brown; maxillae and labium medium
brown, distally white. Abdomen (Fig. 21C,D)

52

MEMOIRS OF THE QUEENSLAND MUSEUM

sepia brown, iridescent; frontally with 1
pair of small white dots, and 1 small
lateral pair; dorsally with 2 pairs of
white patches on top and 2-3 in front of
the spinnerets; laterally with 3-4 white
stripes; ventrally dark brown; with 1
pair of white stripes and 1 white stripe
from epigastric fold to near tracheal
spiracle. Legs pale brown; coxae I-IV
white; trochanter I-IV brown; femur
I-IV proximal part white, distal part
dark brown; tibia I distally white.

Eyes: AME smallest, PLE largest; eye
group width 0.57 of headwidth; AME
0.08; ALE 0.14; PME 0.14; PLE 0.16;
AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.04; PME-PLE 0.10;
ALE-PLE 0.04; eyes group AME-PME
0.34; AME-AME 0^.20; PME-PME
0.32. Clypeus 0.44 high.

Male palp (Fig. 21 A,B): tegulum (LTA)
a short spine, as wide as long; sperm
duct inverted U-shaped; EP at least
twice as long as wide at base (Fig. 6D).
Female (paratype). Total length 4.48.
Ceph 2.08 long, 1.48 wide, 0.96 high;
cl/cw 1 .40; sternum 0.92 long, 0.92 wide;
sl/sw 1 ; abdomen 2.40 long, 1 .76 wide.
Colour: as in male.

Eyes: AME smallest, or PLE largest;
eye group width 0.55 of headwidth;
AME 0.08; ALE 0.12; PME 0.12; PLE
0.14; AME-AME 0.04; AME-ALE
0.04; PME-PME 0.04; PME-PLE 0.10;
ALE-PLE 0.04; eyes group AME-PME
0.34; AME-AME 0.20; PME-PME
0.28. Clypeus 0.48 high.

Epigyne: wider than long, with
semicircular CO (Fig 21E,F).

Variation: ventral white patches on
abdomen may be connected.

FIG 21 . Tropasteron thompsoni. A,B, palp; A, lateral; B, ventral;
scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral; scale 1mm;
E,F, epigyne; E, ventral; F, dorsal; scale 0.25mm.

DISTRIBUTION. Queensland, Wet
Tropics blocks 6, 7, 8 and 9 (Fig. 30B). Ranges
from Mt Formartine south through the Lamb
Range to the northern parts of the Atherton
Tableland and Hugh Nelson Range. Rainforest.

Tropasteron tribulation sp. nov.

(Figs 6E, 22, 31 A)

ETYMOLOGY. For the type locality, a noun in apposition.

MATERIAL. HOLOTYPE: d , Cape Tribulation, NE Qld,
3.5km W (Site 7), 680m, rainforest, 16°05’S, 145°27'E,
2-7 Oct 1 982, GM, DY, GT, pyrethrum knockdown (QM

S57096). PARATYPES: NE Qld: Id, as holotype, 2-7 Oct
1982, GM, DY, GT, pyrethrum knockdown (QM S4288);
I d, as previous, 23 Sep-7 Oct 1982, pitfall (QM S4283);
1 d, as previous, 2.5km W (Site 5), 180m, 23 Scp-7 Oct
1 982, pitfall (QM S4256); 1 9 , as previous, 2.7km W (Site
5 A) 400m, Jan 83, GM, sieved litter (QM S4269); 1 $ , as
previous, 1.5km W (Site 3), 150m, 5-9 Jan 1983, baited
pitfall (QM S4270); 1 9, as previous, 2.7km W (Site 5A),
400m. 5-9 Jan 1983, baited pitfall (QM S3765); Id, as
previous, 2km W (Site 4), 200m, 23 Sep- 7 Oct 1 982, GM,
DY, GT, pitfall (QM S4289); Id, as previous, transect, site
6 500m, rainforest, 17 Nov 98, GM, pyrethrum knock-
down trees (QM S47503); Id, Fritz Ck, Bloomfield,

NEW QUEENSLAND SPIDER GENUS

53

FIG. 22. Tropasteron tribulation. A,B, palp; A, lateral; B,
ventral; scale 0.5mm; C, abdomen dorsal; scale 1 mm; D,E,
epigyne; D, ventral; E, dorsal; scale 0.25mm.

rainforest, 15°52'S, 145°21’E, 31 Dec 75, M. Gray (AM
K.S0490); 39,1 <3, Shiptons Flat, 15°47'33”S,
145°13’5i”E, 16-21 Nov 1975, RR, VD, pitfall (QM
S4467); l?,lc3, as previous (AM KS83924); Id, Mt
Misery road. Site 1, 730m, 15°53’S, 145°LVE, 6 Dec
1990-17 Jan 1991, ANZSES expedition, pitfall (QM
S57089); 1 <3, West Claudie R, Iron Ra, 50m, rainforest,
12°45’S, 143°14‘E, 3-10Dec 1985,GM,DC(QMS40844).

DIAGNOSIS. Differs from all other species by
abdomen with 3 pairs of frontal and without
fronto- lateral white spots. Male palp with long
spine (LTA).

DESCRIPTION. Male (holotype 2nd measure-
ment). Total length 3.4-3.76. C’eph 1 .8- 1 .92 long,

1 .32- 1 .36 wide, 0.72-0.76 high; cl/cw 1 .36- 1 .41;
sternum 0.84-0.92 long, 0.84 wide; sl/sw 1-1 .09;
abdomen 1.6-1.84 long, 1-1.20 wide.

Colour: carapace orange brown. Sternum pale
brown, orange brown; chelicerae pale brown,
distally dark brown; maxillae and labium pale
brown, distally white. Abdomen (Fig. 22C) sepia
brown; frontally with 3 pairs of white dots;
dorsally with 3 pairs of white patches on top and
1-2 in front of the spinnerets; laterally with 1-2
white patches first small, second elongate;
ventrally pale to dark brown; mottled with white,
with 1 pair of white patches near tracheal
spiracle. Legs pale brown; coxae I-1V white;
trochanter I-IV pale to brown; femur I-IV
proximal part white becoming continuously dark
brown distally.

Eyes: AME smallest; eye group width 0.53-0.64
of headwidth; AME 0.08-0.09; ALE 0.10; PME

0.1-0.13; PLE 0.11-0.13; AME-AME 0.02-0.04;
AME-ALE 0.04; PME-PME 0.04-0.06; PME-
PLE 0.08; ALE-PLE 0.04; eyes group AME-
PME 0.03-0.32; AME-AME 0.18-0.22;
PME-PME 0.26-0.30. Clypeus 0.4-0.44 high.

Male palp (Fig. 22A,B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct
semicircular.

Female (paratype). Total length 3.88-4.08. Ceph
1.88-2.08 long, 1.28-1.36 wide, 0.88-0.96 high;
cl/cw 1.46-1.53; sternum 0.84-0.92 long,
0.8-0.92 wide; sl/sw 1-1.05; abdomen 2.00 long,
1.32-1.40 wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.53-0.54
of headwidth; AME 0.06-7; ALE 0. 1 -0. 1 2; PME
0.1-0.11; PLE 0. 11-0.1 4; AME-AME 0.04; AME-
ALE 0.04; PME-PME 0.06; PME-PLE 0.10;
ALE-PLE 0.04; eyes group AME-PME 0.32;
AME-AME 0.16-0.18; PME-PME 0.26-0.28.
Clypeus 0.52 high.

Epigyne: wider than long, with broadly oval CO
(Fig 22E, F).

DISTRIBUTION. This is the most northerly
species of Tropasteron (Fig. 3 1 A). It occurs from
Iron Range in Cape York Peninsula south to the
Bloomfield River (Block 1 ) and Cape Tribulation
(Block 2) at the northern end of the Wet Tropics.
It is mostly in lowlands but is found up to 680m,
west of Cape Tribulation. Rainforest.

54

MEMOIRS OF THE QUEENSLAND MUSEUM

Tropasteron yeatesi sp. nov.
(Figs 6F, 23, 30A)

ETYMOLOGY. For Dr David Yeates of CSIRO
Entomology, a collector of the types.

MATERIAL. HOLOTYPE: <3, Bellenden Ker
Ra, NE Qld. Summit, 1560m, rainforest,

17°16’S, 145°52’E, 28 Aug -8 Oct 1991, GM,

HJ, pitfall (QM S56995). PARATYPES: NE
Qld: 19,1 <3, same data as holotype (QM
S25718); 19,1 <5, Bellenden Ker Ra, Summit
TV Stn, 1560m, 17 0 16'S, 145°5PE,29 Sep 81,

GM, DC (QM S3732); 29,1 (3, as previous, 29
Apr-2 May 1983, GM, DY (QM S3757);

1 V ,1 (3, as previous, 28 Oct 1983, GM, DY, GT
(QM S4267); 1 6 , Massey Ra 4km W, center of
Bellenden Ker, 1250m, 17°16’S, I45°49’E,9-1 1
Oct 1991, GM. HJ, DC (QM S25783); 49,33,

Mt Bartle-Frcrc, summit creek, 1500m,
rainforest, 17°23'S, 145°48'S, 24 Sep 81, GM,

DC, pitfall (QM S3722); 2 9 , as previous, 0.5km
N of Sth Peak, 17°24’S, 145°49’E, 6-8 Nov
1981, Earthwatch Expedition (QM S47543);

29,2(3, as previous, Sth Peak (QM S47543);

19, as previous, Sth Peak, 1400- 1500m,
rainforest , 17°23’S, I45°49'E, 7-8 Oct 1981.
Earthwatch Expedition, berlesate (QM S47543);

19,1<3, Massey Ra, 12km S Gordonvale,

1300m. rainforest. 17°16'S, 145°49’E, 2 May
1 983, GM, DC, sieved litter (QM S47543); 1 73 ,

Bellenden Ker Ra, Cable Tower 3, 1054m,

17°16’S, !45°5rE. 25-31 Oct 1981, Earthwatch
& Qld Museum, pitfall (QM S27890); 1(3, as
previous, sieved litter (QM S27888); 1(3, as
previous, 1-7 Nov 1981. pitfall (QM S27881);

2 c3 , as previous, 17-24 Oct 1981 (QM S27886);

89,4(3, as previous. Summit TV Stn, 1560m,

PILI (QM S27596): 39.43, as previous, 1-7
Nov 1981 (QM S27589); 19,13. as previous
(AM KS83925); 39,7 c3, as previous, 17-24 Oct
1981, PILI (QM S2759I ); lc3, as previous,

0.5km S of Cable Tower 7, 500m, rainforest (QM
S276 1 3 ); 1 2 <3 , as previous, pitfall (QM S276 1 0);

19,1(3, as previous. Centre Peak Summit,

1 500m, rainforest, 1 1 Apr 79, GM, sieved litter
(QM S27595); 1 9, Bellenden Ker Ra, Summit
TV Stn, 1560m, rainforest, 29 Apr-2 May 1983,

GM, DY (QM S27593); 1<3, as previous (QM
S38519); 2<3, as previous (QM S38538);

19,1(3, as previous (QM S38547), 2 <3, as
previous, 17-Apr-97, GM, E. Russell, sieved

litter (QM S3 191 8); 29,1 c3, as previous. Centre pjQ 23
Peak summit, rainforest, 28 Aug 91, GM, HJ, sca ic 0
sieved litter (QM S3 1 565); 1 c3 , as previous. Top j orsa i

Stn, rainforest. 17°16'S, 145°52’E, 17-18 Apr
1997, GM, J. Ovendere pitfall (QM S40691 ).

DIAGNOSIS. Similar to T. cooki but differs from
all other species by abdomen without white spots
frontally and with 3 to 4 pairs of white patches on

Tropasteron yeatesi. A,B. palp; A, lateral; B, ventral;
5mm; C-F, abdomen variation of colour pattern; C,E,
D,F, ventral; scale 1mm; GH, epigyne; G, ventral; H,

top and 3 in front of the spinnerets can be
enlarged to white chevrons, ventrally with white
chevrons. Male palp with long spine (LTA), at
least twice as long as wide.

NEW QUEENSLAND SPIDER GENUS

55

FIG. 24. Tropasteron andreae. A,B, palp; A, lateral; B, ventral:
scale 0.5mm; C, abdomen dorsal; scale 1mm; D,E epigyne; D,
ventral; E, dorsal; scale 0.25mm.

DESCRIPTION. Male (holotype). Total length
3.6-4.16. Ceph 2.04-2.24 long, 1.4-1.56 wide,
0.88-0.96 high; cl/cw 1.43-1.45; sternum
0.8-0.96 long, 0.8-0.92 wide; sl/sw 1-1.04;
abdomen 1.56-1.92 long, 1.16-1.32 wide.

Colour: carapace, sternum orange brown;
chelicerae yellow to medium brown; maxillae
and labium pale brown, distal ly white. Abdomen
(Fig. 23C-F) sepia brown; dorsally with 4 pairs of
white patches on top and 3 in front of the
spinnerets, can be enlarged to white chevrons;
laterally with 1 big white patch and pale
booklungs; ventrally dark brown; mottled with
white. Legs pale brown; coxae 1-IV white;
trochanter I- IV brown; femur I-IV proximal part
white, distal part dark brown; tibia I-IV
proximally and distally with brown spot.

Eyes: AME smallest, PLE largest; eye group
width 0.54 ofheadwidth: AME 0.08; ALE 0.12;
PME 0. 1 1 -0. 1 3; PLE 0. 1 2-0. 1 4; AME- AME 0.04;
AME-ALE 0.04; PME-PME 0.04; PME-PLE
0.08-0.10; ALE-PLE 0.04; eyes group AME-
PME 0.3-0.34; AME-AME 0.20; PME-PME
0.26-0.30. Clypeus 0.44 high; chilum divided.

Male palp (Fig. 23A,B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct
inverted U-shaped; EP at least twice as long as
wide at base (Fig. 6F).

Female (paratype). Total length 3.96-4.16. Ceph
1.96-2.20 long, 1.36-1.52 wide, 1-1.08 high;
cl/cw 1.44-1.44; sternum 0.84-0.92 long,
0.84-0.92 wide; sl/sw 1-1; abdomen 1.96-2 long,
1.44-1.48 wide.

Colour: as in male.

Eyes: AME smallest, or PLE largest; eye group
width 0.5-0.54 of headwidth; AME 0.07-0.08;
ALE 0.1-0.12; PME 0.11-0.12; PLE 0.11-0.14;
AME-AME 0.04; AME-ALE 0.04; PME-PME
0.04; PME-PLE 0.1-0.14; ALE-PLE 0.04; eyes
group AME-PME 0.32-0.34; AME-AME
0.18-0.20; PME-PME 0.26-0.28. Clypeus
0.44-0.48 high.

Epigyne: wider than long, anteriorly wider, with
flattened broadly oval CO (Fig. 23G,H).

Variation: The abdominal white patches can fuse
to white chevrons.

DISTRIBUTION. Queensland, Wet Tropics
block 10 (Fig. 30A). Known only from the
contiguous massifs of Bellenden Ker, Bartle
Frere and the Massey Range. Almost all
specimens are from above 1000m and it is
common at the maximum altitudes of
1 500- 1600m. Rainforest.

KEY TO SPECIES OF T. ANDREAE-GROV?

1 . Palp dorsal tibial apophysis EP without basal bump.

Abdomen with 1 elongate lateral white stripe not
reaching front, abdomen, epigyne ( F igs 6G, ^.4A-b)

& T. andreae

Palp dorsal tibial apophysis EP with basal bump.
Abdomen with 1 elongate lateral white stripe reaching
front (Fig. 6H-J)

2. Abdomen dorsally with 1 pair of long stripes (Figs 25C,

26C) 3

Abdomen dorsally with 2 pairs of long stripes, abdomen,
palp (Figs 6J, 27 A-C) Trobertsi

3. Abdomen additional with 2 pairs of dorsal horizontal

stripes, abdomen, palp (Figs 6H, 25 A-D) . . T. magnum

Abdomen additional with 3 pairs of dorsal horizontal
stripes, abdomen, palp (Figs 61, 26A-D)

I pseudomagnum

56

MEMOIRS OF THE QUEENSLAND MUSEUM

Tropasteron andreae sp. nov.
(Figs 6G, 24, 32)

ETYMOLOGY For my friend and personal PR
consultant Andrea Hammond.

MATERIAL. HOLOTYPE: 6 , Finch Hatton
NP, ME Qld, 350m. rainforest, on foliage,
21°09'S, 148°38'E, 7-14 Apr 1975, VD, R.
Kohout (QM S57985). PARATYPES: ME Qld:
1 9 , same data as holotype (QM S 1 6742); 1 9 , as
previous, 14-Feb-86, RR, JG (QM S9936).

DIAGNOSIS. The largest species of the
genus, differs from all other species in T
andreae-group in abdominal pattern and
male palp with long strong spine (LTA).

FIG 25.
scale 0.

DESCRIPTION. Male (holotype). Total
length 5.6. Ceph 2.80 long, 2.08 wide,

1 .04 high; cl/cw 1 .34; sternum 1 .20 long,

1.16 wide; sl/sw 1.03; abdomen 2.80
long, 1 .80 wide.

Colour: carapace orange brown,
iridescent. Sternum orange brown;
chelicerae medium brown; maxillae and
labium pale brown, distal ly white.
Abdomen (Fig. 24A, B) sepia brown,
iridescent; frontally with 1 pair of small
white dots; dorsally with 3-4 pairs of
white patches, 1, 2 elongated stripes, 3
sometimes 4 horizontal
half-moon-shaped and 1 elongate in
front of the spinnerets; lateral with 1
white stripe; ventrally dark brown,
iridescent. Legs medium brown, with
darker brown lateral stripes; coxae I-IV
white; trochanter I-IV brown; femur I-IV
proximal part white becoming continuously dark
brown distal ly; tibia I-IV proximally pale,
distally brown.

Eyes: in 2 rows with 4 eyes, both rows procurved;
AME smallest; eye group width 0.59 of
headwidth; AME 0.10; ALE 0.12; PME 0.14;
PLE 0.14; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.08; PME-PLE 0.08; ALE-PLE
0.04; eyes group AME-PME 0.38; AME-AME
0.24; PME-PME 0.36. Clypeus 0.68 high.

Male palp (Fig. 24A, B): cymbium flange short
straight, with distal hook, only about 10% of
cymbium long, and with small rounded
extension, incision in between; tegulum (LTA) a
long spine, at least twice as long as wide; DTA
membranous, semicircular, distal part folded
containing embolus; prolateral extension (PE)
short not longer then base of DTA; retrolaterally
with sharp tip and without any additional

Tropasteron magnum. A,B, palp; A, lateral; B, ventral;
5mm; C,D, abdomen; C, dorsal; D, ventral; scale 1 mm.

extension; sperm duct inverted U-shaped;
embolus thin, semicircular, embolus base
cylindrical; tibia short; ventro-lateral tibial
apophysis with chitinous rim dorsally; EP
distally flattened (Fig. 6G). Femur with 2 strong
spines dorsally.

Female (paratype). Total length 7.04. Ceph 3.44
long, 2.32 wide, 1.28 high; cl/cw 1.48; sternum
1.44 long, 1.40 wide; sl/sw 1.03; abdomen 3.60
long, 2.80 wide.

Colour: as in male.

Eyes: AME smallest; eye group width 0.56 of
headwidth; AME 0.14; ALE 0.16; PME 0.18;
PLE 0.20; AME-AME 0.02; AME-ALE 0.06;
PME-PME 0.08; PME-PLE 0.18; ALE-PLE
0.06; eyes group AME-PME 0.48; AME-AME
0.30; PME-PME 0.44. Clypeus 0.88 high.

NEW QUEENSLAND SPIDER GENUS

57

FIG. 26. Tropasteron pseudomagnum. A,B, palp; A, lateral; B,
ventral; scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral;
scale 1mm.

MATERIAL. HOLOTYPE: <J, Cape
Tribulation, NE Qld, 3km W (Site 6), 500m,
rainforest, 16°05’S, 145°27’E,5-9Jan 1983, GM,
baited pitfall (QM S4264).

DIAGNOSIS. Differs from T. pseudo-
magnum and T. robertsi in abdomen with
3 pairs of white patches, first elongated,
2, 3 horizontal and 2 in front of the
spinnerets.

DESCRIPTION. Male (holotype). Total
length 4.6. Ceph 2.40 long, 1.96 wide,
0.88 high; cl/cw 1 .22; sternum 1 .20 long,

1 wide; sl/sw 1 .20; abdomen 2.20 long,
1.56 wide.

Colour: carapace pale brown. Sternum
orange brown; chelicerae pale brown,
distally dark brown; maxillae and labium
pale brown, distally white. Abdomen
(Fig. 25C,D) sepia brown; frontally with
1 pair of small white dots; dorsally with 3
pairs of white stripes, first long, 2, 3
horizontal and 2 in front of the spinnerets;
lateral with 1 white stripe reaching front;
ventrally dark brown; with 1 pair of
elongate white patches. Legs pale brown,
with darker brown lateral stripes; coxae
I-IV white; trochanter I-IV white; femur
1-1V pale with brown ring distally.

Eyes: AME smallest; eye group width
0.46 of headwidth; AME 0.07; ALE 0. 1 2;
PME 0.12; PLE 0.12; AME-AME 0.05;
AME-ALE 0.06; PME-PME 0.08;
PME-PLE 0.14; ALE-PLE 0.06; eyes
group AME-PME 0.36; AME-AME
0.19; PME-PME 0.32. Clypeus 0.64
high.

Epigyne: much wider than long, with broadly
lyriform CO (Fig. 24D,E). Female palpal claw
strong with more then 6 teeth.

Male palp (Fig. 25 A, B): tegulum (LTA) a
long spine, at least twice as long as wide;
sperm duct inverted, U-shaped; EP distally
flattened (Fig. 6H).

Variation: in one specimen the 4th horizontal
patch on abdomen is reduced.

DISTRIBUTION. Known only from Finch Gatton
Gorge within Eungella National Park (Fig. 32).
Rainforest.

DISTRIBUTION. Queensland, Wet Tropics
block 2 (Fig.32). Known only from a single
collection at mid-altitude on the mountain slopes
west of Cape Tribulation. Rainforest.

Tropasteron pseudomagnum sp. nov.

(Figs 6H, 26, 32)

Tropasteron magnum sp. nov.
(Figs 6H, 25A-D, 32)

ETYMOLOGY. Latin magnum , large. T. magnum is one
of the largest spiders in this genus.

ETYMOLOGY. For its similarity to T. magnum.

MATERIAL. HOLOTYPE: <$, Mt Spurgeon, NE Qld,
7km N of (camp 2), 1250m, 15°28’18”S, 145°13’18 M E,
17-19 Oct 1991, GM, DC, L. Roberts, pitfall (QM
S57098). PARATYPES: NE Qld: 2d, same data as

58

MEMOIRS OF THE QUEENSLAND MUSEUM

holotype (QM S25798); 1 d , same locality, GM,
HJ (QM S25805).

DIAGNOSIS. Differs from T. magnum
and T. robertsi in abdomen with 4 pairs of
white patches, first long, 2-4 horizontal
and 1 in front of the spinnerets.

DESCRIPTION. Male (holotype). Total
length 4.6. Ceph 2.40 long, 1.84 wide, 1
high; cl/cw 1.30; sternum 1.04 long, 1
wide; sl/sw 1.04; abdomen 2.20 long,

1.52 wide.

Colour: carapace pale brown. Sternum
sepia brown; chelicerae pale brown,
distally dark brown; maxillae and labium
pale brown, distally white. Abdomen
(Fig. 26C,D) sepia brown; frontally with
1 pair of small white dots; dorsally with 4
pairs of white patches, first long, 2-4
horizontal and 1 in front of the
spinnerets; lateral with 1 white stripe
reaching front; ventrally dark brown;
with 1-2 pairs of elongate white patches.

Legs medium brown, with darker brown
lateral stripes; coxae I-IV white;
trochanter I-IV white; femur I-IV pale
with brown ring distally.

Eyes: AME smallest; eye group width
0.55 of headwidth; AME 0.09; ALE
0.11; PME 0. 1 2; PLE 0. 1 2; AME-AME
0.04; AME-ALE 0.04; PME-PME 0.06;
PME-PLE 0.12; ALE-PLE 0.04; eyes
group AME-PME 0.34; AME-AME
0.22; PME-PME 0.30. Clypeus 0.64
high.

Male palp (Fig. 26A,B): tegulum (LTA) a
long spine, at least twice as long as wide; sperm
duct semicircular; EP distally flattened (Fig. 6H).

FIG 27. Tropasteron robertsi. A,B, palp; A, lateral; B, ventral;
scale 0.5mm; C,D, abdomen; C, dorsal; D, ventral; scale 1 mm.

patches, 1 , 2 long stripes, third horizontal and 3 in
front of the spinnerets.

DISTRIBUTION. Queensland, Wet Tropics
block 4 (Fig. 32). Known only from high altitude
on the remote northwest sector of the Carbine
Tableland. Rainforest.

DESCRIPTION. Male (holotype). Total length
4.68. Ceph 2.36 long, 1.80 wide, 1 high; cl/cw
1.31; sternum 1.04 long, 1 wide; sl/sw 1.04;
abdomen 2.32 long, 1.48 wide.

Tropasteron robertsi sp. nov.

(Figs 6J, 27, 32)

ETYMOLOGY. For Lewis Roberts, one of the collectors
of the types.

MATERIAL. HOLOTYPE: d, Mt Finnigan, NE Qld,
1100m, rainforest, 15°49’S, 145°17’E, 28-30 Nov 1985,
GM, DC, L. Roberts (QM S3799). PARATYPES: NE Qld:
1 d, same locality, 27 Sep 1975, RR (QM S4470).

DIAGNOSIS. Differs from T. magnum and T.
pseudomagnum in abdomen with 3 pairs of white

Colour: carapace pale brown. Sternum sepia
brown; chelicerae pale brown, distally dark
brown; maxillae and labium medium brown,
distally white. Abdomen (Fig. 27C.D) sepia
brown; frontally with 1 pair of small white dots;
dorsally with 3 pairs of white patches, 1, 2 long
stripes, third horizontal and 3 in front of the
spinnerets; lateral with 1 white stripe reaching
front; ventrally dark brown. Legs medium
brown, with darker brown lateral stripes; coxae
I-IV white; trochanter I-IV white; femur I-IV
pale with brown ring distally.

NEW QUEENSLAND SPIDER GENUS

59

FIG 28. Maps of the northern Queensland Wet Tropics. A, limits of the 1 7 mountain/rainforest blocks according
to Monteith ( 1 995), modified in Bouchard (2002): 1 , Mount Finnigan; 2, Thornton Peak; 3, Windsor Tableland;
4, Carbine Tableland; 5, Hann Tableland; 6, Black Mountain; 7, Lamb Range; 8, Walsh/Hugh Nelson Ranges; 9,
Atherton Tableland; 10, Mount Bellenden Ker; 11, Malbon Thompson Range; 12, Walter Hill Range; 13,
Kirrama/Cardwell Ranges; 14, Seaview Range; 15, Hinchinbrook Island; 16, Paluma/Blue Water Ranges; 17,
Mount Elliot. In brackets, number of species occuring in the rainforest block. B, distribution range of
Tropasteron spp.

Eyes: AME smallest; eye group width 0.46 of
headwidth; AME 0.08; ALE 0.10; PME 0.12;
PLE 0.12; AME-AME 0.04; AME-ALE 0.04;
PME-PME 0.06; PME-PLE 0.10; ALE-PLE
0.06; eyes group AME-PME 0.34; AME-AME
0.20; PME-PME 0.30. Clypeus 0.60 high.

Male palp (Fig. 27 A, B): tegulum (LTA) a long
spine, at least twice as long as wide; sperm duct
semicircular; EP distally flattened (Fig. 6J).

DISTRIBUTION. Queensland, Wet Tropics
block 1 (Fig. 32). Restricted to the summit
plateau of Mt Finnigan. Rainforest.

DISCUSSION

With 22 species, Tropasteron is one of the
largest genera in the Asteron - complex and the
only one that has radiated in rainforests (Fig.
28B). Most species of Tropasteron are found only
in montane rainforests; however, 7! Cleveland , T.
fox and T. raveni are also found in adjacent open
forest, and T. luteipes have been found only in the
open forest, at Mt Spurgeon, (ca. 1100m)
surrounded by rainforest.

PHYLOGENETIC RELATIONS. All species of
Tropasteron are closely related in their uniform

60

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 29. Distribution of Tropasteron species A, T. cardwell , T. Halifax: B, T. Cleveland , T. fox, T.raveni.

palps and epigynes but distinct from all other
genera of the Asteron- complex by their
synapomorphies, mentioned in Table 1 (2, 6, 8, 9,
10, 11, 12, 13). Hence, it is difficult to find
phylogenetic relationships within the separate
species-groups, because parallel evolution
obviously took place when ancient aridity events
caused the loss of forest corridors connecting
each block (Reid & Storey, 2000). Therefore, a
complete phylogenetic analysis cannot be done.
Nevertheless, there are quite a few character
states (Table 1 ), which help to reconstruct some
phylogenetic relations. Two quite distinct
species-groups — T. cleveland-group and T.
andreae-g roup — have evolved.

The combination of the most plesiomorphic
characters:

(1) abdomen with 2 pairs of white dots on top
and 1-3 dots in front of the spinnerets (Figs 1 A,
12C, 19C), as it is found also in the outgroup
Pentasteron and some other genera of the
Asteron-compXex e.g. in Asteron , Leptasteron.

(3) male palp with a short external prong of
dorsal tibial apophysis (EP) (Figs 5A,D,F, 6B),
occur only in 2 (I Cleveland , T. raveni) of 18
species in T. cleveland-gvoxxp. All 16 other
species have longer EP or a more evolved colour
pattern with additional white dots.

T. andreae- group seems to be the more
apomorphic group, because of (7) the more
evolved cymbial flange with hook CH (Fig.
24A), (4) the dorsal tibial apophysis with EP
distally flattened (Fig. 6G-J) and (12) the
lyriform copulatory opening (CO) (Fig. 24D). T.
andreae sp. nov. (5) without any basal bump
(BB) on the external prong of the dorsal tibial
apophysis seems to be the most plesiomorphic
species in the T. andreae- group. All other more
evolved species possess this basal bump.

Final phylogenetical results cannot be discussed
here because the knowledge of relationships
from all genera in the Asteron- complex are not
yet known.

NEW QUEENSLAND SPIDER GENUS

61

FIG. 30. Distribution of Tropasteron species A, 7 eacham, T. malbon , 7 yeatesi; B , T daviesae , 7 julatten ,
7. palmerston , 71 thompsoni.

DISTRIBUTION AND BIOGEOGRAPHY

Methods used for collecting Tropasteron
specimens were similar (pitfall trapping and hand
collecting) and sampling effort has been
effectively uniform across all sites. Hence, gaps
or absences are taken here to be real and not
simply an indication of heterogeneous sampling.
Based on the examined material of Tropasteron ,
the following conclusions can be made.

All species of the 71 andreae- group are highly
endemic to single localities (Table 2). They are
allopatric on the three northern massifs and
include a southern outlier at Eungella (71
andreae). Their main centre of diversity is in the
most northern rainforest blocks: 1 Mt Finnigan
(7 robertsi ), 2 Thornton Peak (71 magnum ), 4
Carbine Tableland (71 pseudomagnum). At all
these sites, species of the T. andreae- group are
sympatric with those of the T. Cleveland- group.

The much bigger and more diverse T. cleveland-
group, with 1 8 species, occurs from Iron Range
with its centre in Wet Tropics and also a southern

outlier at Carmila (7 raveni). Of the other two
species that occur outside the WTWHA, most
material of 7. tribulation is from Cape
Tribulation and Shiptons Flat with a northern
outlier from Iron Range ( 1 6 ) and the southern 7
Cleveland to which mountain it is endemic both in
areas of rainforest and open forest at Cape
Cleveland. None of the species at the limits of the
genus distribution are sympatric with any other
species. As in other examined taxa with low
vagility (e.g. beetles: Baehr, 1995; Bouchard,
2002; Reid & Storey, 2000) the main centre of
diversity of 7 cleveland-g roup is in the northern
part of the mountain/rainforest blocks oi North
Queensland (defined by Monteith, 1995), with
2-4 species in each block: 14 (Seaview Ra.: 7
Halifax, T. fox), 13 (Cardwell Ra.: 7 fox, 7
cardwell), \2 (Walter Hill Ra.: 7 daviesae, 7
palmerston ), 11 (Malbon Thompson Ra.: 7
cardwell , 7 cooki, T. malbon), 10 (Mt Bellenden
Ker: 7 yeates , 7 malbon ), 9 (Atherton Tableland:
7. eacham , 7. julatten , 7. thompsoni ), 8

62

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 3 1 . Distribution of Tropasteron species A, I luteipes , 7 monteithi , 7 tribulation ; 7 cooki , 7! haetherae ,

7! splendens.

(Walsh/Hugh Nelson Ranges: 7! eacham , 7
julattem, T. thompsoniO), 7 (Lamb Ra.: 71 coo/:/,
7 splendens , 7! thompsoni ), 6 (Black Mt: 71
julatten , 71 thompsoni ), 4 (Carbine Tableland: 71
heather ae, 71 monteithi , 71 luteipes ), 2 (Thornton

Peak: 71 heatherae, 71 tribulation ), 1 (Mt
Finnigan: 7 heatherae , 71 splendens , 7.
tribulation ).

Margin rainforest blocks have only 1 or no
species 17 (Mt Elliot 0), 16 (Paluma Ra.: 7

TABLE 1 . Character states and their phylogenetical value in Tropasteron.

Character

Plesiomorphic

Apomorphic

1

Abdominal pattern

2 pairs of white spots

with additional frontal and dorsal pairs

2

DtiA

roughly triangular

with 2 sharp tips (IP, EP)

3

DtiA external prong EP

short, about as long as wide

long, at least twice as long as wide

4

DtiA. EP. tip

sharp

distal ly flattened

5

DtiA, EP, basally

without bump

with basal bump

6

Cymbium retrolaterally

with flange

with additional small rounded extension

7

Flange on cymbium

straight

hook-shaped

8

Flange on cymbium

at least 1/3 of cymbium long

10-15% length of cymbium

9

LTA

not separated from tegulum

spine-like separated from tegulum

10

Base of embolus

not separated from tegulum

separated from tegulum, cylindrical

11

DTA

simple, spoon-shaped

semicircular, distally folded with short PE

12

Epigyne CO

inverted u or V-shaped

semicircular, to flat oval or lyriform

13

Epigyne CD

CD short only few loops

elongate with big semicircular distal loops

NEW QUEENSLAND SPIDER GENUS

63

FIG. 32. Distribution of Tropasteron species T. andreae , T. magnum , T.
pseudomagnum , T robertsi.

halifax), 15 (Hinchinbrook I: T.
cardwell ), 5 (Hann Tbld.: 0), 3
(Windsor Tbld.: T. luteipes).

In contrast to the above
mentioned beetles, only about 40
% of the species (9 species) are
endemic to 1 single rainforest
block. The distribution ranges of
most species are over 2 sometimes
3 adjacent rainforest blocks
(Table 2). Hence, the main centre
of species diversity is in the
northern central part of the
rainforest blocks in the Wet
Tropics of Northern Queensland.

According to their distribution
pattern, both T. andreae- group
and T. Cleveland- group must have
diverged or evolved before
Eungella/Carmila and Mt
Cleveland — where the most
plesiomorphic species, T.
andreae , I Cleveland and T.
raveni , exist — were isolated
from the main rainforest block.

Significantly, sympatric species
have divergent abdominal patterns.

Within the WTWHA, synapo-
morphies could be found only for
the I andreae and T. Cleveland
group, but not for groups of
species within each species group.

All other species differ from each
other in small differences in palp
(length of LTA, length of EP) and
or in colour pattern but no
evolutionary pattern is evident. At
least within Tropasteron , parallel
evolution obviously took place,
when, as a result of ancient arid incursions, the
rainforest blocks split up.

ACKNOWLEDGEMENTS

I thank Michael Gray and Graham Milledge,
Australian Museum (Sydney), Robert Raven and
Wendy Hebron, Queensland Museum
(Brisbane), for loans of the material and support
of the work. For an excellent working
atmosphere I thank Valerie Davies, for inform-
ative and critical comments Geoff Monteith, for
being patient my children Johanna and Ursula
Baehr, and for preparing the maps, providing
critical comments on the manuscript and help
with the scanning electron microscope, Robert

Raven. Last, 1 thank Marshall from the
Headlands Chalet (North Stradbroke Island) for
providing a wonderful place and spirit tor
finishing the manuscript. The study is funded by
the Australian Biological Resources Study
Participatory Program.

LITERATURE CITED

BAEHR, B. & JOCQUE, R. 1 996. A revision of Asteron ,
starring male palpal morphology (Araneae,
Zodariidae). Proceedings of the XII I International
Congress of Arachnology, Geneva, 3-8
September 1995. Revue suisse Zoology, hors
seriel: 15-28.

2000. Revisions of the genera in the Asteron-
complex (Araneae, Zodariidae). The new genera

64

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Distribution of the Tropasteron species in
the Mountain Rainforest Blocks (Fig. 28 A), defined
by Monteith (1995) modified in Bouchard (2002).
Recorded altitudes are given. Number of examined
specimens (in brackets); species in bold occur only at
one block. Species are listed from North to South.

Species (specimens)

Altitude

Locality

T. robertsi (2)

1100m

1

T. tribulation ( 1 8)

50-730m

Iron Ra., 1, 2

T. magnum (2)

500m

2

T. pseudomagnum (4)

1250m

4

T. splendens (24)

700- 1200m

1,7

T heatherae (50)

440- 1330m

2,4

T. luteipes ( 1 2)

1100-1 180m

3,4

T. monteithi (63)

600-1 100m

4

T. thompsoni (28)

700- 1200m

6, 7, 8, 9

T. julatten (36)

750-1 100m

6, 8,9

T. cooki (23)

850- 1000m

7,11

T eacham (27)

750- 1000m

8,9

T. malbon (19)

500-900m

10, 11

T. yea test ( 106)

1054- 1560m

10

T. daviesae (22)

800- 1000m

12

T. palmers ton (15)

670m

12

T. cardwell (74)

250-800m

11, 13, 15

T. fox (60)

500-650m

13, 14

T. Halifax (87)

720- 1050m

14, 16

T. Cleveland (42)

500-560m

Mt Cleveland

T. raven i (44)

350m

C Old, Carmila

T. andreae (3)

350m _

Eungclla NP

Cavasteron and Minasteron. Records of the
Western Australian Museum 20: 1-30.

2001. Revisions of the genera in the Asteron-
complex (Araneae, Zodariidae). The new genera
Pentasteron , Phenasteron , Leptasteron and
Subasteron. Memoirs of the Queensland
Museum 46(2): 359-385.

BAEHR, B, 2003. Three new endemic genera of the
Asteron- complex (Araneae: zodariidae) from
Australia: Basasteron , Euasteron and Spinasteron
gen. nov. Memoirs of the Queensland Museum
(this volume).

BAEHR, M. 1995. Revision of Philipis (Coleoptera:
Carabidae: Bembidiinae), a genus of arboreal tachyne
beetles from the rainforest of eastern Australia.
Taxonomy, phylogeny and biogeography.
Memoirs of the Queensland Museum 38(2): 3 1 5-38 1 .

BOUCHARD, P. 2002. Phylogenetic revision of the
flightless Australian genus Apterotheca Gebien
(Coleoptera: Tenebrionidae: Colcometopinae).
Invertebrate Systematics 16: 449-554.

DALLWITZ, M.J., PAINE, T.A. & ZURCHER, E.J.

1998. Interactive keys. Pp. 201-212. In Bridge, P.,
Jeffries, P., Morse, D.R. & Scott, P.R. (eds)
Information technology, plant pathology and
biodiversity. (CAB International: Wallingford).

DAVIES, V.T. 1995. A new spider genus (Araneae:
Amaurobioidea: Amphinectidae) from the wet
tropics of Australia. Memoirs of the Queensland
Museum 38(2): 463-469.

1 999. A new spider genus from North Queensland
(Araneae: Amaurobioidea: Kababinae). Journal
of Arachnology 27(1): 25-36.

DAVIES, V.T. & LAMBKJN, C.L. 2000a. Malarina a
new spider genus (Araneae: Amaurobioidea:
Kababinae) from the wet tropics of Queensland.
Memoirs of the Queensland Museum 45(2):
273-283.

2000b. Wabua a new spider genus (Araneae:
Amaurobioidea: Kababinae) from North
Queensland. Memoirs of the Queensland
Museum 46(1): 129-147.

JOCQUE, R., 1991. A generic revision of the spider
family Zodariidae (Araneae). Bulletin of the
American Museum ofNatural History 201: 1-160.

JOCQUE, R. & BAEHR, B. 2001. Revisions of the
genera in the /U7eraw-complex (Araneae,
Zodariidae). Asteron Jocque and the New Genus
Pseudastemn. Records of the Australian Museum
53:21-36.

MONTEITH, GB. 1995. Distribution and altitudinal
zonation of low vagility insects of the Queensland
Wet Tropics. A report to the Wet Tropics
Managemant Authority. Part 4. (Queensland
Museum: Brisbane).

1997. Revision of the Australian flat bugs of the
Subfamily Mezirinae (Insecta: Hemiptera:
Aradidae). Memoirs of the Queensland Museum
41(1): 1-169.

RAVEN, R.J. 1 978. Systematics of the spider subfamily
Hexathclinae (Dipluridac: Mygalomorphae:
Arachnida). Australian Journal of Zoology
Supplementary Series 65: 1-75

2000. A new species of funnel-web spider
Hadronyche (Hexathelidae: Mygalomorphae)
from North Queensland. Memoirs of the
Queensland Museum 46(1): 225-230.

REID, C.A.M. & STOREY, R.I. 2000. Revision of the
dung beetle genus Temnoplectron Westwood
(Coleoptera : Scarabaeidae: Scarabaeini). Memoirs
of the Queensland Museum 46( 1 ): 253-297.

STOREY, R.I. & MONTEITH, GB. 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.

PSYDRINE GROUND BEETLES (COLEOPTERA: CARABIDAE: PSYDRINAE),
EXCLUDING AMBLYTELINI, OF EASTERN QUEENSLAND RAINFORESTS

MARTIN BAEHR

Baehr, M. 2003 06 30: Psydrine ground beetles (Coleoptera: Carabidae: Psydrinae),
excluding Amblytelini, of eastern Queensland rainforests. Memoirs of the Queensland
Museum 49(1): 65-109. Brisbane. ISSN 0079-8835.

The psydrine ground beetles of eastern Queensland, excluding the tribe Amblytelini, are
reviewed. Species of Laccocenus Sloane, Teraphis Castelnau, Trephisa Moore, and Meonis
Castelnau are enumerated, the Queensland species of Mecyclothorax Sharp revised, and
Sitaphe Moore and Raphetis Moore fully revised. The following new taxa are described:
Mecyclothorax inflatus from southern parts of Atherton Tableland, M. inflatus spinifer from
the Walter Hill Range, M. impressipennis from Isley Hills north of Bellenden Ker Range, M.
storeyi frerei from Bartle Frere, M. lewisensis uncinatus from Mt Hemmant and Mt Halcyon
north of Thornton Peak, Sitaphe trapezicollis from Bellenden Ker/Bartle Frere Ranges, S.
parvicollis from Bartle Frere Range, S. minuta from Lambs Head, Mt Williams, and Isley
Hills north of Bellenden Ker Range, S. hamifera from Cardwell Range, S. incurvicollis from
the Walter Hill Range, 5. parallelipennis from Carbine and Windsor Tablelands, S. ovipennis
from Thornton Peak, Raphetis curia from Springbrook Plateau in southeastern Queensland,
and four subspecies of R. gracilis Moore, all from northeastern Queensland: R. g. frerei from
Bellenden Ker/Bartle Frere Ranges and vicinity, R. g. spinosa from the surroundings of Mt
Lewis, R. g. spurgeoni from north of Mt Spurgeon, and R. g. suharmata from Mt Spurgeon
and Plane Crash Site, the last three subspecies being from different parts of Carbine
Tableland

Within the three revised genera similar patterns of distribution are recognised: all species are
montane, and the many extremely similar taxa each occupy single or closely adjacent
mountain tops or tablelands. This distribution pattern reflects similar patterns in various
invertebrate groups of low vagility occurring in the Wet Tropics of North Queensland. The
high level similarity is strong evidence of a rather recent - probably even Pleistocene
allopatric speciation caused by vicariancc events due to repeated spreading and retreat of the
montane rain forests during the glacial and intcr-glacial periods.

Within the strictly northern Sitaphe , and also in the storey/ -group of Mecyclothorax ,
relationships are still obscure. In Raphetis a fairly distinct gradient exists between the
plesiotypic southern R. curta , the intermediate R. darlingtoni from mid-eastern Queensland,
and the apotypic R. gracilis- complex in north Queensland. □ Coleoptera, Carabidae,
Psydrinae, Mecyclothorax, Sitaphe, Raphetis, new species, Australia, Queensland,
distribution.

Martin Baehr, Zoologische Staatssammlung, Miinchhausenstr. 21, D-81247 Miinchen,
Germany (e-mail: martin.baehr@zsm.mwn.de); 15 April 2003.

By courtesy of Geoff Monteith of Queensland
Museum, Brisbane, I received specimens of the
psydrine genera Mecyclothorax Sharp, Sitaphe
Moore and Raphetis Moore collected during the
extensive rain forest sampling program carried
out by him and his co-workers on various
mountain tops along the east coast of Queensland
during the last 20 years. When I requested some
voucher specimens of Raphetis for comparison
during the course of a forthcoming revision of the
psydrine tribe Amblytelini, Geoff Monteith asked
me to review all specimens because there appeared
to be several different - probably undescribed
— forms. We agreed, then, that it would be best to
combine descriptions of new forms of all genera

in a single comprehensive paper to give an
overview of the geophile psydrines of the rain
forests of eastern Queensland.

This paper, therefore, includes a revision of
Queensland Mecyclothorax , and complete
revisions of Sitaphe and Raphetis. Other psydrine
species that occur near the Queensland/New
South Wales border (mostly on Lamington Plateau)
are mentioned for completeness. Queensland
species of Meonis Castelnau, a genus that urgently
merits a thorough revision, are mentioned
without further commentary.

The tribe Amblytelini is very speciose in
eastern Queensland (c. 40 taxa at the present state
of knowledge) but is omitted from this paper

66

MEMOIRS OF THE QUEENSLAND MUSEUM

because a general revision of that tribe will be
printed elsewhere (Baehr, in press a).

More general information about extent,
distribution, and phylogeny of the Psydrinac is or
will be available from Baehr (1999. in press a).
Here I will direct the readers’ attention to the use
of taxonomic categories higher than genus. I use
the term subfamily for the whole psydrine
complex, and tribe for such units as Amblytelini,
Mecyclothoracini. etc. Other authors rank the
tribes as subtribes and the subfamily as a tribe.
This is, however, a matter of opinion and is of no
serious relevance to the present treatment.

Raphetis and Sitaphe were described in
Moore’s generic revision of the Australian
Psydrinae (Moore, 1963) which is still the basis
of all work on this group in Australia. In it, Moore
described a single species of Sitaphe and two
species of Raphetis , the types of which I have
examined. Moore (1984) later described two
outstanding species of Mecyclothorax from rain
forests of North Queensland, and these are
reviewed in the present paper.

In tropical Queensland, apart from the tribe
Amblytelini, only the three genera that are herein
revised occur. Curiously enough, the many
mountain tops and tablelands south of Atherton
Tableland except for Lamington Plateau and
adjacent ranges on the Queensland/New South
Wales border, although having been sampled
with comparable intensity during the last decade,
apparently lack a similarly diverse psydrine
fauna. It seems, thus, that except for Raphetis
darlingtoni Moore at Eungella, west of Mackay,
the Psydrinae are almost absent from the many
rainforest patches south of the Wet Tropics, and
that this absence indicates a real distribution gap.

MATERIAL AND METHODS

Almost all material for this study was collected
by Geoff Monteith and his co-workers of the
Queensland Museum, Brisbane (QM), and the
bulk of the material is in that collection, except
for duplicates lodged in the working collection of
the author (CBM) in Zoologische Staatssammlung,
Munchen. Altogether 542 specimens of
Mecyclothorax of the storey i-g roup, 290
specimens of Sitaphe and 38 specimens of
Raphetis were available for this study. In addition,
about 80 specimens of southern Queensland
species of Laccocenus , Mecyclothorax , Trephisa ,
Teraphis and Meonis were noted.

The types of B.P. Moore’s species were kindly
loaned from the Australian National Insect

Collection, Canberra (ANIC). Further abbrev-
iations of collections are: CMC, Collection B.P.
Moore, Canberra, in ANIC; DPI, Department of
Primary Industries, Mareeba.

Specimens for male genitalia dissection were
soaked in a moist jar overnight, then the genitalia
were cleaned briefly in hot 4% KOH. The
photographs were made using SPOT Advanced
for Windows 3.5 and subsequently were worked
with Corel Photo Paint 10.

Holotype label data are given in full but for
other specimens collectors’ names and other
terms are abbreviated as follows: DC, D.J. Cook;
DY, D.K. Yeates; GM, G.B. Monteith; GT, G.I
Thompson; HJ, H.A. Janetzki; LR, L. Roberts;
PB, P. Bouchard; RS, R. Sheridan; SM, S.R.
Monteith; Pyr., pyrethrum knockdown; QM
Berl., Queensland Museum Berlesate No.; Rf,
rainforest; Tbld, Tableland.

Descriptions. Because of similarity in external
and genitalic characters within most taxa of
Mecyclothorax , Sitaphe , and Raphetis , generally
one species of each genus is fully described, then
for the other taxa only characters that differ are
noted. This is done so as to reduce repetition, but
was not applied to species that differ more
substantially.

Measurements. Measurements were taken using
a stereo microscope with an ocular micrometer.
Length has been measured from apex of labrum
to apex of elytra. Lengths, therefore, may slightly
differ from those of other authors, especially
Moore (1963, 1984). Length of pronotum was
measured along midline, width of pronotum at
w idest part, w idth of base of pronotum at the
extreme tips of the basal angles, though in those
species of Sitaphe that have the lateral margin
incurved to the basal angle, width of base was
measured immediately in front of the basal angle.
In the measurements of Raphetis species length
of eye includes a small dark coloured ring of
ocellae behind the light area. Ratios are some-
what variable in most species, but generally offer
good indication of relative shape.

Taxonomic Principles. Interpretation of the
taxonomic status of the many highly similar
populations is difficult in all genera that occur in
the Wet Tropics. For the time being, at least until
additional distribution information is available, I
have decided to treat those populations that show
slight though apparently constant differences as
subspecies, when they are allopatric, and as
species, when they are obviously sympatric (e.g.
in Sitaphe). In a couple of taxa the morphological

PSYDRINE GROUND BEETLES

67

differences, in either the male genitalia or
external structure, are very weak and do not
justify description as species. Because several
other carabid genera (and also some non-carabid
groups) on mountain tops in the Wet Tropics of
northeastern Queensland are likewise segregated
into many closely related but apparently well
distinguished units (be they species or sub-
species!) on adjacent mountain ranges (e.g.,
Darlington, 1961a, 1961b; BaehrT 1995b;
Monteith, 1997; Bouchard, 2002), this treatment
seems to accommodate the complex situation
best. Other non-morphological methods may be
useful to interpret the taxonomic status of the
different populations. For further comments see
‘Discussion’.

Although ranges are included in the keys, range
should not be used as a distinguishing character
per se, even when the ranges of most taxa are
apparently restricted and well separated.

Laccocenus Sloane, 1890

Laccocenus Sloane, 1890: 644; C'siki 1929: 484; Moore
1963:286, Moore et al. 1987: 153.

TYPE SPECIES. Laccocenus ambiguus Sloane, 1 890.

REMARKS. 1 his is the sole Australian genus of
the tribe Nomiini s.str. Until recently monotypic,
an additional species has been found in a cave in
southeastern New South Wales and is being
described at present (Moore, in press).

Laccocenus ambiguus Sloane, 1890

Laccocenus ambiguus Sloane, 1890: 646; Csiki 1929: 484;
Moore 1963: 286; Moore et al. 1987: 153

REMARKS. A rather common species living in
and under rotten wood or under bark of fallen
logs in subtropical to temperate rain forest.

DIS1 RIBU I ION. Northeastern New South Wales
(type locality: Dunoon, near Lismore) and at the
following high elevation localities (QM speci-
mens) along the Macpherson Range, Queensland:
(east to west) Tomewin Range; Repeater Station,
Springbrook; Mt Bithongabel and Westcliff
Track, Lamington Plateau; Mt Chinghee; Mt
Superbus; Bald Mountain, via Emu Vale.

Mecyclothorax Sharp, 1903

Mecyclothorax Sharp. 1903: 243; Csiki 1929: 487; Moore
1963: 286; Moore 1984: 161; Moore et al. 1987: 147.

TYPE SPECIES. Cvclothorax montivagus Blackburn,
1878.

REMARKS. A genus of the tribe Mecyclothoracini
that also includes Neonomius Moore in Australia.

Mecyclothorax is widely distributed throughout
eastern, southern and southwestern Australia,
and altogether 16 species were described from
Australia. Moore ( 1 984), who published a partial
revision of the genus (the ambiguus- group) and
described two new species from North Queens-
land, stated that the genus urgently needs revision.

Outside Australia, Mecyclothorax is widely
distributed in New Guinea, New Caledonia,
Borneo, Java, Hawaii, Tahiti, New Zealand,
Norfolk Island, Lord Howe Island and several
subantarctic islands, e.g. Amsterdam and St. Paul
(Baehr 1 992, 1 995a, 1 998, 1 999, 2000, in press b.
Baehr& Lorenz 1999, Britton 1948, Darlington
1962, 1971, Deuve 1987, Jeannel 1944,
Louwerens 1949, Mandl 1969, Moore 1984,
1985, 1992, Moore et al. 1987, Perrault 1978,
1992). In Australia a distinct, probably apotypic,
group of tropical species is well separated from
the range of the main body of the genus in
Australia which is essentially southern.

KEY TO THE QUEENSLAND TAXA OF
MECYCLOTHORAX SHARP

1 . Pronotum almost orbicular, lateral margin shortly excised

in front of basal angles, elytra longer, striae deeply
punctate. Size >5mm. Central and southeastern

Queensland punctipennis (Macleay)

Pronotum more or less cordate, lateral margin not excised
in front of basal angles, elytra shorter and wider, striae
mostly barely punctate. Size <3. 5mm 2

2. Clypeus bisetose; base of pronotum coarsely punctate,

basal angles without seta, anterior transverse impression
deep: aedeagus abnormal, side-inverted and turned to the

right side 3

Clypeus quadrisetose; base of pronotum not or barely
punctate, basal angles usually with setae, anterior
transverse impression barely indicated; aedeagus
normal, turned to the left side * 4

3. Pronotum not sinuate in front of base; aedeagus larger,

apex shorter and wider, internal sac with two small
spinose areas in front (Fig. 1A). Bellcnden Ker Range.
Massey Range storey i storey i Moore

Pronotum slightly sinuate in front of base; aedeagus
smaller, apex longer and narrower, internal sac without
spinose areas in front (Fig. IB). Bartlc Frere Range
storeyi frerei subsp. nov.

4. Lateral margins of pronotum not perceptibly sinuate

posteriorly: genital ring with very elongate apex (Fig.
IE) 5

Lateral margins of pronotum perceptibly sinuate
posteriorly; genital ring, when known, with shorter apex
(Figs 1C-D) 6

5. Spinose fields within apex of orificium of aedeagus

smaller, situated at left and right sides (Fig. IE).
Southwestern Part of Atherton Tbld: Mt Fisher, Mt Hugh
Nelson, Mt Father Clancy, Malaan Rd nr Palmerston
Hwy inflatus inflatus sp. nov.

Spinose fields within apex of orificium of aedeagus very
large, situated only at the right side (Fig. IF). Upper

68

MEMOIRS OF THE QUEENSLAND MUSEUM

Boulder Creek at Walter Hill Range

inflatus spinifer subsp. nov.

6. Four inner striae of elytra impressed and coarsely

punctate. Isley Hills, ne. Atherton Tbld

impressipennis sp. nov.

Inner striae of elytra at most lightly impressed and finely
punctate 7

7. Apex of aedeagus rounded off, genital ring with longer

apex (Fig. 1C); pronotum with comparatively wider
base, ratio base/apex >1.20. Carbine Tbld, Thornton
Peak, Mt Pieter Botte, north of Thornton Peak
.............. lewisensis lewisensis Moore

Apex of aedeagus sharply spined, genital ring with
considerably shorter apex (Fig. ID); pronotum with
comparatively narrower base, ratio base/apex <1. 1 7. Mt
Hcmmant, Mt Halcyon, Roaring Meg Ck, mountain tops
near Cape Tribulation, all north of Thornton Peak
lewisensis uncinatus subsp. nov.

Mecyclothorax punctipennis (Macleay, 1871)

Cyclothorax punctipennis Macleay, 1871: 105.
Mecyclothorax punctipennis, Csiki, 1929: 487; Moore, 1984:

162; Moore ctal., 1987: 149.

Cyclothorax obsoletus Blackburn, 1889: 1389.

REMARKS. A common species in southeastern
Queensland, where it is found in subtropical rain
forest on Lamington Plateau, Main Range, Bunya
Mountains, and further north to about Gayndah
(the type locality) where it has been recollected
recently (QM). One rather recent record is
available from Blackdown Tableland further
north (QM). There, and at scattered localities in
low country, the species also occurs in more open
habitats. It lives on the ground, but also on and
sometimes even under bark of logs and standing
trees. Moore ( 1 984) demonstrated the differences
between M. punctipennis and the rather similar
M arnbiguus Erichson, under which name M.
punctipennis was still noted by Csiki (1929).

DISTRIBUTION. Whole southern Australia from
southern half of Western Australia to southern
Queensland as far north as Tropic of Capricorn
(Moore et al., 1 987). Recently recorded also from
Tasmania (Baehr, 2000).

Mecyclothorax storeyi Moore, 1984

Mecyclothorax storeyi Moore, 1984: 164; Moore et al., 1987:
149.

Small, conspicuously coloured species living
at high altitude in rain forest litter. So far known
from Bellenden Ker, Bartle Frere, and Massey
Ranges at the eastern margin of Atherton
Tableland. As the population living on Mt Bartle
Frere shows significant differences in shape and
structure of the male aedeagus, it is described as a
distinct subspecies.

M. storeyi is peculiar in possessing strikingly
abnormal male genitalia, because aedeagus and

parameres are side-inverted and moreover, are
turned to the right side of the beetle, which is
opposite to all other species of Mecyclothorax
and to Psydrinae in general. During ample
dissections of almost 100 species and several
hundred specimens of the amblytelines Ambly -
telus, Dystricho thorax, and Epelyx ; 1 found a
similar inversion only twice in single specimens
of two species that normally possess nonnal
shaped aedeagi turned to the left side of the body.
Moore (1984, fig. 15) recognised this inversion,
but confused the parameres, describing the left
one as ‘small, styloid, setose laterally’ and the
right one as ‘larger, conchoid, setose apically’.
Nevertheless the parameres are normal, but the
whole male genitalia are inverted which means
that the left paramere in situ is the right one
morphologically, and vice versa.

To clarify the situation, and because the species
includes two subspecies that differ in certain
characters of the aedeagus, and finally, because
the setosity of the parameres is incorrectly
figured in the description, the genitalia of both
subspecies are (re)described and figured herein.

DIAGNOSIS. Small species, distinguished by
the side-inverted aedeagus with straight instead
of downcurved apex, bisetose clypeus (as usual
for the genus), absence of the posterior pronotal
seta, deeply impressed anterior transverse sulcus
of pronotum, and rather narrow base of the
basally coarsely punctate pronotum.

REMARKS. With respect to several differences
between this and the following small, flightless
species from North Queensland, M. storeyi is a
rather isolated species within this group, whereas
all following species form a distinct group of very
closely related taxa.

Mecyclothorax storeyi storeyi Moore, 1984
(Figs 1A, 2A, 5)

Mecyclothorax storeyi Moore, 1984: 164; Moore et al. 1987*

149.

MATERIAL. HOLOTYPE: 6 , N Qld, Mt Bellenden Ker
(summit), from leaf litter, 1911977, B.P. Moore (ANIC).
PARATYPES: 35 ex. from same locality (in ANIC, CMC
DPI, and QM).

NEW RECORDS: 6 6 , 24 9 , Bellenden Ker, Centre Peak
Summit, I O.iv. 1 979, GM. QM Berl. No. 9 17.16S 145.5 IE
Rf, 1500m Sieved litter (CBM, QM); I 9, same loc.,

1 O.iv. 1 979 GM, QM Berl. 12 Rf, Stick brushings (QM); 9
6,1 9 , same loc., 1 1 .iv. 1979 GM, QM Berl. Rf, Sieved
litter (QM); 15 6,5 9, same loc., 1 1 .iv. 1979 GM /QM
Berl. 1 4 ,Rf, Sieved litter (QM); 1 6 , same loc., 1 1 .iv. 1 979
GM, QM Berl. 15, Rf, Stick brushings (QM); 2 9, same

PSYDRINE GROUND BEETLES

69

loc., 1 1 .iv. 1979 GM, QM Berl. 16, Rf, Stick brushings
(QM); 4 <$ , 1 9 , same loc., 1 1 .iv. 1 979 GM, QM Berl. 1 7,
Rf, Sieved litter (QM); 1 <$,4 9, same loc., 28.viii.1991
GM & HJ, QM Berl. No. 852, Rf, 1560m , Sieved litter
(QM); 1 9 , Bellenden Ker, Cable Tower No. 3, 12.iv. 1 979
GM, QM Berl. 2 1 1 7. 1 6S 1 45.52E Rf, 1 000m Sieved litter
(QM); 4 <$, 5 9, Bellenden Ker Summit 10.vi.1980 GM.
QM Berl. No. 220 17°16’S 145 D 52'E Rf, 1561m Sieved
Litter (QM); 8 <$#, 4 9 , Bellenden Ker Range. NQ
Summit TVStn., 1560m, 17.16S 145.5 IE, Nov. 1-7, 1981
Earthwatch/QM, QM Bed. 334, Rf Sieved litter (QM); 5
c$#,2 9 , same loc., Nov. 1-7, 1981 Earthwatch/QM, QM
Berl. 335, Rf Sieved litter (QM); 8 <$#, 5 9, same loc.,
Nov. 1-7, 1981 Earthwatch/QM, QM Berl. 336, Rf Sieved
litter (CBM. QM); 8 <$#, 6 9 ,samc loc., Nov. 1-7, 1981
Earthwatch/QM, QM Berl. 337 Rf Sieved litter (QM); 7
($#, 2 9 , same loc., Nov. 1-7, 1981 Earthwatch/QM, QM
Berl. 338, Rf Sieved litter (QM); 1 9 , same loc. Nov. 1-7,
1981 Earthwatch/QM, QM Berl. 343, Rf Stick & Moss
brushings (QM); II <3, 8 9 , same loc. Oct 25-31, 1981
Earthwatch/QM, QM Berl. 371, Rf Sieved litter (CBM,
QM); 5 dm, 3 9 , same loc. Oct 25-31, 1981
Earthwatch/QM, QM Berl. 372, Rf Sieved litter (QM); 5
<$,3 9, same loc. Oct 25-31, 1981 Earthwatch/QM, QM
Berl. 374, Rf Dracophyllum litter (QM); 9 d, 5 9 , same
loc. Oct 25-31, 1 98 1 Earthwatch/QM, QM Berl. 375, Rf
Sieved litter (QM); 2 d, 3 9 , same loc.Oct 20-23, 1981
Earthwatch/QM, QM Berl. 376, Rf Sieved litter (QM); 8
d , 6 9 , same loc. 28.x. 1 983 GM, DY & GT, QM Berl. No.
601, Rf, Sieved litter(QM); 11 d,9 9, same loc. 28.x. 1983
GM, DY & GT, QM Berl. No. 602, Rf, Sieved litter (CBM,
QM); 2 d, same loc., 16 April 1999 GM & SM, QM Berl.
993, Rf, Moss ex trees & logs (QM); 1 d, same loc., 17
April 1997 GM & Russell, QM Berlesate 930 17°16'S
145°52’E Rf, Sieved leaf litter (QM); 6 d, 3 9, same loc.

1 .xii. 1 998 GM, QM Berl. 977, Sieved litter (QM); 5 d, 1
9 , AUST: Qld: NE: Bellenden Ker, 1 994 Crash. 1 .xii. 1 998
GM, QM Berl. 978 17°16’S 145°5PE Rf, 1325m Sieved
litter (QM);2 9, Mt Bellenden Ker Qld rain forest
12.xii. 1976 Wal ford- Muggins / Mecyclothorax storeyi
Moore (CBM); 1 d, Massey Range, NQ 12km S
Gordonvale 2.v. 1 983 GM, DC, QM Berl. No. 573 17.16S
1 45.59E Rf, 1 300m sieved litter (QM );6 d , 3 9 , NE Qld
Thornton Peak, via Daintree, 20-22.be. 1981 GM & DC,
QM Berl. 30 1 Rf, 1 000- 1 300m Sieved litter & moss (QM)
(probably wrong label! See further discussion under M.
lewisensis).

DIAGNOSIS. As the species was described from
individuals from Bellenden Ker Range, this
population is the nominate subspecies. It is
distinguished from the southern population
living on Mt Bartle Frere, M. s.frerei subsp. nov.,
by considerably larger aedeagus bearing a shorter
and wider apex and two small spinose fields at the
opening of the internal sac, and by wider base of
pronotum the lateral margins of which, on
average, are less sinuate near base.

DESCRIPTION. Measurements. Length: 2.8-
3.25mm; width: 1.35-1. 5mm; Ratios. Length/

width of 9th antennomere: 1. 8-2.0; width/length
of pronotum: 1.24-1.30; width base/apex of
pronotum: 1.10-1.15; width pronotum/head:
1.51-1.60; length/width of elytra: 1.24-1.27;
width elytra/pronotum: 1.36-1.38.

Colour. As in the southern subspecies,
colouration of the nominate subspecies is very
diverse. Although head and pronotum are always
black, colour of the elytra can vary from almost
completely black, to completely reddish, to
blackish or dark reddish with light margin, or
with light humeral area only, or quadrimaculate
with light humerus and light apex.

Male genitalia (Fig. 1A). Genital ring short and
wide, highly asymmetrically triangular, with
characteristic angle laterally, apex narrow and
rather elongate. Aedeagus side-inverted, turned
to the right side (in beetle), narrow and elongate
(in genus), lower surface evenly concave. Apex
fairly elongate, wide, straight, evenly rounded
off. Internal sac rather complexly folded, with
several narrow, sclerotized plates within. On left
side of apical end of (inverted) internal sac with
two strongly spinose fields. Both parameres
comparatively elongate, triangularly convex,
with narrow, elongate apex. Left (in situ right!)
paramcre larger than right (in situ left!), with 1 -2
short apical setae. Right (in situ left!) paramere
with 2 apical and 5-8 moderately elongate setae
along the apical half of lower margin.

Female genitalia (Fig. 2A). Stylomere 1 with
one, rarely two elongate ensiform seta(e) at
lateral part of ventro-apical margin. Stylomere 2
rather short, with short apex and two large
dentiform ventro-lateral ensiform setae of about
similar size below middle of lateral margin. Near
apex with a large, oblong pit and a short
nematiform seta originating from that pit. In
middle of dorso-median surface with a large,
dentiform, dorso-median ensiform seta. Lateral
plate with a densely setose area at median apical
margin.

Variation. A rather variable subspecies with
respect to relative shape and, in particular, to
colour and distinctness of pattern of elytra. Also
puncturation of base of pronotum, and degree of
puncturation and depth of elytral striae vary to
some extent.

DISTRIBUTION (Fig. 5). Bellenden Ker and
Massey Ranges at the eastern margin of Atherton
Tableland.

COLLECTING CIRCUMSTANCES. Generally
collected by Berlese extraction or by sieving of

70

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 1 . Male genitalia o ^ Mecyclothorax spp. A, M. storeyi storeyi Moore, aedeagus, parameres, and genital ring;
B, M. storeyi frerei subsp. nov., aedeagus and parameres; C, M, lewisensis lewisensis Moore, aedeagus!
parameres, and genital ring; D, M lewisensis uncinatus subsp. nov., aedeagus, parameres, and genital ring; E,
M. injlatns inflatus sp. nov., aedeagus, parameres and genital ring; F, M. inflatus spinifer subsp. nov., aedeagus
and parameres. Scales: 0.25mm.

ground litter from upland rain forest. A few were
collected by ‘stick brushings’ and in pitfall traps.
Most specimens are from the absolute summits of
the respective ranges.

Mecyclothorax storeyi frerei subsp. nov.
(Figs IB, 3A-B, 4A, 5)

ETYMOLOGY. The name refers to the type locality, Mt
Bartle Frere, Queensland’s highest peak.

MATERIAL. HOLOTYPE: <J, QMT21216, NE Qld, Mt
Bartle-Frerc, summit creek, 24.ix.1981 G Monteith & D.
Cook, QM Berl. 304 Rf. 1500m Sieved litter (QM).
PARATYPES: 2 6 , 3 9, same data (CBM, QM); 9 <J, 14
9 , Mt Bartle Frere, N Qld. 0.5km N of Sth Peak 6-8 Nov.
1981 Earthwatch/QM, QM Berl. 357 17.24S 145.49E Rf
Sieved litter (CBM, QM); 10 6,6 9, Mt Bartle Frere, N
Qld. NW/Centre Peak ridge 7-8.X.1981, 1400- 1500m
Earthwatch/QM, QM Berl. 358 17.23S 1 45. 48ERf Sieved
litter (CBM, QM); 6 6 , 10 9 , Mt Bartle Frere, N Qld. Sth
Peak Summit, 1 620m 6-8 Nov. 1 98 1 Earthwatch/QM, QM

PSYDRINE GROUND BEETLES

71

FIG 2. Stylomcre 2 and apex of stylomere 1. A,
Mecyclothorax storey i storey /; B, M. lewisensis
lewisensis ; C, Sitaphe rotundata Moore; D, Raphetis
curta sp. nov.; E, R. gracilis gracilis Moore. Scales;
0.1mm.

Berl. 354 17.24S 145.49E Sieved litter (CBM, QM); 3 <3,
same loc. 6-8 Nov. 1981 Earthwatch/QM, QM Berl. 359,
Sieved litter (QM).

DIAGNOSIS. This subspecies is distinguished
from the nominate subspecies by considerably
smaller aedeagus bearing a longer and narrower
apex and lacking the spinose fields at the opening
of the internal sac, and by narrower base of
pronotum, the lateral margins of which are more
distinctly sinuate towards base.

DESCRIPTION. Measurements. Length: 2.65-
3.15mm; width: 1.2- 1.45mm; Ratios. Length/
width of 9th antennomerc: 1 .9-2. 1 ; width/length
of pronotum: 1.29-1.32; width base/apex of
pronotum: 1.07-1.10; width pronotum/head:
1.43-1.50; length/width of elytra: 1.25- 1.29;
width elytra/pronotum: 1.33-1.34.

Colour (Fig. 4A). Black, though colouration of
elytra as variable as in nominate subspecies:
uniformly black, or reddish, of with light lateral
margin, or more or less conspicuously quadri-
maculate.

Head. As in nominate subspecies with clypeus in
middle remarkably convex, frontal furrows deep,
curved, prolonged onto clypeus, eyes convex,
laterally produced, with large orbits of c. 1/3 of
length of eye, with eyes separated from frons and
vertex by very deep sulcus. Surface absolutely
smooth.

Pronotum (Fig. 3A). As in nominate subspecies,
though anteriorly even wider and remarkably
convex, base relatively narrower and perceptibly

sinuate in front of the obtuse basal angles. Base
coarsely punctate and basal angle without basal
lateral seta, as in nominate subspecies.

Elytra (Fig. 3B). As in nominate subspecies.
Scutellar stria always lacking, sutural stria deeply
impressed and distinctly punctate, outer striae far
less impressed, commonly only punctate.

Lower surface. As in nominate subspecies.

Legs. As in nominate subspecies.

Male genitalia (Fig. IB). As in nominate
subspecies, though aedeagus smaller, with longer
apex, on lower surface slightly less curved, and
without the spinose fields at apex of the
(inverted) internal sac. Right (in situ left!)
paramere with only 4-5 setae on lower margin.
Female genitalia. As in nominate subspecies.
Variation. Little variation noted, except for
colouration of elytra, and depth of outer elytral
striae.

DISTRIBUTION (Fig. 5). Mt Bartle Frere, on the
eastern fringe of Atherton Tableland.

COLLECTING CIRCUMSTANCES. Most
specimens sampled by ‘sieving litter’ in montane
rainforest. This subspecies was collected only on
the summit plateau, above 1500m.

Mecyclothorax lewisensis Moore, 1984

Mecyclothorax lewisensis Moore, 1984: 165; Moore et al.

1987: 148.

REMARKS. Small, conspicuously coloured
species living at high altitude in rain forest litter.
So far known from Mt Lewis and certain local-
ities at the southern fringe of Carbine Tableland,
also from Thornton Peak and Mt Pieter Botte
north of Daintree River. As the northernmost
population living on Mt Hemmant, Mt Halcyon,
and Mt Sorrow near Cape Tribulation shows
significant differences in shape of the male
aedeagus, this population is described as a
distinct subspecies.

In contrast to M. storeyi, M. lewisensis has
normally shaped and directed male genitalia,
that, however, show rather apomorphic features
in the apex of the aedeagus and the left paramere.

There is some confusion as to presence or
absence of the basal prothoracic seta. Moore
(1984: 165-166) stated that 10 specimens he
examined from Thornton Peak lack this seta,
which is unusual and opposite to all other
specimens that I have examined. Through
courtesy of G. Monteith I had the opportunity to
see all QM material including the 10 mentioned

72

MEMOIRS OF THE QUEENSLAND MUSEUM

specimens from Thornton Peak. Thereby it
appeared that only one specimen belongs to M.
lewisensis which lacks the basal prothoracic seta
though not the puncture, whereas the nine other
specimens belong to M. storeyi. Apparently,
some confusion of labelling has occurred, because
it is extremely improbable that M. storeyi should
occur on Thornton Peak. As a conclusion,
Moore's (1984) statement that M. lewisensis is
highly variable in this character does not apply.

Because M. lewisensis includes two subspecies
that differ in certain characters of the aedeagus,
and moreover, because structures of the internal
sac are omitted and the setosity of the parameres
is incorrectly figured in the description (Moore,
1 984, fig. 1 6), the genitalia of both subspecies are
(re)described and figured herein.

DIAGNOSIS. Small, with cordate prothorax that
bears sharply angulate basal angles; clypeus
quadrisetose (as in all following species): dis-
tinguished by the apically very elongate male
genital ring. Further distinguished from M.
storeyi Moore by not side- in verted aedeagus and
the basal pronotal seta; from M. impressipennis
sp. nov. by narrower, less quadrate elytra with far
less deeply punctate inner striae and by narrower
base of pronotum; and from M. inflatus sp. nov.
by narrower base of pronotum bearing angulate
basal angles, and by less spinose apex of internal
sac of aedeagus, longer right paramere, and
presence of a membranous area near apex of left
paramere.

Mecyclothorax lewisensis lewisensis Moore
(Figs 1C, 2B, 6)

Mecyclothorax lewisensis Moore, 1984: 165; Moore et al.,

1987: 148.

MATERIAL. HOLOTYPE: d, N Qld, Mt Lewis,
16.i. 1 975, B.P. Moore (ANIC). PARATYPES: 23 ex. from
same locality (in ANIC, CMC, and QM).

NEW RECORDS. 1 d, 2 9, Australien, Qld. Mt Lewis,
1000m 6.1.1982 M. Baehrl Mecyclothorax lewisensis
Moore (CBM); 7 d, 4 9, Mt Lewis, road end 1 8.xi. 1 997
GM, QM Bert. 950, 16°30’S 145°15’E, Rf 1 120m Sieved
litter (CBM, QM); 1 d, 2 9, Mt Lewis Rd 29km from
Hwy 29.xi. 1 997 DC, QM Berl. 964 1 6°3 1 'S 1 45° 1 6’E, Rf
1210m Leaf litter (QM); 3 9, 2.5km N Mt Lewis via
Julatten 3jg.1983 DY & GT, QM Berl. No. 610 16.34S
145. 16E Rf, 1040m Sieved litter (QM); 2 d, 1 9, same
loc., 3.xi. 1 983 DY & GT, QM Berl. No. 61 1, Rf, 1040m
Sieved litter (QM); 2 d , same loc., 3.xi. 1983 DY GT, QM
Berl. No. 612, Rf, 1040m Sieved litter (QM); 2 d, 7.5km N
Mt Lewis, via Julatten, 8.ix. 1 98 1 GM & DC, QM Berl. 279
Rf, 1200m Sieved litter (QM); 1 d, 2 9 5.5km N Mt
Lewis, via Julatten, 8.ix. 1 98 1 GM & DC, QM Berl. 276 Rf,

FIG 3. Pronota and elytra of Mecyclothorax spp. A, B,
M. storeyi frerei subsp. nov.; C, D, M. lewisensis
uncinatus subsp. nov.; E, F, M. impressipennis sp.
nov.; QM inflatus spinifer subsp. nov.; H, M. inflatus
inflatus sp. nov.

1 100m Sieved litter (QM); 1 d, same loc., 8.ix. 198 1 GM
& IX, QM Berl. 275 Rf, Sieved litter (QM); 3 6 , 1 9 , Mt
Lewis, 18km N 23.xi.1998 GM, QM Berl. 975 16°30’S
145°16’E, Rf. 1300m Moss and litter(QM); 5 d,5 9 , Mt
Demi, 7km SW Mossman 29.x. 1983 DY GT, QM Berl.
No. 604 16.30S 145.19E Rf 1100m Sieved litter (CBM,
QM); 1 9, same loc., 29.x. 1983 DY GT, QM Berl. No.
603, Rf, Sieved litter (QM); 9 d , 3 9 , Hilltop. 5.5.km N Mt
Lewis. 13.Lx.1981 GM & DC, QM Berl. 297 Rf, 1200m
Sieved litter (CBM, QM); 13 d, 5 9,sameloc.,13.ix.l981
GM & DC, QM Berl. 297 Rf, 1200m Sieved litter (CBM,
QM); 2 9 , The Bluff, 1 1 km W of Mossman 29 April 1 983
GM, DY, QM Berl. No. 551 16.27S 145.16E Rf, 1050m
litter (QM); 1 d,2 9 , same loc., 29 Apnl 1983 GM, DY,
QM Berl. No. 554, Rf, 900- 1000m litter (QM); 1 9 , same
loc.29 April 1983 GM, DY, QM Berl. No. 555, Rf,
900- 1000m litter (QM); 1 9 , same loc., 29 April 1983

PSYDRINE GROUND BEETLES

73

GM, DY, QM Berl. No. 556, Rf, 900- 1000m litter (QM); 2
<J , 1 9 , same loc., 2.xi. 1983 GM, DY & GT, QM Berl. No.
609, Rf, 1 000m litter (QM); 1 9 , Carbine Tbld, N Qld
Mossman Bluff Camp, 30.xi.l990, 1000m GM & HJ
Pitfall Traps (QM); 6 <3, 3 9, Devils Thumb area 10km
NW Mossman 9.x. 1 982 GM, DY & GT, QM Berl. No. 456
1 6.34S 1 45. 1 7E Rf 1 000- 1 1 80m Sieved litter (QM): 2 6 . 6
9 , same loc. 9.x. 1 982 GM, DY & GT, QM Berl. No. 457,
Rf 1150m Sieved litter (QM); 2 d, 5 9, same loc.,
9.x. 1982 GM, DY & GT, QM Berl. No. 461, Rf. Sieved
litter (QM); 1 d, 1 9, same loc., 10.X.1982GM, DY&GT,
QM Berl. No. 459, Rf 1 1 80m Sieved litter (QM); 7 d , 7 9 ,
same loc.. 9- 10.x. 1982 GM. DY & GT, QM Berl. No. 455,
Rf 1000- 1 1 80m Sieved litter (CBM, QM); 2 d, same loc.,
9.x. 1982 GM, DY & GT, QM Berl No. 455, Rf,
1000- 11 80m Sieved litter/ Mecy>clothorax lewisensis
Moore Det. GM 1 989 (CBM); 1 9 , Devil’s Thumb - Pauls
Luck, 1 2km WNW Mossman, NQ 27 Ail 1 989 - 1 5.i. 1 990
ANZSES Expedition Site 12, 1300m, pitfall (QM); 9 d, 9
9 , Upper Whvanbeel Creek 5.ix. 1992 GM, QM Berl. No.
859 16°23' 145°17\ 1 1 50m Rf, litter (QM); 1 d,7kmNMt
Spurgeon (Camp 2) 17- 19.x. 1991. 1200- 1 250m 16°22’S
145°13'E GM, DC & LR Pitfall Traps (QM); 6 d, 2 9,
same loc., GM & HJ, QM Berl. No. 857 Rf. 1 250m Sieved
litter (CBM, QM); 4 d, 1 9 , 4km NNE Mt Spurgeon
15.x. 1991 GM & HJ,QM Berl. No. 854 16°24’S 145°13’E
Rf. 1250m Sieved litter (QM); 1 9, 3.5km NNE Mt
Spurgeon 15-20.X.199I, 1350m 16°24’S 145 D 13’E GM,
HJ, DC & LR. PITFALLS (QM); 1 d, Stewart Ck, 4km
NNE Mt Spurgeon (Camp 1), 1250-1 300m 16°24'S
145°1 3’E 1 5-20.X. 1991. PITFALLS GM, DC & LR (QM);

1 d, 2km SE Mt Spurgeon via Mt Carbine, N Qld.
20.xii.88-4.i. 1 989 GM. GT & ANZSES 1 1 00m, RF, pitfall
(QM ); 1 d, Thornton Peak, via Daintree, 20-22.ix.1981
GM & DC, QM Berl. 30 1 Rf, 1 000- 1 300m Sieved litter &
moss (QM); 3 d, 5 9 . Thornton Peak summit, via
Daintree 24-27.ix.1984 GM & SM, QM Berl. 662 Rf,
11 00- 1 300m Sieved litter & moss (CBM, QM): 3 d.
Granite Outcrops 0.5km E. Mt Pieter Botte 5.x. 1982 GM,
DY & GT, QM Berlesate No. 450 16.05S I45.23E Rf,
780m Sieved litter (CBM, QM).

DIAGNOSIS. As the species was described from
individuals from Mt Lewis, the population
showing similar genitalic morphology is the
nominate subspecies. It is distinguished from the
northern subspecies M. /. uncinatus subsp. nov.
by evenly rounded apex of aedeagus, generally
narrower pronotum with wider base, and, on
average, longer elytra.

DESCRIPTION. Measurements. Length: 2.8-
3.25mm; width: 1.35-1. 5mm; Ratios. Length/
width of 9th antennomere: 1 .65-1 .8; width/length
of pronotum: 1.32-1.36; width base/apex of
pronotum: 1.21-1.25; width pronotum/head:
1.48-1.54; length/width of elytra: 1.22-1.27;
width elytra/pronotum: 1.37-1.42.

Colour. Rather variable, blackish to more or less
dark piceous, elytra commonly lighter,

reddish-piceous to reddish, more or less
inconspicuously quadrimaculate, though apical
spots always present and rather distinct,
invariably somewhat oblique.

Pronotum. Contrary to the description (Moore
1984, p. 165) all examined specimens bear the
posterior lateral seta, including all specimens
from Thornton Peak. Most probably Moore
included in this species by error a wrongly
labelled sample of M. storeyi that, unfortunately,
also included a single specimen of M. lewisensis
with setae broken on both sides (series examined
by me).

Male genitalia (Fig. 1C). Genital ring elongate,
triangular, slightly asymmetric, apex narrow and
very elongate. Aedeagus normal, turned to the
left side (in beetle), rather short and compact (in
genus), lower surface almost straight, but
suddenly curved down in front of apex. Apex
short and wide, obtusely triangular, suddenly
turned down. Internal sac rather complexly
folded, with two narrow elongate spine-shaped
sclerotized plates within. On left side and at roof
of (inverted) internal sac with a strongly spinose
field each. Parameres very dissimilar, left
comparatively stout, with spine-like, thin apex
and small membranous area between basal part
and apex, with a single short apical seta. Right
paramere narrow and elongate, with tapering
apex, with 1 apical seta, c. 10 rather elongate
setae on lower margin, and 2-3 shorter setae on
apical third of upper margin.

Female genitalia (Fig. 2B). Stylomere 1 with 2-3
elongate ensiform setae at lateral part of
ventro-apical margin. Stylomere 2 rather short,
with short apex and 2 large dentiform
ventro-lateral ensiform setae of about similar size
below middle of lateral margin. Near apex with a
large, oblong pit and a short nematifoim seta
originating from that pit. In middle of
dorso-median surface with a large, dentiform,
dorso-median ensiform seta. Lateral plate with a
densely setose area at median apical margin.

Variation. A rather variable subspecies with
respect to relative shape of pronotum and elytra,
colour and distinctness of colour pattern, and
depth and degree of puncturation of elytra.
Especially the "specimens from Mt Spurgeon in
western part of Carbine Tableland tend to have
rather well impressed and coarsely punctate
elytra, though this is not regarded as more than a
local variation. All examined specimens from Mt
Spurgeon and Thornton Peak possess the

74

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 4. Habitus ofMecyclothorax. A, M. storeyi frerei subsp. nov.; B, M. lewisensis uncinatus subsp. nov.; C, M.
impressipennis sp. nov.; D, M. inflatus inflatus sp. nov. Lengths: 3.0mm; 2.95mm; 3.2mm; 3.05mm.

scutellar pore and seta, whereas these are
apparently absent in all other populations.

DISTRIBUTION (Fig. 6). Carbine Tableland
north of Mossman (Mt Lewis, Mt Demi,
Mossman Bluff, Devils Thumb, Mt Spurgeon),
and Thornton Peak and Mt Pieter Botte, both
north of Daintree River, North Queensland.

COLLECTING CIRCUMSTANCES. Most were
collected from rain forest leaf litter by Berlese
extraction, with a few from ground pitfall traps.
These are clearly ground dwellers invariably
occurring on the highest peaks of mountains and
tablelands and not descending below about
750m.

Mecvclothorax lewisensis uncinatus subsp. nov.
(Figs ID, 3C-D, 4B, 6)

ETYMOLOGY. The name refers to die sharply hooked
apex of acdeagus.

MATERIAL. HOLOTYPE: d, QMT2I587, AUST: NE
Qld, Mt Halcvon 24 Nov 1993 G Montcith & H. Janetzki,
QM BERLESATE No. 864 16°03’S 145°25'E Rf, 870m
Leaf litter (QM). PARATYPES: 2 <3,2 9, same data
(CBM, QM); 3 d, 4 9, Mt Hcmmant 27.xi.1993 GM &
HJ. QM Berl. No. 865 16°07'S I45°25’E Rf, 1050m
Sieved leaf'litter & moss (CBM, QM); 1 d , Roaring Meg
Ck 6km W Cape Tribulation 5.x. 1 982 GM, DY & GT, QM
Berl. No. 453 16.05S 145.24E Rf 710m Sieved litter
(QM); 1 d , 4.5-5km W of Cape Tribulation (Top Camp)

PSYDRINE GROUND BEETLES

75

1.x. 1982 GM, DY & GT, QM Bcrl. No. 442 16.05S
145.26E Rf, 760-780m Sieved litter (QM).

DIAGNOSIS. Distinguished from the nominate
subspecies by the hook-shaped apical part of
aedeagus bearing a sharply spined apex,
considerably shorter apex of male genital ring,
generally wider pronotum with narrower base,
and. on average, shorter elytra.

DESCRIPTION. Measurements . Length: 2.9-
3.1mm; width: 1.35- 1.45mm; Ratios. Length/
width of 9th antennomere: 1 .6-1.8; width/length
of pronotum: 1.34-1.41; width base/apex of
pronotum: 1.14-1.17; width pronotum/head:
1.45-1.54; length/width of elytra: 1.19-1.24;
width elytra/pronotum: 1.33-1.37.

Colour (Fig. 4B). As in nominate subspecies,
maculate pattern in all examined specimens
distinct.

Head. As in nominate subspecies with remark-
ably protruding eyes and small orbits.

Pronotum (Fig. 3C). As in nominate subspecies,
though at the average slightly wider in anterior
half and with relatively narrower base. Posterior
lateral setae always present.

Elytra (Fig. 3D). As in nominate subspecies,
though at the average slightly shorter. In all
examined specimens all striae, including sutural
stria, barely impressed and rather finely punctate.
Contrary to most populations of the nominate
subspecies, except for those occurring on Mt
Spurgeon and Thornton Peak, sutural pore and
seta always present.

Lower surface and legs. As in nominate subspecies.

Male genitalia (Fig. ID). Generally similar to
those of nominate subspecies, though apical part
of genital ring considerably shorter and apex of
aedeagus more curved downwards and with
acute spine at end.

Female genitalia. As in nominate subspecies.

Variation. Little variation noted, except for slight
differences in depth of puncturation of elytral
striae.

DISTRIBUTION (Fig. 6). Mt Hemmant, Mt
Halcyon and Roaring Meg Creek, north of
Thornton Peak, North Queensland.

COLLECTING CIRCUMSTANCES. All
specimens collected by sieving leaf litter and
moss in montane rainforest. Therefore, this
subspecies apparently lives in litter at the ground
and in moss near the bases of trees. Collections

were made between 710m and 1050m, mostly at
the tops of the respective mountains.

Mecyclothorax imprcssipcnnis sp. nov.
(Figs 3E-F, 4C, 5)

ETYMOLOGY. The name refers to the deeply impressed
four inner striae.

MATERIAL. HOLOTYPE: 9, QMT2142I, AUST: NE
Qld, Isley Hills I Dec 1 993 G Monteith & H. Janetzki, QM
Berlesate No 866 17°03’S 145°42’E Rainforest, 1050m
Sieved litter & moss (QM). PARATYPE: 1 9 , same data
(CBM).

DIAGNOSIS. Small species with cordate
prothorax that bears sharply angulate basal
angles, clypeus quadrisetose; distinguished from
all other species of this group by the very coarsely
punctate, well impressed four to five inner elytral
striae. Further distinguished from M. storeyi
Moore by much wider pronotum with wider base
that bears the posterior marginal setae; from M.
lewisensis Moore by wider, more quadrate elytra
with deeply punctate four inner striae and by
wider base of pronotum; and from M. in flatus sp.
nov. by narrower base of pronotum bearing
angulate basal angles.

DESCRIPTION. Measurements. Length: 3.2mm;
width: 1 .52- 1 .55mm; Ratios. Length/width of 9th
antennomere: 1.5; width/length of pronotum:
1.35-1.38; width base/apex of pronotum: 1.24-
1.27; width pronotum/head: 1.49-1.56; length/
width of elytra: 1.19-1.21; width elytra/
pronotum: 1.40-1.41.

Colour (Fig. 4C). Very dark piceous to almost
black, lateral margins of pronotum and four
inconspicuous spots on the elytra reddish.
Antennae, palpi and legs light reddish to dark
yellowish. Lower surface dark piceous, lateral
and terminal margins of abdomen reddish.

Head. Distinctly narrower than pronotum. Eyes
but moderately projecting, orbits rather large, c.
1/3 of length of eye. Eyes separated from Irons by
a narrow furrow. Frontal furrows elongate, deep,
evenly curved, almost attaining position ot
posterior supraorbital seta. Clypeal suture deep,
clypeus quadrisetose, punctures large and deep.
Labrum anteriorly straight, six-setose.
Mandibles of moderate size, seta in outer scrobe
elongate. Mentum with wide, obtuse, triangular
tooth. Two mental setae and four gular setae very
elongate. Glossa rather narrow, bisetose, para-
glossae membranous, by far surpassing glossa.
Lacinia elongate, sparsely spinose at inner
margin. Terminal palpomeres asetose. Antenna

76

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 5. Distribution of species of Mecyclothorax (part).

short, median antennomeres c.

1.5 X as long as wide. Posterior
supraorbital setae situated shortly
behind posterior margin of eye.

Upper surface of head absolutely
smooth, highly glossy.

Pronotum (Fig. 3E). Rather wide,
gently cordate. Apex almost
straight, apical angles barely
produced. Lateral margins evenly
curved, in front of base gently
sinuate, base straight. Basal
angles angulate, almost rect-
angular. Marginal sulcus
anteriorly narrow, widened
towards base. Both, apex and
base not margined. Median line
fine, not attaining apex nor base.

Basal grooves deep, linear.

Anterior transversal sulcus barely
indicated, basal transverse sulcus
shallow. Base with few scattered
punctures. Both marginal setae
present, the anterior one situated
slightly in front of middle, the
posterior one at basal angle.

Surface absolutely smooth,
highly glossy.

Elytra (Fig. 3F). Short and wide,
convex, rather quadrate. Humeri
wide, evenly rounded, lateral
margin anteriorly gently convex. Base
completely bordered. Scutellar stria interrupted,
scutellar pore and seta present, situated in 1st
interval. Four inner striae well impressed,
coarsely punctate, striae shortened towards base
and apex. Outer striae increasingly superficial,
finely punctate. Inner intervals considerably
convex. Dorsal puncture conspicuous, at inner
margin of 3rd interval, situated slightly in front of
middle. Marginal series consisting of 7-8 anterior
and 6 posterior setae that are widely separated in
middle. At end of 3rd and 5th intervals with a
short seta each. Intervals absolutely smooth,
highly glossy.

Low er surface. Metepistemum short, even wider
than long, terminal abdominal stemite in male
bisetose, in female quadrisetose.

Legs. Fairly elongate. Squamosity of male
anterior tarsus unknown.

Male genitalia. Unknown.

Female genitalia. As in M. 1. lewisensis.

Variation. Only some variation of depth of elytral
striae noted.

DISTRIBUTION (Fig. 5). Isley Hills, northwest
of Bellenden Ker Range. Known only from type
locality.

COLLECTING CIRCUMSTANCES. Collected
by pyrethrum knockdown on trees and logs in
upland rainforest.

Mecyclothorax inflatus sp. nov.

ETYMOLOGY. The name refers to the very wide elytra.

REMARKS. This new species is distributed
through the southern and southwestern parts of
Atherton Tableland. In the southernmost part of
its range (Walter Hill Range) a population exists
that differs by the armature of the internal sac of
the male aedeagus and is described as separate
subspecies. Some apparent geographic variation
also occurs in the nominate subspecies with
respect to external morphology (mainly in shape
of pronotum). However, at the present this is not
regarded as of major taxonomic value and the
different populations have not been attributed

PSYDRINE GROUND BEETLES

77

FIG 6. Distribution of species of Mecyclothorax (part).

subspecific rank (see also chapter ‘Variation’ in
the description of the nominate subspecies).

DIAGNOSIS. Small species with wide prothorax
that is barely sinuate posteriorly and bears rather
obtuse basal angles, clypeus quadrisetose.
Further distinguished from M storeyi Moore by
much wider pronotum with wider base that bears
the posterior marginal setae; from M. lewisensis
Moore by wider pronotum with wider base, and
by shorter genital ring; and from M impressi-
pennis sp. nov. by far less coarsely punctate
elytral striae.

Mecyclothorax inflatus inflatus sp. nov.
(Figs IE, 3H, 4D, 5)

MATERIAL. HOLOTYPE: cj, QMT21595, Mt Father
Clancy, 10km S Millaa Millaa NE Qld 4.V.1983 G
Montcith & D. K. Yeates, QM Berlesate No. 581 17.35S
145. 38E Rainforest, 840m Sieved litter (QM).
PARATYPES: 1 9 , same data (QM); 2 6 , 6 9 , Mt Father
Clancy, 9km S Millaa Millaa. 6.xii.l988 GM GT, QM
Berl. 812 145.33’E 17.35’S Rf. 1000m Sieved litter.
(CBM, QM); 2 6 , 2 9 , AUST: Qld: NE: Mt Hugh Nelson,
summit 7.ii.l999 GM & DC, QM Berl. 990 17°31’S

145°33’E Rf 1200m Sieved litter (QM);

1 d,3 9 , AUST: Qld: NEQMaalanRd,
1.5km S Palmerston Hwy 26.xi.1994.
GM, QM Berl. No. 879 17°36’S,
145.42’E Rf. 750m Sieved litter (CBM,
QM); 1 6, 2 9, AUST: Qld: NE Mt
Fisher, summit 8.ii. 1999 GM & DC,
QM Berl. 991 17°34’S 145°33’E Rf,
1360m Sieved litter (CBM, QM).

DIAGNOSIS. Distinguished
from southern subspecies, M.
inflatus spinifer subsp. nov., by
slightly shorter and wider elytra
and less spinose apex of internal
sac of aedeagus.

DESCRIPTION. Measurements.
Length: 2.8- 3.25mm; width:
1.35- 1.65mm; Ratios. Length/
width of 9th antennomere:
1.6-1.85; width/length of
pronotum: 1.38-1.48; width
base/apex of pronotum: 1.32-
1.36; width pronotum/head:
1.49-1.65; length/width of elytra:
1.16-1.19; width elytra/pronotum:
1.34-1.40.

Colour (Fig. 4D). Head and
pronotum dark piceous to almost
black, elytra piceous to
reddish-piceous. Lateral margins
of pronotum and four very
inconspicuous spots on the elytra reddish.
Antennae, palpi and legs light reddish to dark
yellowish. Lower surface dark piceous, lateral
and terminal margins of abdomen reddish.

Head. Distinctly narrower than pronotum. Eyes
but moderately projecting, orbits rather large, c.
1/3 of length of eye. Eyes separated from frons by
a narrow furrow. Frontal furrows elongate, deep,
evenly curved, almost attaining position of
posterior supraorbital seta. Clypeal suture deep,
clypeus quadrisetose, punctures large and deep.
Labrum anteriorly straight, six-setose. Mandibles
of moderate size, seta in outer scrobe elongate.
Mentum with wide, obtuse, triangular tooth. Two
mental setae and four gular setae present, very
elongate. Glossa rather narrow, bisetose, para-
glossae membranous, by far surpassing glossa.
Lacinia elongate, sparsely spinose at inner
margin. Terminal palpomeres asetose. Antenna
short, median antennomeres slightly >1.5 X as
long as wide. Posterior supraorbital setae situated
shortly behind posterior margin of eye. Upper
surface of head absolutely smooth, highly glossy.

78

MEMOIRS OF THE QUEENSLAND MUSEUM

Pronotum. Wide, barely or not cordate. Apex
almost straight, apical angles barely produced.
Lateral margins evenly curved, in front of base
either convex or almost straight, not sinuate, base
straight. Basal angles either angulate, though
wide, or almost obtuse. Marginal sulcus
anteriorly narrow, widened towards base. Both,
apex and base not margined. Median line fine, not
attaining apex nor base. Basal grooves deep,
linear. Anterior transversal sulcus barely
indicated, basal transverse sulcus shallow. Base
with few scattered punctures. Both marginal
setae present, the anterior one situated slightly in
front of middle, the posterior one at basal angle.
Surface absolutely smooth, highly glossy.

Elytra (Fig. 3H). Short and wide, convex, gently
oviform. Humeri wide, evenly rounded, lateral
margin convex throughout. Base completely
bordered. Scutellar stria interrupted or almost
wanting, if present, situated in 1st interval,
scutellar pore and setae wanting. Inner striae
including sutural stria indicated as rows of fine
punctures, or even wanting, not at all impressed,
outer striae not perceptible. Intervals absolutely
depressed. Dorsal puncture inconspicuous,
situated slightly in front of middle, at inner
margin of 3rd interval when this is present.
Marginal series consisting of 7-8 anterior and 6
posterior setae that are widely separated in
middle. At end of 3rd and 5th intervals with a
short seta each. Intervals absolutely smooth,
highly glossy.

Legs. Fairly elongate. Male anterior tarsus
slightly widened, asymmetrically squamose on
1st - 3rd tarsomeres.

Male genitalia (Fig. 1 E). Genital ring moderately
elongate, asymmetrically triangular, apex
moderately wide, fairly elongate. Aedeagus
normal, turned to the left side (in beetle), rather
short and compact (in genus), lower surface
almost straight, but suddenly curved down in
front of apex. Apex short and wide, suddenly
curved down and to the rear, with acute spine at
end. Internal sac rather complexly folded, with a
narrow elongate spine-shaped sclerotized plate
within. On left side of bottom and on right side at
roof of (inverted) internal sac with a rather small
spinose field each. Parameres fairly dissimilar,
left comparatively stout, with rather short,
tapering, spine-like apex, with a single short
apical seta. Right paramere narrow and elongate,
with tapering apex, with 2 apical setae, 8-10
rather elongate setae along lower margin, and 2-5
shorter setae on apical third of upper margin.

Female genitalia. As in M. /. lewisensis.
Variation. Some geographical variation ot
relative shape of pronotum can be noted between
the populations recorded from Mt F isher and Mt
Hugh Nelson (both near Millaa Millaa) and those
from Mt Father Clancy and Malaan Rd near
Palmerston Hwy, further southeast. I he
examined specimens from Mt Fisher and Mt
Hugh Nelson possess large pronota with the
lateral margins evenly curved towards the obtuse
basal angles, whereas in the available specimens
from Mt Father Clancy and Malaan Road the
pronotum is less voluminous and has the lateral
margin little convex to almost straight near the
more angulate basal angles. Further collecting in
the area between may clarify the taxonomic
situation. As male genitalia do not show any
striking differences, for the present these
populations are regarded as infrasubspecific
units of no taxonomic value. Probably, they are
members of a morphological cline. However, for
better comparison and possible future
distinction, the measurements and ratios of both
populations are added below:

Mt Fisher/Mt Hugh Nelson (N = 5): Length:
3.0-3.25mm; width: 1 .52-1. 65mm; Ratios. Length/
width of 9th antennomere: 1.7-1.85; width/length
of pronotum: 1.45-1.48; width base/apex of
pronotum: 1. 29- 1.34; width pronotum/head: 1.59-
1.65; length/width of elytra: 1.16-1.17; width
elytra/pronotum: 1 .34- 1 .36.

Mt Father Clancy/Malaan Rd (N = 5): Length:
2.8-3. 1mm; width: 1.35- 1.52mm; Ratios. Length/
width of 9th antennomere: 1 .6-1.7; width/length
of pronotum: 1.37-1.44; width base/apex of
pronotum: 1.32-1.36; width pronotum/head: 1.49-
1.58; length/width of elytra: 1.18-1.19; width
elytra/pronotum: 1.35-1.40.

DISTRIBUTION (Fig. 6). Southwestern part of
Atherton Tableland: Mt Fisher, Mt Hugh Nelson,
Mt Father Clancy, and Malaan Road south of
Palmerston Hwy., North Queensland.

COLLECTING CIRCUMSTANCES. Collected
by berlese extraction from rainforest leaf litter.

Mecyclothorax inflatus spinifer sp. nov.

(Figs IF, 3G, 5)

ETYMOLOGY. The name refers to the remarkably laige
spinose fields at the entrance of the internal sac of the male
aedeagus.

MATERIAL. HOLOTYPE: <5, QMT21599, NE Qld,
Upper Boulder Ck via Tully 27.x. 1983 Monteith, Yeates &
Thompson, QM Berlesate No. 600, 17.50S 145.54E Rf,

PSYDRINE GROUND BEETLES

79

TABLE I . Mecyclothorax spp. measurements.

Species

N

body length
(mm)

ratio length/
width 9th
antennomere

ratio width/
length
pronotum

ratio width
base/apex
pronotum

ratio width
pronotum/
head

ratio length/
width elytra

ratio width
elytra/

pronotum p

M. storeyi storeyi

10

2.8-3.25

1. 8-2.0

1.24-1.30

1.10-1.15

1.51-1.60

1.24-1.27

1.36-1.38

M. storeyi frerei

10

2.65-3.15

1.9-2. 1

1.29-1.32

1.07-1.10

1.43-1.50

1.25-1.29

1.33-1.34 |

M. lewisensis lewisensis

10

2.8-3.25

1.65-1.8

1.32-1.36

1.21-1.25

1.48-1.54

1.22-1.27

1.37-1.42

A/. lewisensis uncinatus

10

2.9-3. 1

1.65-1.8

1.34-1.41

1.14-1.17

1.45-1.54

1.19-1.24

1.33-1.37

M. impressipennis

2

3.2

1.5

1.35-1.38

1.24-1.27

1. 49-1. 56

1.19-1.21

1.40-1.41

M. injlatus injlatus

10

2.8-3.25

1.6-1.85

1.38-1.48

1.32-1.36

1.49-1.65

1.16-1.19

1.34-1.40

M. inflatus spinifer

10

2.8-3.05

1.7-1. 8

1.36-1.40

1.30-1.37

1.53-1.59

1.20-1.23

1.31-1.36 1

900m Sieved litter (QM). PARATYPES: 3 <5, 5 9, same
data (CBM, QM); 1 9, same loc., 26.x. 1983 GM, DY &
GT Pyr. in RF. (QM).

DIAGNOSIS. Distinguished from the northern
nominate subspecies by slightly longer and
narrower elytra, narrower pronotum at the
average, and remarkably spinose apex of internal
sac of aedeagus.

DESCRIPTION. Measurements. Length: 2.8-
3.05mm; width: 1.35- 1.48mm; Ratios. Length/
width of 9th antennomere: 1.7- 1.8; width/length
of pronotum: 1.36-1.40; width base/apex of
pronotum: 1.30-1.37; width pronotum/head:
1.53-1.59; length/width of elytra: 1.20-1.23;
width elytra/pronotum: 1.31-1.36.

Colour. As in nominate subspecies, though all
examined specimens with rather light coloured
elytra and extremely faded pattern.

Head. As in nominate subspecies.

Pronotum (Fig. 3G). As in nominate subspecies,
though pronotum generally even narrower and
lateral margins posteriorly even less convex than
in the Mt Father Clancy/Malaan Rd population.
Therefore, basal angles ungulate and distinct.

Elytra. As in nominate subspecies, though
slightly longer and narrower. Scutellar pore and
seta lacking. Striation extremely inconspicuous,
in some specimens barely recognisable.

Lower surface. As in nominate subspecies.

Legs. As in nominate subspecies.

Male genitalia (Fig. IF). Very similar to those of
nominate subspecies, though internal sac at end
with two large, remarkably spinose fields, both
situated at the right side.

Female genitalia. As in nominate subspecies.
Variation. Very little variation noted.

DISTRIBUTION (Fig. 6). Southernmost
Atherton Tableland: Upper Boulder Creek area in

Walter Hill Range. Known only from type
locality.

COLLECTING CIRCUMSTANCES. Collected
by sieving litter and by pyrethrum knockdown in
upland rainforest on trees and logs. Probably this
subspecies generally lives on the ground, but also
in moss at the bases of rainforest trees. So far
collected at 900m.

MEASUREMENTS AND RATIOS IN
MECYCLOTHORAX SHARP

For better comparison of the species the
measurements and ratios of all species and
subspecies are compiled in Table 1.

REMARKS. The diversity of Mecyclothorax in
the Wet Tropics of North Queensland is shown to
be greater than previously indicated. Certainly,
the various taxa of the storeyi- group are
thoroughly distinct from those species occurring
in the southern half of Australia (the ambiguus-
group), and in part, they are very closely related
inter se. In view of shape of aedeagus and
complex armature of the internal sac in all
northern taxa, these probably form a rather apotypic
group within the Australian Mecyclothorax , but
at the same time in external and genitalic
morphology they show a certain grade of
similarity with the species occurring in New
Guinea (Baehr, 1995a, 1998, 2002c) and New
Caledonia (Deuve, 1987, Baehr, pers. obs.). So, it
would be conceivable that the New Guinean and
New Caledonian Mecyclothorax (and perhaps
also those occurring further north in Java and
Borneo) should have been derived from ancestors
that were related to the storeyi- stock and were
coming originally from northern Queensland.

Apart from M. storeyi which is unique for its
strange, side-inverted aedeagus and generally
lack of posterior marginal prothoracic seta, the
other three species are still very closely related.

80

MEMOIRS OF THE QUEENSLAND MUSEUM

This is demonstrated by the morphology of the
aedeagus, the duplication of the clypeal seta, and
the shape and structure of pronotum. Even the
most distant taxa, M. lewisensis and M. inflatus ,
separated by about 100km, are still very closely
related.

As with the situation in New Guinea, where in
those areas that have been more extensively
sampled for litter inhabiting Mecyclothorax ,
many species with rather restricted ranges exist
(Baehr, 1995a, 2002c), the Wet Tropics of
northern Queensland also harbours a number of
taxa — most still closely related — in a restricted
region. In several instances, the ranges of
different taxa are spatially close. Certainly this
rapid turnover of ranges is due to the low vagility
of these tiny, flightless, litter inhabiting, montane
beetles. Since none have been taken below 650m
in North Queensland, and most are from above
1,000m, even rather unimportant creek valleys
may act as significant distribution barriers.

A striking example for this range fragment-
ation is the distribution of populations of M.
lewisensis in the Thornton Peak area. The more
northern form (M. lewisensis uncinatus ) occurs
on a mountain block that is no more than 5km
distant from Thornton Peak where the nominate
form occurs, and that is only separated by the
valleys of two creeks that do not descend below
500m. However, these unimportant valleys
apparently are sufficient to act as significant
barriers for montane, rain forest living beetles. In
the same region, this barrier is corroborated by
the occurrence of two different species of blind,
litter-inhabiting water beetles of the genus
Terradessus (Dytiscidae), one on each side of the
valley (Brancucci & Monteith, 1996). In
Mecyclothorax the situation is even more com-
plex. Within the range of the northern population
(A/. lewisensis uncinatus ), namely near Mt Pieter
Botte, the southern nominate population
apparently appears again as demonstrated by the
rounded apex of the aedeagus and the very
elongate genital ring in males collected on Mt
Pieter Botte. This population again occurs in the
immediate neighbourhood of the uncinate form
on mountains near Cape Tribulation, also
separated by only a minor stream valley.

So far, in the North Queensland Mecyclothorax ,
no overlapping of ranges has been detected,
contrary to other genera (e.g. in Sitaphe). This
may be evidence of a rather recent diversification
of this group following recent immigration into
the northern montane rain forests. The high level

of phylogenetic diversification ol the species, on
the other hand, would suggest a longer history of
Mecyclothorax in this area. Additional
knowledge about distribution may bring more
light to this question.

Teraphis Castelnau, 1 867

Teraphis Castelnau. 1867: 41; Castelnau, 1868: 127; Sloane,
1898: 470; Csiki. 1929: 485; Moore. 1963: 283; Moore et

Phersila Sloane, 1903: 591; Sloane, 1920. 156, Csiki, 1929:
485: Moore, 1963: 283; Moore etal., 1987: 151.

REMARKS. A genus of Tropopterini. Most
species occur in southeastern Australia from
southern New South Wales to Tasmania with a
single species reaching the Macpherson Range in
southern Queensland. Sloane’s replacement
name Phersita was unjustified, as Moore (1987)
stated.

Teraphis helmsi (Sloane)

Drimostoma helmsi Sloane, 1890: 647.

Teraphis helmsi, Sloane, 1 898: 47 1 ; Csiki, 1 929: 486; Moore,
1963: 284; Moore etal., 1987: 151.

DISTRIBUTION. The single northern species of
this decidedly southern genus occurs in
northeastern New South Wales (type locality:
Dunoon, near Lismore) and at the following
localities (QM specimens) along the Macpherson
Range within Queensland (east to west): Upper
Tallebudgera Valley; Numinbah Arch; Mt
Asplenium; Mt Huntley. It is curious that it has
never been taken on the well-collected
Lamington Plateau, even though it occurs to the
east and west of that area.

Trephisa Moore, 1963

Trephisa Moore, 1963: 282; Moore et al., 1987: 152.

REMARKS. A genus of Tropopterini, described
by Moore for a unique, small-eyed, elongate
species.

Trephisa parallcla Moore

Trephisa parallela Moore, 1963: 282; Moore et al., 1987:
152.

REMARKS. At the time of Moore et al. (1987)
the unique species of the genus was still known
only from the type locality at Binna Burra on the
Lamington Plateau. An additional specimen has
been collected recently by G. Monteith at
Springbrook Plateau about 10km SE of Binna
Burra, which belongs to the same tableland
system. The species is apparently endogeous, as
all five recorded specimens were collected under
deeply embedded rocks.

PSYDRINE GROUND BEETLES

81

DISTRIBUTION: Lamington and Springbrook
Plateaus, southeastern Queensland.

Sitaphe Moore, 1963

Sitaphe Moore, 1963: 284; Moore et at, 1987: 150.

TYPE SPECIES. Sitaphe rotundata Moore, 1963, by
monotypy.

DIAGNOSIS. Easily distinguished from all other
Australian Tropopterini by the short, oval-shaped
form, rather trapezoidal pronotum that is widest
at or near the basal angles, and oviform, scarcely
striate elytra. Other diagnostic characters may be
taken from Moore's (1963) description of the
genus.

Pronotum shorter and wider, anteriorly less trapezoidal
(Fig. 8E); right paramere angulate near base, rather
straight in apical half (Fig. 7F). Upper Boulder Creek
area in Walter Hill Range incunncollis sp. nov.

6. Smaller species, length 4. 1 -4.7mm and elytra short and

wide (ratio length/width < 1.15) and aedeagus and
parameres short (Fig. 3 A). Northeastern part of Atherton
Tbld: Lambs Head, Mt Williams, Isley Hills
minuta sp. nov.

Either larger species, length 4.9-5.6mm and aedeagus and
parameres elongate (Fig. 7G); or length 4.6-5. 3mm and
elytra elongate ( ratio length/width >1.18) 7

7. Elytra shorter, laterally more parallel, less oviform (Fig.

81); aedeagus and parameres elongate (Fig. 7G). Carbine

and Windsor Tblds parallelipennis sp. nov.

Elytra longer, laterally more convex, oviform (Fig. 8L);
aedeagus and parameres much shorter (Fig. 7H).
Thornton Peak ovipennis sp.nov.

REMARKS. Highly apotypic, known only from
the Wet Tropics. All taxa of this genus live in
crevices of logs and dead trees in tropical upland
rain forest (G.B.Monteith, pers.comm.), and
extend to the summits of the highest peaks in the
region.

KEY TO THE TAXA OF THE GENUS
SITAPHE MOORE

l.

2 .

3.

4.

5 .

Pronotum markedly trapezoidal, at basal angles not at all
incurved (Figs 8D, 9B). Bellenden Ker/Bartle Frere

Ranges trapezicollis sp. nov.

Pronotum less trapezoidal, at base at least slightly

incurved (Figs 8A,B,E,G,H,J,K, 9A, 9C) 2

Basal angle of pronotum obtuse, without distinct denticle,
lateral margin barely incurved, pronotum veiy wide
(ratio width/length >1.60), with wide base (ratio width
base/apex >2.0) (Figs 8A, 9A). Eastern and southern
fringes of Atherton Tbld: Bellenden Ker/Bartle
Frere/Massey Ranges, Malbon Thompson Range, Mt
Hypipamee, Mt Fisher, Millaa Millaa Falls, Mt
Kooroomool rotundata Moore

Basal angle of pronotum with small denticle and more or
less deep incurvation in front (Figs 8E,G.H,J,K), or basal
angle rectangular and lateral margin in front straight or
even slightly sinuate (Figs 8B, 9C): pronotum less wide
(ratio width/length <1.60), with narrower base (ratio

width base/apex <2.0) 3

Basal angle of pronotum rectangular and lateral margin in
front straight or even slightly sinuate (Figs SB, 9C),
surface of pronotum rather depressed, base in middle
remarkably impressed. Bartlc Frere Range
parvicollis sp.nov.

Basal angle of pronotum with small denticle and more or
less deep incurvation in front (Figs 8E,GH,J,K), surface
of pronotum rather convex, base in middle less deeply
impressed 4

Lateral margin of pronotum near base suddenly incurved,
basal angle conspicuously dentiform (Figs 8E,J). ... 5
Lateral margin of pronotum near base but slightly
incurved, basal angle faintly denticulate (Figs 8G,H,K).

6

Pronotum longer and narrower, anteriorly more regularly
trapezoidal (Fig. 8J); right paramere less angulate near
base, more curved in apical half (Fig. 7E). Cardwell
Range hamifera sp.nov.

Sitaphe rotundata Moore
(Figs 2C, 7A, 8 A, 9A,D, 11)

Sitaphe rotundata Moore, 1963: 284; Moore et al. 1987: 150.

MATERIAL. HOLOTYPE: 6 , Mt Bartlc Frere W. slope,
Q. 3-5000’, Dec. 57’ Darlingtons/ Sitaphe gen. nov.
rotundata sp. nov. holotype d Det. B.P. Moore 63
(ANIC). PARATYPE: 1 9, same data;' Sitaphe gen. nov.
wtundata sp. nov. paratype 9 Det. B.P. Moore ‘63
(ANIC).

NEW RECORDS. 1 9, Bellenden Ker, 1994 Crash Site.
17°16'S 145°5rE 1 .xii.1998 GM. Pyr. trees. 1325m
(OM):2 9, Bellenden Ker Range, NQ Summit TV Stn.,
1560m Oct 25-3 L 1981 Earth watch/QM, QM Bcrl. 372
17 16S 1 45.5 1 E Rf Sieved litter (QM); 1 d, 1 9,sameloc.,
29 Aor-3 May, 1983 GM, DY (QM); 1 9, Bellenden Ker,
Centre Peak Summit, 1 0.iv. 1 979 GM, QM Bed. N 12
17.16A 145.5 IE Rf, 1500m Stick brushings (QM); 4 9,
Bellenden Ker Range, Cable Tower 3, 1054m Oct 17-24
1981 Earthwatch/QM (QM); 4 d, same loc.,17 Oct-5
Nov. 1981 Earthwatch/QM (CBM, QM); 3 d, Mt Bartle
Frere NW/Centre Peak 16 Sept. 1982, 1500m GM & SM
(OMV 3 d, same loc., 24.ix.1981 GM & DC (QM); 2 * ,
same loc., 7-8.xi.1981, 1400-1500m Earthwatch/Qld
Museum (QM); 1 9, Mt Bartle Frere O.^km N ! of Stlt
Peak 6-8 Nov. 1 98 1 , 1 500m Earthwatch/QM Pyr. (QM). 3
d . 2 9 , Mt Bartle Frere, N Qld. Sth. Peak Summit 1 620m
6-8 Nov., 1981 Earthwatch/QM (QM); 1 9 , same loc., 6-8
Nov ,1981 Earthwatch/QM Pyr. (CBM); 4 d , 3 9 i Same
loc 29.xi.1998. GM. DC’, PB. (CBM, QM); 3 d, Mt
Bartle Frere west face, 1 000- 1400m 7.x. 1980 GB & SM
(OMV 1 d. Mt Bartle Frere, N Qld. Central Ridge. 1500m
27.xii.1989 GM & SM (QM); 7 d, 3 9, NE. Q: 17°16’S
1 45°49’E Massey Range, 4km W ot Centre Bellenden Ker
9-11.X.1991- 1250m GM, HJ&DC(CBM.QM);3 ?,NE.

0 17°14’S 145°48’E Massey Range. 6km NW of Centre
Bellenden Ker 1 1 - 1 2.x. 1 99 1 . 1 1 50m GM, H.I & DC (QM);
2 6 North Bell Peak, 1000m Malbon Thompson Ra„ N
Old. 1 9-22.xi. 1 990, 800-900m GM & GT (QM);2 6 , same
loc 900- 1000m, 15-16.ix.1981 GM & DC (QM);1 2,
same loc 130ct 1982,850-1000mGM,DY&GT(QM);

1 6 1 2, South Bell Peak Malbon Thompson Ra.,N Qld.
20-2 l.xi.l 990, 900m GM & GT Pitfall Traps (QM); 2 6,2

82

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 7. Male genitalia of Sitaphe spp. A, S. rotundata Moore, aedeagus, parameres, and genital ring; B, S.
trapezicollis sp. nov., aedeagus and parameres; C, S. parvicollis sp. nov., aedeagus and parameres; D, S. minuta
sp. nov., aedeagus and parameres; E, S. hamifera sp. nov., aedeagus and parameres; F, S. incurvicollis sp. nov.,
aedeagus and parameres; Q S. parallelipennis sp. nov., aedeagus and parameres. H, S. ovipennis sp. nov.'
aedeagus and parameres. Scales: 0.5mm.

2, Mt Hypipamee Nat. Park 5.x. 1980 GM, QM Berl. 237
Rf, 950m Stick brushing (CBM, QM); 1 d, same loc.,
960m’ 24 July 1982 S. & J. Peck, SBP86, Rf streamside
litter ( ANIC); 1 d , Tower S. of Crater N P 1 6.v. 1 995 GM,
QM Berl. No. 886 17°27’S, 145°29’E Rf, 1230m Stick
brushing (QM); 1 d, Millaa Millaa Falls, via Millaa
Millaa, "N Qld. 12 Aug. 1968. R. Cantrell / Sitaphe
rotundata Mre Del. B.P. Moore’ 69 (QM); 1 d, 1 2, Mt
Fisher, 7km SW Millaa Millaa NQ (Whiteing Road)
5 .v. 1983, 1200m GM, DY (QM); 3 d, 2 2, Mt Fisher,
1050-1 1 00m 7km SW Millaa Millaa, NQ. 27-29 Apr.,
1982 GM, DY & EXT (CBM, QM); 1 d,Mt Fisher, l/2km
NW 17°33'S 1 45°33 ’E 8.ii. 1 999. GM 1280m. Pyr. -trees &
logs, Rf. (QM); 1 d, Mt Fisher, summit. 1360m 17°33’S
145°33’E 8.ii.l999. RfGM. Pyr.-trees & logs. (QM); 1 2,
Mt Kooroomool, summit. 7km S. 1 7°54 , S 145°4L’E
3-4.xii.1998 GM DC PB 1050m, Rf. (QM).

DIAGNOSIS. Distinguished by the very wide
pronotum with wide base. Further distinguished

from syntopic S. trapezicollis sp. nov. by basally
incurv ed lateral margins of pronotum and by the
remarkably curved right paramere. Distin-
guished from species with similarly shaped
pronota by having basal angles not sinuate and
not at all angulate.

DESCRIPTION. Measurements. Length: 4.6-
5.8mm: width: 2.40-3.25mm; Ratios. Length/
width of 9th antennomere: 2.2-2. 5; width/length
of pronotum: 1.60-1.67; width base/apex of
pronotum: 1.98-2.07; width pronotum/head:
2.02-2.12; length/width of elytra: 1.12-1.15;
width elytra/pronotum: 1 .25- 1 .30.

Colour. Overall black as in all other species.
Head. Very similar to that of other species.

Pronotum (Figs 8A, 9A). Very wide, generally
trapezoidal, though lateral margins evenly

PSYDRINE GROUND BEETLES

83

convex throughout, slightly incurved to basal
angles, therefore, widest diameter at a short
distance in front of basal angles. However, lateral
margins not excised at basal angles which are
obtuse, neither angulate, nor dentiform.

Elytra. Of average relative length, in middle
rather parallel-sided though still gently convex,
elytra not markedly egg-shaped. Usually two
internal striae slightly depressed and rather finely
punctate, both inner intervals in basal half gently
convex. Outer striae decreasingly impressed or
even almost absent, barely punctate.

Legs and low er surface. As in the other species.

Male genitalia (Fig. 7A). Genital ring
moderately elongate, asymmetrically triangular,
apex moderately wide, rather short. Aedeagus,
comparatively short (in genus), lower surface
gently curved. Apex very short, barely
surpassing (inverted) internal sac. Orificium
turned to right side. Internal sac moderately
folded, with two narrow, sclerotized plates
within, the upper one more spine-like, the lower
one wider and near apex slightly spinose.
Parameres elongate, fairly dissimilar, left
comparatively stout, rather convex, with
suddenly tapering, rather elongate apex, asetose.
Right paramere very narrow and elongate, deeply
sinuate on lower side, with tapering apex,
asetose, though with a series of minute hairs at
lower margin near apex.

Female genitalia (Fig. 2C). Stylomere 1 asetose
at apical rim. Stylomere 2 rather short, with short
apex and 2 large dentiform ventro-lateral
ensiform setae of about similar size below middle
of lateral margin. Near apex with a large, oblong
pit and a nematiform seta originating from that
pit. In middle of dorso-median surface with a
large, dentiform, dorso-median ensiform seta.
Lateral plate with a densely setose area at median
apical margin.

give no clear information, this species probably
lives rather on the bark or in moss of logs and
trunks than in the forest litter.

Sitaphe trapezicollis sp. nov.

(Figs 7B, 8D, 9B,E, 12)

ETYMOLOGY. The name refers to the remarkably
trapezoidal shape of pronotum.

MATERIAL. HOLOTYPE: d, QMT21669, Bcllenden
Ker Range, NQ Summit TV Stn.. 1 560m Oct. 25-3 1 , 1981
Eartliwatch/Qld Museum (QM). PARATYPES: 3 d, 4 9,
same data (QM); 7 d , 6 9 , same loc., 1 7 Oct.-Nov. 5, 1 98 1
17.16S 145.5 IE Earthwateh/QM Pyr. (CBM, QM); 1 d, 1
9 , same loc.Oct. 17-24. 1981 Earthwatch/QM (QM); 1 9,
same loc., 30 Nov-2.xii.1998 HAND GM, PB& DC,
1500m 1991 (QM); 1 d, l 9, same loc., Nov. 1-7, 1981
Earthwatch/QM, QM Berl. 342 Rf Stick brushings (QM);
1 d , same loc.. 28.x. 1 983 GM, DY & GT Pvt. in RF (QM);
5 d , 4 9 , same loc., 29 Apr.-3 May, 1 983 GM, DY (CBM,
QM); 1 9, same loc., 28.x. 1983, GM. DY & G1 (QM); 4
d 2 9 , same loc., 28.x. 1983 GM DY & GT Pyrethrum.
Rf. (QM); 1 d, same loc., 8.x. 1991. 1560m GM & HJ
Pyrethrum, trees & logs (QM); 4 d, 2 9, same loc.,
8.x. 1991 . 1 560m GM, HJ & DC (QM); 1 9 , Mt Bellenden
Ker Centre Peak Summit NE Qld 10-1 2.iv. 1979 1500m
GM (QM); 2 d. NEQ: 17°16 T S 145°5PE Bellenden Ker
summit 30 Nov-2.xii. 1998 HAND GM, PB & DC, 1500m
1991 (QM);7 d, 5 9, Mt Bartlc Frere, N Qld. Sth. Peak
Summit, 1620m 6-8 Nov., 1981 Earthwatch/QM Pyr.
(CBM. QM);5 d, 4 9 , same loc,, 29.xi.1998. GM.
Pyrethrum, trees/logs. 1 620m (CBM, QM).5 d , 3 9 .same
Idc., 29.xi. 1 998, GM, DC &PB. (CBM, QM); 2 d , 2 9 , Mt
Bartle Frere, NW/Centre Peak ridge, 7-8 xi. 1981,
1 400- 1 500m Earthwatch/Qld Mus. (QM ); 4 d , 2 9 , same
loc 16 Sept. 1982, 1500m GM & SM (QM); 2 d, Mt
Bartle Frere, NQ 0.5km N of Sth. Peak, 6-8 Nov. 1981,
1500m Earthwatch/QM Pyr. (QM); 1 d, 1 9, 17°24'S
145°49’E Bartle Frere, Top camp 1500m, 29.xi.1998 GM,
Pyr. trees, R.F.(QM).

DIAGNOSIS. Easily distinguished by the strongly
trapezoid prothorax that is not at all incurved at
basal angle.

DISTRIBUTION (Fig. 10). Almost the whole of
Atherton Tableland and surroundings, North
Queensland: Bellenden Ker and Bartle Frere
Ranges, Massey Range, Malbon Thompson
Range, Mt Hypipamee, Millaa Millaa Falls, Mt
Fisher, Mt Kooroomool.

COLLECTING CIRCUMSTANCES. All
specimens collected in montane rainforest,
generally above 1050 m, with many specimens
caught at the summits of the ranges. Mostly
sampled by pyrethrum knockdown, but also by
‘stick brushing’, ‘pitfall trapping’ and ‘on tree’.
Although collecting circumstances as labelled

DESCRIPTION. Measurements. Length: 4.9-
6.2mm; width: 2.70-3.45mm; Ratios. Length/
width of 9th antennomere: 2.25-2.5; width/length
of pronotum: 1.49-1.53; width base/apex of
pronotum: 2.02-2.07; width pronotum/head:
2.03-2.11; length/width of elytra: 1.12-1.13;
width elytra/pronotum: 1.29-1.34.

Colour. Black, elytra in some specimens very
dark piceous. Labrum and mandibles dark
reddish, palpi and antennae reddish. Femora and
tarsi reddish, tibiae reddish-piceous. Lower
surface of anterior body dark piceous, abdomen
reddish-piceous.

84

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 8. Pronota and elytra of Sitaphe spp. A, S.
rotundata Moore; B. C, S. parvicollis sp. nov.; D, S.
trapezicollis sp. nov.; E, F, S. incurvicollis sp. nov.; G
S. minuta sp. nov.; H, I, S. parallelipennis sp. nov.; J,
S. hamifera sp. nov.; K, L, S. ovipennis sp. nov.

Head. Half as wide as pronotum. Eyes of
moderate size, little protruding, posteriorly
enclosed by the orbits that are about 1/3 of length
of eyes. Eyes separated from frons by a narrow,
straight furrow. Frons with two elongate, curved
furrows medially of eyes that are prolonged to

clypeal suture. Neck separated from frons by a
shallow transverse furrow. Anterior supraorbital
seta situated close to eye slightly in front of
middle of eye, posterior seta situated at or just
behind posterior margin of eye and slightly
moved on upper part of head. Clypeal suture
deeply impressed. Labrum anteriorly slightly
concave, 6-setose and with some shorter hairs
around the anterior angles. Mandibles elongate,
though of median size in genus, straight, inner
margin straight for a long distance, then gently
incurved, with acute apex, with elongate seta in
scrobe. Right mandible with conspicuously large
tooth in middle of inferior margin. Mentum with
apical ly rounded triangular tooth. Submentum
bisetose. Gula quadrisetose. Glossa short,
narrow, bisetose, paraglossae hyaline, surpassing
glossa. Lacinia elongate, inner margin with few
strong spines, apex markedly incurved, very
acute. Both palpi glabrous, maxillary palpus with
elongate, fusiform terminal palpomere, labial
palpus shorter and stouter, apex transverse.
Antennae rather short and stout (in genus), just
attaining base of pronotum, pilose from half of
4th antennomere, central antennomeres <2.5 x
as long as wide. Surface of head absolutely
smooth, without any indication of
microsculpture or puncturation, remarkably
glossy.

Pronotum (Figs 8D, 9B). Remarkably trapezi-
form. Wide, though comparatively narrow in
genus. Apex slightly concave, anterior angles
slightly produced, obtusely rounded, sides
evenly though comparatively little curved, not
incurved towards base, therefore widest
immediately at basal angles. Base in lateral third
remarkably oblique, exactly adapted to the
oblique base of elytra. Basal angles angulate, not
dentiform, angle c. 100°. Lateral borders
coarsely margined, apex and base more Finely
margined. Lateral channel barely indicated.
Median line distinct, though shallow, neither
reaching apex nor base. Basal grooves very
shallow, rather punctiform, situated close to
middle. Both, anterior and posterior transverse
sulci absent. Base in middle not impressed.
Anterior lateral seta situated shortly behind
middle, posterior lateral seta situated slightly
inside of basal angle. Surface absolutely smooth,
without any indication of microsculpture and
puncturation, remarkably glossy.

Elytra. Wide, convex, considerably wider than
pronotum, wide at humeri, reversely oviform.
Humeri angulate, basal border deeply excised.
Lateral margins convex throughout, elytra widest

PSYDRINE GROUND BEETLES

85

at anterior fourth, than evenly narrowed. In
anterior half lateral margin faintly convex, barely
sinuate at position of crossing of epipleurae.
Basal margin complete, lateral margin narrow
throughout. Scutellar puncture and seta present,
at base of outtumed 1st stria. Sutural stria short,
inside 1st stria. Two inner striae at least in
anterior half impressed, finely punctate or even
gently crenulate. External striae decreasingly
less distinct, barely punctate, outer striae barely
recognisable. 8th stria only in apical half present.
Near apex 1st, 2nd, and in particular 7th striae
well impressed, 7th stria forming an elongate,
fairly deep furrow. At most 1st and 2nd intervals
feebly convex in basal half, outer intervals
depressed or not recognisable. Disk impunctate.
Marginal series consisting of two groups of 7-8
and 6 setiferous punctures, respectively, which
are rather widely separated in middle. Inside of
deepened 7th stria with two additional punctures
very near to apex. Some of the marginal setae
very elongate. Intervals absolutely smooth,
without any traces of microreticulation, highly
glossy.

Lower surface. Elytral epipleurae anteriorly very
wide. Metepistemum short and small, slightly
longer than wide at apex. Lower surface
impunctate. Terminal stemite in male bisetose, in
female quadrisetose along margin, and with two
shorter setae in middle somewhat removed from
margin.

Legs. Fairly elongate. 5th tarsomere with one pair
of very elongate setae beneath. Anterior tarsus in
male barely wider than in female, 1st - 3rd
tarsomeres very lightly squamose beneath.
Claws large, smooth.

Male genitalia (Fig. 7B). Genital ring as in S.
rotundata. Aedeagus as in S. rotundata , though
even shorter and slightly more curved. Left
paramere basally narrower than in S. rotundata ,
with evenly tapering apex. Right paramere less
elongate than in S. rotundata , for less sinuate on
lower side.

Female genitalia. Very similar to those of S.
rotundata.

Variation. Some variation noted in size, relative
shape of pronotum, and depth of elytral striae.

DISTRIBUTION (Fig. 11). Bellenden Ker and
Bartle Frere Ranges at the eastern margin of
Atherton Tableland.

COLLECTING CIRCUMSTANCES. Mostly
collected by pyrethrum knockdown in upland
rainforest, some also by hand collecting.

Generally they have been captured not below
1500m, with most specimens collected on
summit peaks and ridges. This species probably
lives rather on the bark or in moss of logs and
trunks than in the forest litter.

Sitaphe parvicollis sp. nov.

(Figs 7C, 8B-C, 9C,F, 12)

ETYMOLOGY. The name refers to the unusually narrow
and small pronotum.

MATERIAL. HOLOTYPE: £, QMT21688, Mt Bartle
Frere, N Qld. Sth. Peak Summit, 1620m 6-8 Nov., 1981
Earthwatch/Qld Museum Pyr. (QM). PARATYPE: 1 9,
same loc., 29.xi.1998. GM, DC, PB (CBM).

DIAGNOSIS. Immediately distinguished by the
narrower pronotum that is widest at basal third
and bears almost straight lateral margins in basal
third to two-fifth.

DESCRIPTION. Measurements. Length: 5.4-
6.0mm; width: 2.75-2. 95mm; Ratios. Length/
width of 9th antennomere: 2,7-2.75; width/length
of pronotum: 1 .4 1 ; width base/apex of pronotum:
1.92; width pronotum/head: 1.93-1.95;
length/width of elytra: 1.18-1.19; width
elytra/pronotum: 1.35-1.37.

Colour. As in S. trapezicollis.

Head. As in S. trapezicollis , though mandibles
remarkably elongate, and also antennae longer
than in all other species.

Pronotum (Figs 8B, 9C). Not as trapeziform as in
other species. Moderately wide, though
comparatively narrow in genus. Apex slightly
concave, anterior angles slightly produced,
obtusely rounded, sides evenly rounded in
anterior three thirds, then oblique, or even faintly
concave, slightly narrowed to basal angles, not
incurved at angles. Pronotum widest about at
posterior two thirds. Base laterally barely
oblique, slightly overlapping the gently oblique
base of elytra. Basal angles angulate, very
slightly produced laterally, gently dentiform,
angle almost right. Lateral borders coarsely
margined, apex and base more finely margined.
Lateral channel barely indicated. Median line
distinct, though shallow, neither reaching apex
nor base. Basal grooves rather deep, oblique,
situated close to middle. Both transverse sulci
barely recognisable, though basal region in
middle remarkably impressed. Anterior lateral
seta situated shortly behind middle, posterior
lateral seta situated slightly inside of basal angle.
Surface absolutely smooth, without any

86

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 9. Head and pronotum of Sitaphe. A, S. rotundata Moore; B, S. trapezicollis sp. nov.; C, S. parvicollis sp.
nov.; habitus of Sitaphe : D, S. rotundata Moore; E, S. trapezicollis sp. nov.; F, S. parvicollis sp. nov. Lengths:
4.9mm; 5.4mm; 5.4mm.

indication of microsculpture and puncturation,
remarkably glossy.

Elytra (Fig. 8C). As in S. trapezicollis , though
longer and more regularly oviform. Base far less
oblique than in other species, but otherwise rather
similar. In the holotype inner striae barely
impressed, outer striae barely recognisable, in the
paratype at least four inner striae well impressed,
intervals between clearly convex, outer striae
finer but still recognisable.

Lower surface. As in 5. trapezicollis.

Legs. As in S. trapezicollis.

Male genitalia (Fig. 7C). Genital ring as in S.
rotundata. Aedeagus as in S. rotundata , though
apex slightly longer. Left paramere very similar,
but right paramere less deeply sinuate on lower
surface than in S. rotundata.

Female genitalia. As in S. rotundata.

Variation. Due to scarce material little variation
noted in shape, but striking variation in
development and depth of elytral striae.

DISTRIBUTION (Fig. 1 1 ). South Peak of Bartle
Frere Range at the eastern margin of Atherton
Tableland, North Queensland.

COLLECTING CIRCUMSTANCES. Specimens
were collected by hand and by pyrethrum from
logs and tree trunks.

Sitaphe minuta sp. nov.

(Figs 7D, 8G, 11)

ETYMOLOGY. The name refers to the very small size of
this species.

MATERIAL. HOLOTYPE: d, QMT21849, NEQ:
17°02’S 145°40 1 E Lambs Head (east end), 1180m
29. xi. 1993. Monteith, Janetzki & Cook (QM).
PARATYPES: 1 9 , same data (QM); 1 9 , Lambs Head,

PSYDRINE GROUND BEETLES

87

lOkmW Edmonton, NQId. 12-13.xii.1988, 1200m GM&
GT(QM); 1 <J, same loc.,8-9.i.!990,GM&SM (CBM); 1
9 , same loc., 10-12.xii.1989, 1200mGM,GT&HJ(QM);
1 d, Isley Hills, 17°03’S 145°42’E 1050m 30.xi.1993; 1
9, Mt Williams, 900-1000m 16°55’S 145°40’E
2-3.xii.1993. DC, GM & HJ (QM).

DIAGNOSIS. Small, rather short species.
Distinguished from the most similar species, S.
rotundata Moore, by smaller size, narrower
pronotum with relatively narrower base,
distinctly incurved margin at base, and dentiform
basal angles; and from likewise similar S.
ovipennis sp. nov. by smaller size, narrower base
of pronotum and considerably shorter elytra.

DESCRIPTION. Measurements . Length: 4.1-
4.7mm; width: 2. 2-2. 5mm. Length/width of 9th
antennomere: 1 .9-2.0; width/length of pronotum:
1.52-1.56; width base/apex of pronotum:
1.81-1.86; width pronotum/head: 1.87-1.92;
length/width of elytra: 1.13-1.15; width
elytra/pronotum: 1.29-1.33.

Colour. As in S. trapezicollis.

Head. As in S. trapezicollis , though antennae
even shorter.

Pronotum (Fig. 8G). Moderately trapeziform,
because lateral margins very convex. Wide,
though comparatively narrow in genus, base
comparatively narrow. Apex slightly concave,
anterior angles slightly produced, obtusely
rounded, sides evenly curved over their whole
length, considerably incurved towards base,
therefore widest shortly in front of basal angles.
Base in lateral third rather oblique, adapted to the
oblique base of elytra. Basal angles developed as
tiny denticles. Lateral borders coarsely
margined, apex and base more finely margined.
Lateral channel barely indicated. Median line
distinct, though shallow, neither reaching apex
nor base. Basal grooves moderate, slightly
oblique, situated close to middle. Anterior
transverse sulcus absent, posterior sulcus
perceptible in the slightly impressed median part
of base. Anterior lateral seta situated at or shortly
behind middle, posterior lateral seta situated
slightly inside of basal angle. Surface absolutely
smooth, without any indication of
microsculpture and puncturation, remarkably
glossy.

Elytra. As in S. trapezicollis , though base slightly
less oblique. Lateral margins evenly curved,
elytra widest at basal fifth. Two or three inner
striae distinctly impressed and finely punctate,
intervals between gently convex. Outer striae
decreasingly distinct, barely punctate.

Lower surface. As in S. trapezicollis.

Legs. As in S. trapezicollis.

Male genitalia (Fig. 7D). Genital ring as in S.
rotundata. Aedeagus and parameres very much
as in S. rotundata , though right paramere slightly
shorter and less deeply sinuate.

Female genitalia. As in S. rotundata.

Variation. Some variation in depth and extent of
puncturation of elytral striae recognised.

DISTRIBUTION (Fig. 11). Lambs Head, Mt
Williams, and Isley Hills, all adjacent parts of the
Lamb Range mountain massif, North
Queensland.

COLLECTING CIRCUMSTANCES. This
species probably also lives rather on the bark or in
moss of logs and trunks than in the forest litter.
All specimens were sampled above 900m, most
were collected at the highest tops of the
respective mountains.

Sitaphe hamifera sp. nov.

(Figs 7E, 8J, 11)

ETYMOLOGY. The name of this species refers to the
dentiform basal angles of die pronotum.

TYPE MATERIAL EXAMINED. HOLOTYPE: <3,
QMT2 1 776, Cardwell Range, NE. Qld Mt Macalister area,
1000m 19.xii. 1986 Monteith, Thompson & Hamlet
pyrethrum knockdown (QM).

DIAGNOSIS. Medium sized, pronotum
comparatively narrow, decidedly trapezitorm,
with remarkably incurved lateral margins
towards base, with dentiform basal angles.
Further distinguished from all other species
except for S. parallelipennis sp. nov. by rather
parallel-sided, not oviform elytra. From S.
parallelipennis it is distinguished, inter alia , by
the narrower pronotum with comparatively wider
base, and by more regularly convex lateral
margins in middle of elytra.

DESCRIPTION. Measurements. Length:
5.2mm; width: 2.6mm; Ratios. Length/width of
9th antennomere: 2.25; width/length of
pronotum: 1.47; width base/apex of pronotum.
1.98; width pronotum/head: 1.89; length/width
of elytra: 1.14; width elytra/pronotum: 1.31.
Colour. As in S. trapezicollis.

Head. As in S. trapezicollis.

Pronotum (Fig. 8J). Trapeziform, because lateral
margins in anterior two thirds oblique and rather
straight. Comparatively narrow in genus, though
base comparatively wide. Apex faintly concave,

88

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 10. Distribution of Sitaphe spp. (part).

anterior angles slightly produced,
obtusely rounded, sides very little
curved in anterior two thirds, then
remarkably convex and deeply
incurved towards base, therefore
widest well in front of basal
angles, almost at posterior sixth.

Base laterally gently obliquely
convex, rather adapted to the
moderately oblique base of elytra.

Basal angles developed as tiny
denticles. Lateral borders
coarsely margined, apex and base
more finely margined. Lateral
channel barely indicated. Median
line distinct, though shallow,
neither reaching apex nor base.

Basal grooves deeply punctiform,
obliquely extended anterio-
medially, situated close to middle.

Anterior transverse sulcus absent,
posterior sulcus perceptible in the
slightly impressed median part of
base. Anterior lateral seta situated
about at middle, posterior lateral
seta set inside of basal angle.

Surface absolutely smooth, with-
out any i ndication of microsculpture
and puncturation, remarkably
glossy.

Elytra . As in S. trapezicollis , highly convex,
though shape far less oviform. Base slightly less
oblique. Lateral margins more evenly curved,
elytra widest about at middle, apical curvature
very short. Three inner striae distinctly impressed
and finely punctate, intervals between gently
convex. Outer striae decreasingly distinct, barely
punctate.

Lower surface. As in S. trapezicollis.

Legs. As in S. trapezicollis.

Male genitalia (Fig. 7E). Genital ring and
aedeagus much as in S. rotundata. Left paramerc
with considerably shorter apex than in S.
rotundata , right paramere slightly less deeply
sinuate on lower surface.

Female genitalia. Unknown.

Variation. Unknown.

DISTRIBUTION (Fig. 12). Mt Macalister in
Cardwell Range, North Queensland.

COLLECTING CIRCUMSTANCES. The
holotype was collected by py rethrum knockdown
on logs or trees at 1 000m, which is close to the top
of this mountain.

Sitaphe incurvicollis sp. nov.

(Figs 7F, 8E,F, 12)

ETYMOLOGY. Refers to the deeply incurved lateral
margins in the basal part of the pronotum.

MATERIAL. HOLOTYPE: <J, QMT21782, Upper
Boulder Creek 11km N Tully, NE Qld. 5-7 Dec 1989.
1000m Monteith, Thompson & Janetzki (QM).
PARATYPES: 1 9 , Upper Boulder Creek 6km N Tully,
NE Qld. 4.xii. 1989. 100-500mGM,CiT& HJ(QM); 1 6,
same loc., 650-900m, 24-27.X. 1983 GM, DY & GT (QM);
2 9, same loc., 500-600m 24-27.X.1983 GM, DY & GT
(CBM, QM); 1 6 , Tully Falls. N Qld. 8.xii.l990, 750m
GM, GT & HJ Pyr. Logs & Trees (QM); 1 <j,
Koombooloomba N Q. 1/53 GB. / M. 101. / Sitaphe
rotundata Moore det. B.P. Moore '69 (ANIC).

DIAGNOSIS. Medium sized, rather short. Due to
the basally incurved lateral margin and small,
dentiform basal angle of the pronotum probably
most closely related to S. minuta sp. nov. and S.
hamifera sp. nov. Distinguished from S. minuta
by larger size and wider pronotum with wider
base; and from S. hamifera by less triangular and
much wider pronotum, and by more oviform
elytra the widest diameter of which is close to the

PSYDRINE GROUND BEETLES

89

DESCRIPTION. Measurements. Length:
5.0-5.4mm; width: 2.65-2. 8mm. Length/width of
9th antennomere: 2. 1-2.2; width/length of
pronotum: 1.54-1.59; width base/apex of
pronotum: 1.88-1.94; width pronotum/head:
2.02-2.09; length/width of elytra: 1.11-1.13;
width elytra/pronotum: 1 .28- 1.31.

Colour. As in S. trapezicollis , though elytra
distinctly piceous which colour, however, may be
caused by incomplete pigmentation due to recent
hatching.

Head. As in S. trapezicollis , though eyes slightly
larger and orbits shorter.

Pronotum (Fig. 8E). Moderately trapeziform,
because lateral margins rather convex. Wide,
though comparatively narrow in genus, but base
comparatively wide. Apex slightly concave,
anterior angles slightly produced, obtusely

punctate.

Lower surface. As in S. trapezicollis.
Legs. As in S. trapezicollis.

Male genitalia (Fig. 7F). Genital ring and
aedeagus as in 5. rotundata. Left paramere with
shorter apex than in S. rotundata, right paramere
slightly shorter and with remarkably straight
apex.

Female genitalia. As in S. rotundata.

Variation. Little variation in size and shape.
Variation in depth and extent of puncturation ol
elytral striae recognised.

DISTRIBUTION (Fig. 12). Upper Boulder
Creek at eastern end of the Walter Hill Range, to
the south of the Atherton Tableland, North
Queensland.

90

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 12. Distribution of Sitaphe spp. (part).

COLLECTING CIRCUMSTANCES. This
species probably lives on the bark or in moss of
logs and trunks. Altitude records from 500-
1 000m, with most 500-750m. These altitudes are
lower than for most other species.

Sitaphe parallelipennis sp. nov.

(Figs 7G, 8H, 81, 10)

ETYMOLOGY. Refers to the rather parallel shape of
elytra.

MATERIAL. HOLOTYPE: <J, QMT21838, NE Qld:
16°22’S 145°13'E 7km N Mt Spurgeon (Camp 2)
17- 19.x. 1991. 1200- 1250m Monteith, Janetzki, Cook &
Roberts (QM). PARATYPES: 3 <3, 12 9, same data

(CBM, QM); 4 <3, 7 9 . Stewart Ck 4km
NNE Mt Spurgeon (Camp 1), 16°24’S
145°13’E 1250-1 300m 15-20.X.1991.
GM, HJ, DC & LR (CBM, QM); 2 d , Mt
Spurgeon Summit 16°26’S 145°12’E
21. xL 1997. 1320m GM, DC & Burwell
(QM); 2 d, 2 9, Mt Spurgeon Summit
16°26’S 145°12'E 21.xi.1997. 1320m
GM. Pyrethrum, trees & logs (QM); 1 d,
2km SE Mt Spurgeon 16°27'S 145°12’E
13-14.X.1991. 1100m GM. HJ & DC
(QM); 2 d, 2 9, Black Mtn. 4.5km N of
Mt Spurgeon. 16°24'S 145°12'E
1 200- 1 250m 1 7- 1 8.x. 1991. GM, HJ, DC
& LR (QM). I d, Windsor Tbld, N Qld.
9.i. 1 989, 1 225m E. Schmidt & ANZSES
Site 3, Pyr.(QM); 1 9, same loc.,
27.xii.88 - 9.L1989 E. Schmidt &
ANZSES , pitfall (QM); 2 9. Mt Demi,
7km SE of Mossman 29.x. 1983, 1 100m
DY & GT Pyr. in RF. (QM); 1 9, Mt
Demi summit, 1000m 16°30'S 145°19'S
1 6- 1 7.xii. 1 995 GM & GT (QM); 1 d , Mt
Lewis, Via Julatten, N Qld. 3,500-4000'
27-28.xi.1965, GM (QM); 2 d, 1 9, Mt
Lewis Rd (Hut) 16°31'S 145°16'S 14
July 1996, 1200m GM(QM); 1 d, 2.5km
N Mt Lewis, via Julatten 3.xi.l983,

1 040m D Y & GT Pyr. in RF. (QM ); 1 9 ,
Mt Lewis summit, via Julatten,
IO.ix. 1981 GM & DC, QM Berl. 287 Rf,
1200m Log debris (QM); 2 d, same
loc., via Julatten, 9.ix.l981 GM & DC
(QM); 1 9, Mt Lewis Rd end, 10km N
Mt Lewis, N Qld. 25.xi.1990. 1100m.
GM. GT, DC. RS & HJ (QM); 2 9,
5.5km N of Mt Lewis, via Julatten
1 1 00m, 8.ix.I981 GM&DC(QM);2 9,
Hilltop, 18km N Mt Lewis I6°30’S
1 45° 16 E 23.xi.l998 GM. Pyr. trees.
1300m, RF(QM); 1 d, Carbine Tbld, NE
Qld Plane Crash Site 27-28.xi. 1990,
1330m GM, GT, DC, RS & HJ Pitfall
Traps & Hand (QM); 1 9, Nr Plane
Crash I lkm NW Mossman, N Qld
28.xii.1989, 1240m ANZSES. Pyr.(QM); 1 9, Devil's
Thumb 12km NW Mossman. N Qld 27.xii.1989, 1000m
ANZSES, Pyr.(QM); 4 d, 6 9 , Devils Thumb area, 10km
NW Mossman 9-10 Oct. 1982, 1000-1 180m GM, DY &
Thomson (CBM, QM);1 9, Pauls Luck, Platypus Ck,
13km W Mossman, N Qld. l-2.i.l990, 1 100m ANZSES,
Pyr.(QM); I d, Mossman Bluff Summit 10km W
Mossman, N Qld. 18.xii.1988. 1300m GM & GT
Pyrethrum, Trees & Rocks (QM); 1 d, Mossman Bluff
Track 9- 1 0km W Mossman, NQ 2 1 .xii. 1 989, 1 000- 1 300m
GM, GT, ANZSES (QM);

DIAGNOSIS. Fairly large, rather short, with
basally slightly incurved lateral margins of
pronotum and dentiform basal angles.
Distinguished from all species with similar base

PSYDRINE GROUND BEETLES

91

of pronotum by the remarkably elongate
aedeagus and parameres. Further distinguished
from S. ovipennis sp. nov. by shorter, parallel, and
less oviform elytra.

DESCRIPTION. Measurements. Length: 4.9-
5.6mm; width: 2.6-2. 9mm. Length/width of 9th
antennomere: 2.1-2.25; width/length of
pronotum: 1.56-1.59; width base/apex of
pronotum: 1.88-1.93; width pronotum/head:
1.97-2.01; length/width of elytra: 1.13-1.15;
width elytra/pronotum: 1.26-1.29.

Colour. As in S. trapezicollis.

Head. As in S. trapezicollis.

Pronotum (Fig. 8H). Very similar to that of S.
ovipennis , though slightly wider.

Elytra (Fig. 81). As in S. trapezicollis , though
base slightly less oblique. Lateral margins
anteriorly rather gently curved, shape not
distinctly oviform though in middle rather
parallel-sided. Elytra widest about at basal third,
or even further behind, in median part lateral
margins very little convex, almost straight. Three
inner striae, rarely also 4th stria in part, distinctly
impressed and finely punctate, intervals between
gently convex. Outer striae decreasingly distinct,
barely punctate.

Lower surface. As in S. trapezicollis.

Legs. As in S. trapezicollis.

Male genitalia (Fig. 7G). Genital ring as in S.
rotundata. Aedeagus structurally as in S.
rotundata , but much longer and narrower and
with longer apex which is distinctly curved
down. Both parameres considerably longer than
in S. rotundata , left paramere also narrower, right
paramere far less deeply sinuate on lower
surface.

Female genitalia. As in S. rotundata.

Variation. Rather little variation in size and shape
noted. Only some unimportant variation in depth
and extent of puncturation of elytral striae
recognised.

DISTRIBUTION (Fig. 10). Widespread on
Carbine and Windsor Tablelands, west of
Mossman, North Queensland.

COLLECTING CIRCUMSTANCES. Many
specimens were collected by pyrethrum
knockdown on logs and trees, single specimens
also by pitfall trapping or hand collecting. This
indicates that this species lives on the bark or in
moss on logs and trunks. All specimens were
collected above 1000m, most at or near the
summits of the respective mountains or ranges.

Sitaphe ovipennis sp. nov.
(Figs 7H, 8K, 8L, 10)

ETYMOLOGY. Refers to the ovifonn elytra.

MATERIAL. HOLOTYPE: S. QMT21756, NE Qld,
Thornton Peak, 11km NE Daintree. 1 100- 1200m 30.x.- 1
Nov., 1983 Monteith, Yeates & Thompson (QM).
PARATYPES: 7 d, 6 9 , same data (CBM, QM); 2 d,
Thornton Peak via Daintree. 1000- 1300m. 20-22. ix. 1981
GM & DC (QM); 8 d, 3 9 , Thornton Peak, 1 100-1300m
via Daintree, N Qld. 24-27.ix.1984 GM & SM (CBM,
QM).

DIAGNOSIS. Medium sized, elongate, with
basally slightly incurved lateral margins of
pronotum and dentiform basal angles.
Distinguished from all species with similar base
of pronotum by much longer, decidedly oviform
elytra. Further distinguished from nearby Y
parallelipennis sp. nov. by shorter aedeagus and
parameres.

DESCRIPTION. Measurements. Length: 4.6-
5.3mm; width: 2.40-2.75mm. Length/ width of
?th antennomere: 1.8-1. 9; width/length o
pronotum: 1.52-1.56; width base/apex of
pronotum: 1.86-1.91; width pronotum/head:
1.89-1.94; length/width of elytra: 1.18-l.zU,
width elytra/pronotum: 1.23-1.27.

Colour. As in 5. trapezicollis , though in most
specimens elytra rather piceous which colour,
however, may be caused by incomplete
pigmentation due to recent hatching.

Head. As in S. trapezicollis.

Pronotum (Fig. 8K). Moderately trapeziform,
because lateral margins rather convex. Wide,
though with comparatively narrow base. Apex
gentlv concave, anterior angles slig y
produced, obtusely rounded, sides evenly curved
over their whole length, moderately incurved
towards base, therefore widest shortly m front ot
basal angles. Base in lateral third rather oblique
adapted to the oblique base of elytra. Basal angles
developed as tiny denticles. Lateral borders
coarsely margined, apex and base more tine y
margined. Lateral channel barely indica ecL
Median line distinct, though shallow, neither
reaching apex nor base. Basal grooves deep,
punctiform. obliquely extended anterio-
medially, situated close to middle. Anterior
transverse sulcus absent, posterior sulcus
perceptible in the rather deeply impressed
median part of base. Anterior lateral seta situated
shortly behind middle, posterior lateral seta set
slightly inside of basal angle. Surface absolutely

92

MEMOIRS OF THE QUEENSLAND MUSEUM

smooth, without any indication of microsculpture
and puncturation, remarkably glossy.

Elytra (Fig. 8L). As in S. trapezicollis, though
considerably longer and base slightly less
oblique. Lateral margins rather evenly curved,
shape distinctly oviform with rather elongate
apex. Elytra widest about at basal fifth. Two inner
striae, rarely also 3rd stria in part, distinctly
impressed and finely punctate, intervals between
gently convex. Outer striae decreasingly distinct,
barely punctate.

Lower surface. As in S. trapezicollis.

Legs. As in S. trapezicollis.

Male genitalia (Fig. 7H). Genital ring as in S.
rotundata. Aedeagus as in S. rotundata , though
apex even shorter. Left paramere very similar,
though apex slightly shorter than in S. rotundata ,
right paramere slightly shorter, slightly less
deeply sinuate on lower surface, and apex
straight.

Female genitalia. As in S. rotundata.

Variation. Rather little variation in size and shape
noted. Only some unimportant variation in depth
and extent of puncturation of elytral striae
recognised.

DISTRIBUTION (Fig. 1 0). Thornton Peak, north
of Daintree, North Queensland.

COLLECTING CIRCUMSTANCES. All
specimens were hand collected, mostly from
surfaces of logs and tree trunks as with all species
of Sitaphe (GB.Monteith, pers. comm.). This
species has been collected only above 1000m on
the highest part of the Thornton Peak massif.

MEASUREMENTS AND RATIOS IN
SITAPHE MOORE

For better comparison of the species the meas-
urements and ratios of all species are compiled in
Table 2.

REMARKS. All taxa of Sitaphe are treated as
species, because in one area three taxa, in another
area two taxa are sympatric and most probably
even syntopic. Slight though well distinguished
differences can be noted in external structure,
mainly shape and structure of pronotum, and
shape of elytra. In all species, however, the male
genitalia are very similar in shape and structure.
Only minor differences in relative length of
aedeagus and parameres, shape of apex of
aedeagus, and shape and curvature of the
parameres have been noted.

Probably some of the allopatric taxa here
regarded as species could be as well described as
subspecies, but in view of the still unsatisfactory
knowledge of the number of existing taxa and of
their distribution I have decided to rate all taxa as
species, because it may be conceivable that even
more taxa possess overlapping ranges. Never-
theless, it should be stressed that in areas where
two or three taxa occur sympatrically, the
morphological divergence generally is greater
than between allopatric taxa. This is not too sur-
prising, because there must be some ecological
divergence in sympatric or even syntopic taxa
which, as a consequence, certainly will cause
more striking morphological differences.

PHYLOGENY AND BIOGEOGRAPHY

Since all existing taxa of Sitaphe , and their
respective ranges, are probably not yet known,
any considerations about phylogenetic relations
and biogeographical history of the species are
necessarily provisional. Nevertheless, some
considerations are possible at this stage. These,
however, are still very difficult to assess due to
the extremely close relationship between species.
Since the male genitalia are similar throughout
the genus, with the only divergence being in the
northern S. parallelipennis , which has a longer
aedeagus with bent apex, only external characters
are useful. But many of these are also extremely
uniform throughout the genus, e.g. shape of head,
length of mandibles, size of eyes, surface
structure of elytra. But shape of pronotum and of
elytra may be of some use. There is also the
problem of determining w'hich genus is the
nearest relative of Sitaphe. These obstacles make
any decision about apomorphy versus plesio-
morphy within Sitaphe extremely difficult.

Character states unique for the whole genus
Sitaphe are the elongate mandibles, depressed
eyes, wide, more or less conspicuously
trapezoidal pronotum, wide and short, dorsal ly
convex, oval-shaped elytra, reduction of elytral
striae, and absence of any microreticulation or
puncturation on the surface. However, as stated
above, most of these character states are uniform
throughout the genus. All species possess short
antennae and a remarkably trapezoidal pro-
notum, except for S. parvicollis , in which the
antennae are decidedly longer and narrower and
the pronotum is definitively less w'ide at base. In
the other species the pronotum is firmly attached
to the base of the elytra, and is almost as wide as
the latter. In S. parvicollis the elytra are also
comparatively longer, making this species

PSYDRINE GROUND BEETLES

93

TABLE 2. Sitaphe spp. measurements.

N

body length
(mm)

ratio length/
width 9th
antennomere

ratio width/
length
pronotum

ratio width
base/apex
pronotum
rotundata

ratio width
pronotum/
head

ratio length/
width elytra

ratio width
elytra/
pronotum

S. rotundata

10

4.6-5. 8

2.2-2.5

1.60-1.67

1.98-2.07

2.02-2.12

1.12-1.15

1.25-1.30

S. trapezicollis

10

4.9-6.2

2.25-2.5

1.49-1.53

2.02-2.07

2.03-2.11

1.12-1.13

1.29-1.34

S. parvicollis

2

5.4-6.0

2.7-2.75

1.41

1.92

1.93-1.95

1.18-1.19

1.35-1.37

S. minuta

7

4. 1-4.7

1. 9-2.0

1.52-1.56

1.81-1.86

1.87-1.92

1.13-1.15

1.29-1.33

S. hamifera

1

5.2

2.25

1.47

1.98

1.89

1.14

1.31

S. incurvicollis

7

5.0-5.4

2. 1-2.2

1.54-1.59

1.88-1.94

2.02-2.09

1.11-1.13

1.28-1.31

S. parallelipennis

10

4.9-5.6

2.1-2.25

1.56-1.59

1.88-1.93

1.97-2.01

1.13-1.15

1.26-1.29

S. ovipennis

10

4.6-5.3

1.8-1. 9

1.52-1.56

1.86-1.91

1.89-1.94

1.18-1.20

1.23-1.27

generally longer and narrower than all others.
When compared with other, southern
tropopterine genera, it seems conceivable at the
first glance, then, that this species should be most
plesiotypic within the genus, at least with respect
to the above character states.

This compact, smooth, convex, remarkably
wedge-shaped body structure, in combination
with the narrow head and elongate,
stiletto-shaped mandibles, strongly suggests the
‘cychriform’ body shape of certain snail eating
carabid groups as exemplified by the northern
hemisphere Cychrus and certain Cardbus.
Although virtually nothing is known about
foraging methods and diet of Sitaphe , its highly
specialised body shape suggests either a similar
snail eating habit in all Sitaphe , or a foraging
method of forcing the slender forebody into
crevices in the bark raking for prey and using the
elongate mandibles as pincers. Both habits, in
turn, could explain the highly similar shape and
structure in almost all species.

S. trapezicollis has a strikingly trapezoidal
pronotum, the basal angles of which are not
incurved, and which is very tightly adapted to the
base of the elytra. If we admit that the
morphological trend within the genus is directed
to a most compact, short, ovoid body shape with
almost jointless adaptation of pronotum and
elytra, then S. trapezicollis would seem most
apomorphic.

All other species have the base of prothorax
more or less incurved to form an obtuse (5.
rotundata) or faintly dentiform (S. minuta , S.
parallelipennis , S. ovipennis) or markedly
dentiform basal angle ( S . incurvicollis , S.
hamifera). With respect to this character, S.
rotundata probably represents the most
plesiomorphic state.

However S. parvicollis is longer and narrower
than all other species, has the basal part of
pronotum remarkably deeply impressed, and also
possesses distinctly longer antennae and
mandibles. In combination these features give
this species an even stronger ‘cychriform' body
shape impression than in the other species. From
the speculative assumption that all Sitaphe eat
snails we may further speculate that this special
body shape in S. parvicollis derives from a more
specialised diet and/or foraging mode, e.g.
consumption of snails with narrower and deeper
aperture.

If all these considerations and assumptions
prove right, then S. rotundata would be the
species that is most plesiomorphic, at least with
respect to the character states considered. Sitaphe
trapezicollis and S. parvicollis would be most
apomorphic, S. incurvicollis and S. hamifera
would be somewhat intermediate, whereas S.
minuta and both northern species (S.
parallelipennis and S. ovalipennis) would be
more closely related to S . rotundata. However,
the last two species are unique in at least one
character: S. parallelipennis has fairly dissimilar
male genitalia, and S. ovipennis has longer elytra
than usual.

We could combine these raw assumptions
about relationships with the distribution pattern
of the species to get a glimpse of evolution and
biogeographic history of the genus. Perhaps five
assumptions can be set up:

1) There is no indication where the early stock of
the genus came from, except that, as a member of
an originally cool adapted group, it probably
arrived from the south. The nearest relatives of
Sitaphe live in southeastern Australia and belong
to the Theprisa-Theraphis-Phersita complex, but

94

MEMOIRS OF THE QUEENSLAND MUSEUM

I do not know which of these three genera is
closest.

2) Sitaphe is morphologically highly isolated,
though the species within the genus are extremely
closely related. This suggests that the genus, or at
least its original stock, is a comparatively old
element of the North Queensland rain forests but
that the taxonomic radiation within the genus has
probably occurred comparatively recently.

3) Based the above considerations it seems
conceivable that the widespread S. rotundata is
the most plesiotypic species and may be most
similar to the original stock of the genus. More
apotypic species have more restricted ranges,
either in the north on Carbine Tableland (S.
parallelipennis) and Thornton Peak (5.
ovipennis ), or in the south on Cardwell Range (5.
hamifera) and Walter Hill Range (S. incurvi-
collis ), or in the east on Mts Bellenden Ker and
Bartle Frere (S. trapezicollis , S. parvicollis).
Actually the eastern margin of* Atherton Table-
land is inhabited by four different species
including 5. minuta , that occurs on the mountains
northwest of Bellenden Ker/Bartle Frere, being
more closely related to S. rotundata.

4) The high Bellenden Ker Range (comprising
Mt Bellenden Ker, Mt Bartle Frere and the
Massey Range) is inhabited by three species.
Over part of their ranges these three species are
not only sympatric but actually syn topic and have
been collected within the same fogging or sieving
sample. Thus they may actually live together on
the same log. Characteristically, the most
plesiotypic species ( S . rotundata) again has the
widest range, since it occurs not only on
Bellenden Ker and Bartle Frere, but also on the
contiguous Massey Range. The more evolved S.
trapezicollis occurs on both high peaks, whereas
S. parvicollis , probably the most specialised
species, has been found only on the summit of the
highest peak, Bartle Frere.

5) It is notable, therefore, that the three sympatric
species on the Bellenden Ker Range show the
greatest morphological divergence in the whole
genus. This is presumably the result of an
ecological shift leading to more essential
structural differences than usual.

If the above assumptions are correct, then we
have another example of a generalised scenario
of distribution patterns very similar to that
recognised in some other insect groups (e,g.
Philipis (Baehr 1995b)) in the Wet Tropics:

- One or few plesiotypic species exhibit a wide
distribution over the central Wet Tropics

(Atherton Tableland in the broad sense), also
further to the south.

- Several or many more or less apotypic species
are arranged in an almost semicircular pattern
around the central region or these widespread
species to the north, east, and south of it.

Meonis Castelnau, 1867

Meonis Castelnau, 1867: 69; Castelnau. 1868: 155; Csiki,
1929: 484; Moore, 1963: 288; Moore et ai. t 1987: 152.

DIAGNOSIS. Genus of tribe Meonini (which
includes Raphetis). Moore et al. (1987) included
five species, three of which have been so far
recorded only in the northern half of New South
Wales. However, the genus urgently needs
revision to settle the status of synonymised taxa
as well as to accommodate certain taxa that are
probably yet undescribed.

Meonis ater Castelnau, 1867

Meonis ater Castelnau, 1867: 70; Castelnau, 1868: 156;
Sloane. 1900: 563; Sloane, 1911: 827; Sloane, 1916: 201;
Csiki, 1929: 484; Straneo, 1941: 86; Moore et al., 1987:
152.

Meonis amplicollis Sloane, 1915: 448; Sloane, 1916: 201;
Csiki, 1929: 484; Straneo, 1941: 86; Moore et al., 1987:
152.

REMARKS. Provided the above synonymy is
correct, this species occurs in southeastern
Queensland (e.g. on Lamington Plateau) and
immediately adjacent northeastern New South
Wales (Clarence River area). It is a ground
dwelling species that inhabits leaf litter and is
also found under logs in subtropical to temperate
rain forest. The elongate, porrect mandibles,
cychriform head and glabrous surface suggest
molluscs as the diet for this and other species of
Meonis.

DISTRIBUTION. Southeastern Queensland and
northeastern New South Wales near border.

Meonis niger Castelnau, 1867

Meonis niger Castelnau, 1867: 70; Castelnau, 1868: 156;
Sloane, 1911: 826; Sloane, 1916: 201; Csiki, 1929: 484;
Straneo, 1941: 86; Moore et al., 1987: 152.

DISTRIBUTION. Southeastern Queensland and
adjacent northeastern New South Wales, in
Queensland apparently with a slightly wider
range than the preceding species.

Raphetis Moore, 1963

Raphetis Moore, 1963: 288; Moore et al., 1987: 152.

TYPE SPECIES. Raphetis darlingtoni Moore, 1963, by
original designation.

PSYDRINE GROUND BEETLES

95

DIAGNOSIS. Meonini. As described by Moore
(1963) it is distinguished from Meonis by
glabrous instead of setose paraglossae, spinose
laciniae bearing a strong terminal hook instead of
densely setose lacinae lacking the terminal hook,
weakly bordered base of elytra instead of
unbordered base, and a normal shaped scutellum,
whereas in Meonis the scutellum is tiny.

In most other respects the two genera are very
similar and also share the characteristic body
shape, small eyes, and elongate, porrect
mandibles. Even the male genitalia do not show
much structural differences, apart from some
minor differences of size and shape of aedeagus,
and length of parameres and extent of their
chaetotaxy. Female stylomeres are very uniform
throughout Raphetis , and they only exhibit minor
differences of shape and number of ensiform
setae on rim of first and latero- ventral margin of
second stylomere.

REMARKS. The Meonini, apart from Raphetis ,
includes only Meonis Castelnau in Australia.
Meonis occurs in subtropical and cool temperate
rain forests of southeastern Australia from south-
ern New South Wales to southern Queensland. In
southern Queensland, the two genera are
sympatric on the Springbrook Plateau.

Moore described R. darlingtoni from Eungella
Range west of Mackay and R. gracilis from the
western Atherton Tableland. Until now Raphetis
was regarded as very rare, and the two species
were described from one or two specimens.
Through the systematic sampling of the Queens-
land Museum workers 38 additional specimens
are now available, mainly from four rather
restricted areas: Springbrook Plateau on the
Queensland/ New South Wales border;
Bellenden Ker/Bartle Frere Range at the eastern
slope of Atherton Tableland; the Millaa Millaa /
Mt Fisher area on the western part of Atherton
Tableland (from where R. gracilis Moore was
described); and the Carbine Tableland to the
north of Atherton Tableland. The fifih area from
where the genus was recorded is Eungella
Plateau west of Mackay in central eastern
Queensland, where R. darlingtoni Moore occurs.
Hence the distribution seems to be very frag-
mented, though in view of the quite exhaustive
sampling that has taken place on other
intermediate mountain massifs I suspect that it
may reflect the true distribution.

KEY TO THE TAXA OF RAPHETIS MOORE

1 . Pronotum markedly cordate, deeply sinuate in front of

basal angles; base little wider than apex (Figs 14ABD,
15B-C) 2

Pronotum little cordate, barely sinuate in front of basal
angles; base distinctly wider than apex (Figs 14C, 1 5 A).
Southeastern Queensland curta sp. nov.

2. Pronotum laterally more convex; elytra distinctly striate,
striae coarsely punctate; aedeagus suddenly curved to
apex. Eungella Plateau, central eastern Queensland

darlingtoni Moore

Pronotum laterally less convex; elytra barely striate,
striae, when visible, impunctate; aedeagus more evenly
curved to apex (Fig. 13B-F) 3

3. Apex of elytra with distinct, slightly curved spine (Fig.

1 4G). Mt Lewis area, southern Carbine Tbld . . gracilis

spinosa subsp. nov

Apex of elytra not spinose, at most with tiny denticle (Fig.
14H,I) 4

4. Lower margin of aedeagus slightly sinuate near apex (Fig.

5. Pronotum narrow, barely wider than long, relatively

narrower in relation to head (Fig. 14A); elytra relatively
longer (see appendix). Size generally smaller,
7.0-7. 8mm. Millaa Millaa/Mt Fisher area, western part
of Atherton Tbld gracilis gracilis Moore

Pronotum wider, distinctly wider than long, relatively
wider in relation to head (Fig. 14B); elytra relatively
shorter (see append i x ). S i ze ge n era 1 1 y larger. 7.9-8. 5mm.
Bellenden Ker/Bartle Frere Range, eastern margin of
Atherton Tbld gracilis frerei subsp. nov.

6. Apex of elytra w r ith tiny denticle (Fig. 14H); eyes smaller,

<2 x as long as orbits; pronotum narrower (see
appendix). Plane Crash Site at eastern margin, and Mt

Spurgeon at western margin of Carbine Tbld . gracilis

subarmata subsp. nov

Apex of elytra without denticle, regularly convex (Fig.
141); eyes larger, well >2 x as long as orbits; pronotum
wider (sec appendix). Area north of Mt Spurgeon at
western margin of Carbine Tbld . . gracilis spurgeoni
subsp. nov.

Raphetis darlingtoni Moore, 1963
(Fig. 16)

Raphetis darlingtoni Moore, 1963: 288; Moore et al., 1987.
153.

MATERIAL. HOLOTYPE: d, Eungella Rge. W. of
Mackay Q 2-3000' Nov57 Darlingtons/ Raphetis gen.n.
darlingtoni sp.n. holotype 6 Det. B.P. Moore 63 ( ANIC).

REMARKS. The descriptions of R. darlingtoni
and R. gracilis Moore, are brief and hence
contain some mistakes. The elytra are not tree in
R. darlingtoni , but are fused as in R. gracilis. In
the holotype, however, they arc opened, because
the abdomen had been removed. Separating of
the normally fused elytra also has been observed
in preserved specimens of R. gracilis.

96

MEMOIRS OF THE QUEENSLAND MUSEUM

The parameres are not dissimilar in R.
darlingtoni and similar in R. gracilis , but are
dissimilar in both species, though slightly less so
in R. gracilis , because the left paramere is less
wide and less triangular in the latter species.
Moreover, the right paramere is as well setose on
its lower margin in R. gracilis as it is in R.
darlingtoni.

Hence, the two species are less different in
certain respects than the descriptions indicate.

For better distinction, measurements and ratios
of the holotype of R. darlingtoni are added below.

DIAGNOSIS. Easily distinguished by the
remarkably cordate prothorax, deeply impressed
elytral striae, and deeply curved aedeagus with
elongate, blade-like apex.

DESCRIPTION. Measurements. Length: 6.1mm;
width: 2.55mm. Length eye/orbit: 3.2; width/
length of pronotum: 1.17; width base/apex of
pronotum: 1.15; width pronotum/head: 1.55;
length/width of elytra: 1.37; width elytra/
pronotum: 1.35.

DISTRIBUTION (Fig. 16). Eungella Plateau
west of Mackay, central eastern Queensland.
Known only from type locality. This species has
not been recaptured since description.

Raphetis curta sp. nov.

(Figs 2D, 13 A, 14C,F, 15A,D, 16)

ETYMOLOGY. Refers to the short, compact hind body
compared with the other species.

MATERIAL. HOLOTYPE: d,QMT93419, SEQ:
28°15’S 153°16 T E Springbrook Repeater, 21 Dec 1996.
1000m GB. Monteith Pyrethrum, dead trees (QM).
PARATYPES: 1 6, same loc., GM. Pyr. Tree trunks,
1000m (CBM); 1 9, same loc., 9.XII.1972 GM. & SM /
Raphetis sp. n. deL B.P. Moore ‘74 (QM).

DIAGNOSIS. Easily distinguished by the barely
cordate prothorax, short abdomen, and shorter
and wider, on lower margin distinctly bisinuate,
aedeagus.

DESCRIPTION. Measurements. Length: 5.3-
5.7mm; width: 2.3-2.6mm. Length eye/ orbit:
2.3-2.4; width/length of pronotum: 1.20- 1.23;
width base/apex of pronotum: 1.34-1.40; width
pronotum/head: 1.51-1.58; length/width of
elytra: 1.31-1.35; width elytra/pronotum: 1.36-
1.43.

Colour. Dark piceous to blackish. Lateral margin
of elytra narrowly reddish. Clypeus, labrum, and
mandibles reddish-piceous, palpi and antennae

reddish. Femora and tarsi reddish, tibiae
reddish-piceous. Lower surface of anterior body
dark piceous, abdomen reddish-piceous.

Head (Fig. 15A). Considerably narrower than
pronotum. Eyes comparatively large (in genus),
moderately convex, laterally fairly protruding.
Eyes separated from frons by a narrow, straight
furrow. Frons with two irregular, slightly curved
furrows medially of eyes that are prolonged to
clypeal suture. Furrows medially widened to
some shallow, irregular grooves, at posterior end
laterally bordered by convex ridge. Anterior
supraorbital seta situated close to eye near
anterior border of eye, posterior seta situated well
behind eye and rather moved to upper part of
head. Neck separated by a shallow, transverse
furrow. Clypeal suture distinct. Labrum
anteriorly slightly concave, 6-setose and with
some shorter hairs around the anterior angles.
Mandibles elongate, straight, though shorter than
in related species, inner margin straight for a long
distance, then suddenly incurved, with acute
apex, with elongate seta in scrobe. Mentum with
acute triangular tooth. Submentum bisetose.
Gula quadrisetose. Glossa short, narrow,
bisetose, paraglossae hyaline, surpassing glossa.
Lacinia very elongate, inner margin with few
strong spines, apex markedly incurved, very
acute. Both palpi glabrous, apex transverse.
Antenna rather slender and elongate, surpassing
base of pronotum by about two antennomeres,
pilose from half of 4th antennomere, central
antennomeres c. 2.5 x as long as wide. Surface
of head absolutely smooth, without any
indication of microsculpture, remarkably glossy.

Pronotum (Figs 14C, 15A). Wide, moderately
convex, not markedly cordiform. Apex gently
concave, anterior angles slightly produced,
obtusely rounded, sides evenly curved, widest at
middle, very weakly sinuate in basal half. Basal
angles angulate, almost rectangular, angle c. 95°.
Base laterally slightly oblique. Apex not
margined, base laterally more or less distinctly
margined. Lateral channel narrow, slightly
explanate near base. Median line distinct, well
impressed, neither reaching apex nor base. Basal
grooves straight, rather linear, fairly deep. Both,
anterior and posterior transverse sulci barely
indicated. Anterior lateral seta situated slightly in
front of middle, posterior lateral seta absent.
Surface absolutely smooth, without any
indication of microsculpture and puncturation,
though near median line with some short,
transverse furrows, remarkably glossy.

PSYDRINE GROUND BEETLES

97

FIG 1 3. Male genitalia of Raphetis spp. A, R. curta sp. nov., aedeagus, parameres, and genital ring; B R. gracilis
gracilis Moore, aedeagus, parameres, and genital ring; C, R. gracilis spinosa su sp. nov., ae eagus a
parameres; D, R. gracilis frerei subsp. nov., aedeagus and parameres; E. R. gracilis sp subsp. nov.,
aedeagus and parameres; F, R. gracilis subarmata subsp. nov., aedeagus and parameres. u e

Elytra (Fig. 14F). Wide, convex, considerably
wider than pronotum, widest at humeri. Humeri
gently angulate, lateral margins feebly convex,
elytra widest at anterior fourth or third, than
faintly though evenly narrowed. Lateral part of
apex slightly sinuate at position of crossing of
epipleurae, then evenly convex. Basal margin
complete, lateral margin narrow throughout.
Scutcllary puncture and seta present, at base of
1st stria. Scutellary stria short, inside 1st stria.
Striae extremely fine, barely impressed, though
in one specimen 1st stria fairly distinct. External
striae barely recognisable. Striae barely punctate,
or internal striae very finely punctate. 8th stria
complete, well impressed. Near apex 1st, 2nd,
5th, and in particular 7th striae well impressed,

7th stria forming an elongate, fairly deep furrow.
Intervals depressed, at most 1 st and 2nd intervals
very feebly convex in apical half. Disk
impunctate. Marginal series consisting of two
groups of 6-7 and 6 setiferous punctures,
respectively, which are widely separated in
middle. Inside of deepened 7th stria with two
additional punctures near apex. Some of the mar-
ginal setae very elongate. Intervals absolutely
smooth, without any traces of microreticulation,
highly glossy, slightly iridescent.

Lower surface . Metepisternum short, slightly
longer than wide at apex. Mesepistemum and
metepisternum coarsely though very sparsely
punctate. Terminal stemite in male bisetose, in

98

MEMOIRS OF THE QUEENSLAND MUSEUM

female quadrisetose along margin, though with
some additional shorter setae.

Legs. Fairly elongate. 5th tarsomere asetose
beneath. Anterior tarsus in male barely wider
than in female, lst-3rd tarsomeres asymmetric-
ally squamose beneath.

Male genitalia (Fig. 13 A). Genital ring short and
wide, asymmetrically triangular, with short,
rounded apex. Aedeagus short and compact,
laterally depressed, lower surface markedly
bisinuate, apex short, very wide, convex, slightly
turned. Internal sac rather complexly folded, with
several sclerotised plates within. Both parameres
comparatively short and wide, left larger than
right, with wide, obtuse apex, 2-5 apical setae and
0-2 shorter setae on lower margin. Right
paramere remarkably short, with acute apex and
c. 12 elongate setae at apex and along the whole
of lower margin.

Female genitalia (Fig. 2D). Stylomere 1 with 2
ensiform setae at middle of ventro-apical margin.
Stylomere 2 rather short, with short apex and 2
large dentiform ventro-lateral ensiform setae of
about similar size in middle of lateral margin.
Near apex with a large, oblong pit and a short
nematiform seta originating from that pit. In
middle of dorso-median surface with a large,
dentiform, dorso-median ensiform seta. Lateral
plate with a densely setose area at median apical
margin.

Variation. Some minor variation in relative shape
of pronotum and elytra, depth of elytral striae and
number of setae on the parameres.

DISTRIBUTION (Fig. 16). Springbrook Plateau
at Queensland/New South Wales border.

COLLECTING CIRCUMSTANCES. Collected
by pyrethrum knockdown in upland rainforest on
'tree trunks' and ‘dead trees’. Apparently, this
species lives in crevices of dead wood and,
according to observations made by G. Monteith,
it comes out to forage at the bark surface at night.

Raphetis gracilis Moore
(Fig. 17)

Raphetis gracilis Moore, 1963: 288; Moore et al„ 1987: 153.

REMARKS. Occurs in five populations with
slightly different ranges, and for the present these
are classified as subspecies. For distinctions see
key.

For taxonomic problems see ‘Taxonomic
Principles’ and ‘Remarks’.

DIAGNOSIS. Relatively large, easily distinguished
by the combination of following character states:
extremely elongate mandibles, depressed eyes,
distinctly cordate prothorax, weak striation of
elytra, evenly but not suddenly curved aedeagus,
and rather elongate parameres the right one
bearing setae only in apical part of lower rim.

Raphetis gracilis gracilis Moore
(Figs 2E, 13B, 14A, 17)

MATERIAL. HOLOTYPE: d, Millaa Millaa, Q.
Atherton Tab. Apr. 1 ‘32. 2500 ft/ Australia, Harvard Exp.,
Darlington' Raphetis gen.n. gracilis sp.n. holotype 6 Det.
B.P. Moore ‘63 (AN1C).

NEW RECORDS. 2 c5, NEQ. 17°33‘S 145°33’E Mt
Fisher, l Akm NW 8.ii. 1 999. 1 280m. R/F. GM. Pyr. - trees &
logs. 2 1 70 (CBM, QM. ); 1 <J,2 ? , Mt Fisher, 1 050- 1 1 00m
7km SW Millaa Millaa. N Qld 27-29 Apr., 1982 GM, DY
& DC (QM); 2 d . Millaa Millaa Falls, via Millaa Millaa, N
Qld 12 Aug. 1968 B. Cantrell (QM).

DIAGNOSIS. Medium sized subspecies with
unarmed elytra and narrow, comparatively
elongate, rather cordiform pronotum.
Distinguished from the northern subspecies (R. g.
spurgeoni , R. g. subarmata , R. g. spinosa) by
narrower prothorax and narrower, on lower side
slightly downcurved apex of elytra. Distinguished
from most closely related R. g. frerei by lesser
size, slightly narrower pronotum and con-
siderably longer elytra.

REDESCRIPTION. Measurements. Length:
7.0-7. 8mm; width: 2. 65-2. 90mm. Length eye/
orbit: l. 8-2.0; width/length of pronotum: 1.00-
1.03; width base/apex of pronotum: 1.08-1.11;
width pronotum/head: 1 .35-1 .41 ; length/width of
elytra: 1.49-1.52; width elytra/pronotum:
1.49-1.55.

Colour Black. Clypeus, labrum, and mandibles
reddish-piceous, palpi and antennae reddish.
Legs reddish. Lower surface of anterior body
dark piceous, abdomen reddish-piceous.

Head. Considerably narrower than pronotum.
Eyes comparatively small (in genus), little
convex, laterally little protruding. Orbits
elongate, oblique, straight. Eyes separated from
frons by a narrow, straight furrow. Frons with
rather deep, irregularly impressed, curved
furrows medially of eyes that are prolonged to
clypeal suture. Furrows medially widened to
shallow, irregular grooves, at posterior end
laterally bordered by convex ridge. Anterior
supraorbital seta situated close to eye near
anterior border of eye, posterior seta situated far
behind eye and rather moved to upper part of

PSYDRINE GROUND BEETLES

99

head. Neck separated by a shallow, transverse
furrow. Clypeal suture distinct. Labrum
anteriorly rather deeply concave, 6-setose and
with some shorter hairs around the anterior
angles. Mandibles very elongate, straight, inner
margin straight for a long distance, apex little
curved, with elongate seta in scrobe. Mentum
with acute triangular tooth. Submentum bisetose,
gula quadrisetose, all setae very elongate. Glossa
short, rather narrow, bisetose, paraglossae
hyaline, surpassing glossa. Lacinia very
elongate, inner margin with rather few strong
spines, apex markedly incurved, very acute. Both
palpi elongate, glabrous, apex transverse.
Antenna rather slender and elongate, barely
surpassing base of pronotum, pilose from apical
half of 4th antennomere, central antennomeres c.
2.5 x as long as wide. Surface of head absolutely
smooth, without any indication of micro-
sculpture and puncturation, remarkably glossy.

Pronotum (Fig. 14A). Comparatively narrow,
moderately convex, rather cordiform. Apex
gently concave, anterior angles slightly
produced, angulate, sides evenly curved, widest
at middle, sinuate and almost parallel in basal
fifth. Basal angles angulate, rectangular. Base
laterally slightly oblique. Apex and base not
margined. Lateral channel narrow, barely
explanate near base. Median line distinct, slightly
impressed, neither reaching apex nor base. Basal
grooves straight, rather linear, fairly deep. Both
transverse sulci barely indicated. Anterior lateral
seta situated slightly in front of middle, posterior
lateral seta absent. Surface absolutely smooth,
without any indication of microsculpture or
puncturation, though near median line with some
short, transverse furrows, remarkably glossy.

Elytra. Comparatively elongate, convex, con-
siderably wider than pronotum, rather parallel,
wide at humeri. Humeri slightly angulate, lateral
margins feebly convex, elytra widest at anterior
third or near middle, posteriorly evenly
narrowed. Lateral part of apex slightly sinuate at
position of crossing of epipleurae, then evenly
convex. Apex evenly rounded off. Basal margin
complete, lateral margin narrow throughout.
Scutellary puncture and seta present, at base of
1st stria. Scutellary stria not perceptible. Striae
extremely fine, barely perceptible. External striae
not recognisable. Striae very finely punctate. 8th
stria complete, coarsely punctate, though barely
impressed. Near apex only 1st and 7th striae well
impressed, 7th stria forming an elongate, fairly
deep furrow. Intervals absolutely depressed. Disk
impunctate. Marginal series consisting of two

groups of 7-8 and 6-7 setiferous punctures,
respectively, which are widely separated in
middle. Inside of deepened 7th stria with two
additional punctures near apex. Some of the
marginal setae very elongate. Intervals
absolutely smooth, without any traces of
microreticulation, highly glossy.

Lower surface . Metepisternum rather short,
about 1.25 x as long as wide at anterior margin,
considerably narrowed behind. Lower surface
impunctate. Terminal stemite in male bisetose, in
female quadrisetose along margin, and with two
additional shorter setae in middle slightly
removed from margin.

Legs. Elongate. 5th tarsomere asetose beneath.
Anterior tarsus in male barely wider than in
female, lst-3rd tarsomeres asymmetrically and
sparsely squamosc beneath.

Male genitalia (Fig. 13B). Genital ring short and
wide, asymmetrically triangular, with short,
rounded apex. Aedeagus rather elongate (in
genus), laterally depressed, lower surface almost
evenly concave, slightly sinuate just in front of
apex. Apex short, very wide, convex, though on
lower side slightly angulate, slightly turned.
Internal sac rather complexly folded, with several
sclerotised plates within. Both parameres
comparatively elongate and narrow, left larger
than right, triangular, with obtusely triangular
apex, and 4-5 apical setae, without additional
setae on lower margin. Right paramere with acute
apex and c. 10 elongate setae along the apical
third of lower margin.

Female genitalia (Fig. 2E). Stylomere 1 with 3
ensiform setae at middle ofventro-apical margin,
sometimes with an additional nematiform seta.
Stylomere 2 moderate, with relatively short apex
and 2 large dentiform ventro-lateral ensiform
setae the upper one of which is considerably
larger in middle of lateral margin. Near apex with
a large, oblong pit and a short nematiform seta
originating from that pit. In middle of
dorso-median surface with a large, dentiform,
dorso-median ensiform seta. Lateral plate with a
densely setose area at median apical margin.
Variation. Slight differences in relative shape of
pronotum and elytra.

DISTRIBUTION (Fig. 17). Millaa Millaa/Mt
Fisher area in the western part of Atherton
Tableland, northeastern Queensland.

COLLECTING CIRCUMSTANCES. Taken by
pyrethrum knockdown on trees and logs in
upland rainforest.

100

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 1 4. Pronota, elytra and elytral apices of Raphetis spp. A, R. gracilis
gracilis Moore; B, R. gracilis frerei subsp. nov.; C,E, R. curta sp. nov.;
D,F,I, R. gracilis spurgeoni subsp. nov.; G, R. gracilis spinosa subsp.
nov.; H, R. gracilis subarmata subsp. nov.

Raphetis gracilis frerei subsp. nov.

(Figs 13D, 14B, 15B,E, 17)

ETYMOLOGY. Refers to the distribution on Mt Bartle
Frere.

DESCRIPTION. Measurements.
Length: 7.9- 8.5mm; width:
3.1 -3.3mm. Length eye/ orbit:
1.7-1. 9; width/length of pro-
notum: 1.04- 1.07; width base/
apex of pronotum: 1.10-1.13
width pronotum/hcad: 1.46-1.51
length/width of elytra: 1.42-1.47
width elytra/pronotum: 1.54- 1.58.
Colour. As in nominate
subspecies.

Head. As in nominate subspecies,
though eyes even more depressed.
Pronotum (Figs 14B, 15B). As in
nominate subspecies, though
pronotum, at the average, slightly
wider and with wider base.

Elytra. As in nominate subspecies,
though slightly shorter and wider,
on average laterally slightly more
convex. Inner striae somewhat
variable, either barely indicated as
a row of extremely fine punctures,
or even very gently impressed.
Lower surface. As in nominate
subspecies.

Legs. As in nominate subspecies.
Male genitalia (Fig. 13D). Much
as in nominate subspecies, but aedeagus slightly
longer and both parameres slightly narrower.

Female genitalia. As in nominate subspecies.

Variation. Some minor variation in relative shape
of pronotum and elytra, and in distinctness of the
inner elytral striae.

MATERIAL. HOLOTYPE: d, QMT21872, NEQ.
17^16’S J45°49’E Massey Ra., 4km W of Bellenden Ker,
1 250m 1 0.x. 1991. Pyrethrum, Monteith & Janetzki (QM).
PARATYPES: 2 d, 1 9, same data (QM, CBM); 1 d,
same locality, 1 1.x. 1991 (QM); 2 9, same locality,
9-1 1.x. 1991 (QM); 2 d, 1 9, Mt Bartle Frere. N Qld.
Central Ridge. 1500m 27.xii.1989. GM Pyr. Logs (CBM,
QM); 1 9, Bellenden Ker Range, NQ Cable Tower 3,
1054m 17 Oct.-5 Nov. 1981 Earthwatch&QM (QM).

DISTRIBUTION (Fig. 17). Mt Bellenden Ker,
Mt Bartle Frere, and Massey Range at the eastern
margin of Atherton Tableland, North Queens-
land.

COLLECTING CIRCUMSTANCES. Taken by
pyrethrum knockdown in upland rainforest on
logs. Collected so far between 1050- 1500m.

DIAGNOSIS. Large subspecies with unarmed
elytra and comparatively narrow and elongate,
rather cordiform pronotum. Distinguished from
the northern subspecies (R. g. sputgeonL R. g.
subarmata , R. g. spinosa) by larger size,
narrower prothorax, and narrower, on lower side
slightly downcurved apex of elytra.
Distinguished from most closely related R. g.
gracilis by larger size, slightly wider pronotum
and considerably shorter elytra.

Raphetis gracilis spurgeoni subsp. nov.
(Figs 13E, 14D,E,I, 15C,F, 17)

ETYMOLOGY. Refers to the type locality near Mt
Spurgeon.

MATERIAL. HOLOTYPE: d, QMT21876, NE.Q:
16°22’S 145°13’E 7km N Mt Spurgeon (Camp
2), 17- 19.x. 1991, 1200- 1250m, Monteith, Janetzki, Cook
& Roberts (QM). PARATYPES: 2 d, 1 9, same data
(CBM, QM).

PSYDRINE GROUND BEETLES

101

DIAGNOSIS. Small subspecies with unarmed
elytra and wide, short, moderately cordiform
pronotum. Distinguished from both southern
subspecies (R. g. gracilis , R. g. frerei ) by wider
prothorax and wider, on lower side evenly curved
apex of elytra. Distinguished from most closely
related R. g. subarmata and R. g. spinosa by more
cordiform pronotum with comparatively
narrower base, and by absolute lack of any spine
or denticle at apex of elytra.

DESCRIPTION. Measurements. Length: 6.6-
7.0mm; width: 2.55-2.7mms. Length eye/orbit:
2. 1-2.3; width/length of pronotum: 1.12-1.16;
width base/apex of pronotum: 1.10-1.15; width
pronotum/head: 1.45-1.50; length/width of
elytra: 1.48-1.50; width elytra/pronotum: 1.42-
1.45.

Colour. As in nominate subspecies.

Head. As in nominate subspecies, though eyes
relatively convex.

Pronotum (Figs 14D, 15C). As in nominate
subspecies, though pronotum wide with fairly
narrow base, hence rather cordiform.

Elytra (Figs 14E, 141). As in nominate
subspecies, but with wider, on lower side, evenly
curved apex, and with inner elytral striae
invariably very gently though perceptibly
impressed, though barely punctate. Hence, inner
intervals faintly convex.

Lower surface. As in nominate subspecies.

Legs. As in nominate subspecies.

Male genitalia (Fig. 13E). As in nominate
subspecies, but aedeagus less curved, apex larger
and evenly rounded on lower border, and left
paramere with only 3 setae at apex.

Female genitalia. As in nominate subspecies.
Variation. Slight variation noted in relative shape
of eyes, prothorax, and elytra.

DISTRIBUTION (Fig. 17). North of Mt
Spurgeon near the western edge of the Carbine
Tableland, North Queensland.

COLLECTING CIRCUMSTANCES. Taken by
pyrethrum knockdown on logs and tree trunks in
upland rainforest above 1200m.

Raphetis gracilis subarmata subsp. nov.
(Figs 13F, 14H, 17)

ETYMOLOGY. Refers to the denticle at the apex of the
elytra.

MATERIAL. HOLOTYPE: <J, QMT21866, Carbine
Tableland, NE Qld Plane Crash Site 27-28 Nov 1990,

1330m, Monteith, Thompson, Cook, Sheridan & Janetzki,
Pitfall Traps & Hand (QM). PAR ATYPES: 1 6 , 1 9 , same
data (CBM): 1 6 \ 16°24’S 145°13’E Stewart Ck, 4km
NNE. Mt Spurgeon (Camp 1), 1250-1300m,
15-20.X.1991. GM, HJ, DC & LR (QM); 1 <J, 16°24’S
145°12’E Black Mtn. 4.5km N of Mt Spuigeon, 1250-
1330m, 17-18^.1991. GM, HJ, DC & LR (QM).

DIAGNOSIS. Medium sized subspecies with
slightly denticulate elytra and moderately wide
and short pronotum. Distinguished from both
southern subspecies (R. g. gracilis , R. g. frerei) by
wider prothorax with wider base, and by wider,
on lower side evenly curved apex of elytra.
Distinguished from R. g. spurgeoni by less
cordiform prothorax and presence of a denticle at
the apex of the elytra; and from R. g. spinosa by
narrower, slightly more cordiform pronotum, and
by denticulate rather than spinose apex of elytra.

DESCRIPTION. Measurements. Length: 6.7-
7.9mm; width: 2.65-3. 1mm. Length eye/ orbit:
1. 8-2.0; width/length of pronotum: 1.08- 1.10;
width base/apex of pronotum: 1.14-1.18; width
pronotum/head: 1,47-1.52; length/width of
elytra: 1.45-1.48; width elytra/pronotum:
1.50-1.58.

Colour. As in nominate subspecies.

Head. As in nominate subspecies.

Pronotum. As in nominate subspecies, though
base rather wide.

Elytra (Fig. 14H). As in nominate subspecies,
though elytra rather short, laterally rather convex,
and with a tiny sharply angulate or even
denticulate at apex. Inner striae invariably gently
though perceptibly impressed, though barely
punctate. Inner intervals faintly convex.

Lower surface. As in nominate subspecies.

Legs. As in nominate subspecies.

Male genitalia (Fig. 13F). As in nominate
subspecies, but aedeagus more curved, apex
larger and evenly rounded on lower border, and
left paramere with only 2-3 setae at apex.

Female genitalia. As in nominate subspecies.
Variation. Some slight variation noted in size and
in relative shape of eyes, prothorax, and elytra.

DISTRIBUTION (Fig. 17). Mt Spurgeon and
Plane Crash Site at the western and eastern
margins, respectively, of the Carbine Tableland,
North Queensland.

COLLECTING CIRCUMSTANCES. Specimens
from Mt Spurgeon were collected by pyrethrum
knockdown in upland rainforest, those from
Plane Crash Site by pitfall trapping and by hand

102

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1 5. Head and pronotum of Raphetis. A, Raphetis curta sp. nov.; B, R. gracilis frerei subsp. nov.; C, R. gracilis
spurgeoni subsp. nov.; habitus of Raphetis. D, Raphetis curta sp. nov. E, R . gracilis frerei subsp. nov. F, R.
gracilis spurgeoni subsp. nov.; lengths: 5.3mm; 8.3mm; 7.0mm.

collecting. Captured only above 1250m which is
at maximum altitude on these mountains.

Raphetis gracilis spinosa subsp. nov.

(Figs 13C, 14G, 17)

ETYMOLOGY. Refers to the spinose elytral apices.

MATERIAL. HOLOTYPE: <J, QMT21858, Mossman
Bluff 10km W Mossman, N Qld. 1 7-1 8.xii. 1988.
1100-1300m, Monteith & Thompson (QM).
PARATYPES: 2 <5, NEQ: 16°30 > Sxl45°16’E Hilltop,
18km N Mt Lewis, 23.xi.1998. GM Pyrethrum, trees.
1300m Rf. (CBM); 1 d, 2km SE Mt Spuigeon via Mt
Carbine. N Qld. 20-2 1 .xii. 1 988. 1 1 00m, GM & GT (QM);
3 Mt Lewis Rd end 10km N Mt Lewis 25.xi.1990,
1100m, GM, RS & GT. Pyrethrum, Logs (QM).

DIAGNOSIS. Medium sized subspecies with
spinose elytra and moderately wide and short
pronotum with very wide base. Distinguished

from both southern subspecies ( R . g. gracilis , R.
g. frerei) by wider prothorax with wider base, and
by wider, on lower side evenly curved apex of
elytra. Distinguished from both closely related R.
g. spurgeoni and R. g. subarmata by less
cordiform prothorax, remarkably spinose apex of
the elytra, and shorter and stouter aedeagus;
further on from R. g. spurgeoni by wider base of
pronotum, and from R. g. subarmata by wider
pronotum.

DESCRIPTION. Measurements. Length: 6.95-
8.0mm; width: 2.6-3.2mm. Length eye/ orbit:
2.0-2. 1; width/length of pronotum: 1.11- 1.16;
width base/apex of pronotum: 1.18-1.22; width
pronotum/head: 1.50-1.55; length/width of
elytra: 1.48-1.56; width elytra/pronotum:
1.42-1.44.

Colour. As in nominate subspecies.

PSYDRINE GROUND BEETLES

103

Head. As in nominate subspecies.

Pronotum. As in nominate subspecies, though
pronotum very wide and with wide base.

Elytra (Fig. 14G). As in nominate subspecies,
though elytra narrow in comparison to pronotum.
Apex with conspicuous hook-shaped spine that is
slightly curved outwards. Inner striae very gently
impressed, barely punctate. Inner intervals not or
extremely faintly convex.

Lower surface. As in nominate subspecies.

Legs. As in nominate subspecies.

Male genitalia (Fig. 13C). As in nominate
subspecies, but aedeagus shorter and stouter,
more evenly curved, apex larger and evenly
rounded on lower border, and left paramere with
only 2 setae at apex.

Female genitalia. As in nominate subspecies.
Variation. Some differences in relative shape of
pronotum and elytra, and development of elytra
striae.

DISTRIBUTION (Fig. 17). Mt Lewis, Mossman
Bluff and Mt Spurgeon, Carbine Tableland, North
Queensland.

COLLECTING CIRCUMSTANCES. Taken by
pyrethrum knockdown in upland rainforest on
trees and logs. Collected only above 1 100m.

MEASUREMENTS AND RATIOS IN
RAPHETIS MOORE

For better comparison of the species the
measurements and ratios of all species and
subspecies are compiled in Table 3.

REMARKS ON THE RAPHETIS GRACILIS
COMPLEX

As mentioned in the introduction, the five
different populations of R. gracilis are classified
here as subspecies. At first glance, however, at
least R. gracilis spinosa might be regarded a
separate species because of its conspicuously
spinose elytra and its wide pronotum with
comparatively wide base. Specimens of this
population also look rather different in general
shape from other populations. However, the male
genitalia of R. gracilis spinosa are not
perceptibly different from those of the other
populations of R. gracilis from the Carbine
Tableland. Measurements and ratios of all five
populations also widely overlap. Because of this
complexity it is difficult to make reliable
distinctions between populations in the present
state of knowledge. On the other hand, these

beetles live in rain forest at rather high altitude
and a lowland barrier that is today free from
rainforest exists between at least the southern
populations (R. g. gracilis and R. g. frerei ) and
the northern populations (R. g. spinosa , R. g.
spurgeoni and R. g. subarmata). That these two
groups are sufficiently different to be classified as
separate taxa, is also indicated by the shape of the
aedeagus that is slightly sinuate subapically at its
lower margin in the southern group, and straight
in the northern group, and by the structure of The
internal sac. This particular lowland barrier is
known as the Black Mountain Barrier and has
been shown to be effective in several vertebrate
and invertebrate groups (Schneider et al. 1998).

It is unknown how substantial the barriers were
before the arrival of European settlers in north
Queensland after which the tablelands between
the ranges were extensively deforested. Though
at least some climatic differences must have
existed, even in pre-human times, that probably
prevented interbreeding between the populations.

Hence, at least the southern and northern
groups have been taxonomically separated for an
unknown period. Within the northern group (R. g.
spinosa , R. g. subarmata , and R. g. spurgeoni )
the situation is somewhat different. All three
populations live in rather close proximity, and it
is even conceivable that they overlap in some
areas. Moreover, the environment on the Carbine
Tableland is much more homogenous today than
it is on the Atherton Tableland, and this may have
also been the case in pre-human times . It is
conceivable, then, that these populations have
differentiated rather recently, and may still be in
the course of differentiation with some inter-
breeding still occurring.

PHYLOGENY AND BIOGEOGRAPHY OF
RA PHETIS. Because of comprehensive sampling
by the Queensland Museum of rainforested
mountains in Queensland the actual distribution
of Raphetis is probably rather reliably known
Hence, some considerations about phylogenetic
relations are possible, even though new taxa may
be discovered in future.

Within the Psydrinae in general and with
respect to the closely related genus Meonis in
particular, the following character states in
Raphetis seem to be apomorphic: 1 ) reduction of
elytral striation; 2) lengthening of head; 3)
lengthening of mandibles; 4) reduction of eye
size in comparison to size of orbit; 5) flattening of
eyes; 6) reduction of microreticulation to get a

104

MEMOIRS OF THE QUEENSLAND MUSEUM

highly glossy surface; 7) reduction of pilosity on
lower margin of right paramere.

These character states apply for the whole
genus. But within the genus there are also strong
clinal developments within most of the same
characters.

When applied to the species, R. darlingtoni and
R . curta are seen to be plesiomorphic in almost all
characters when compared with R. gracilis. Only
in the reduced elytral striation is R. curta
apomorphic compared to R . darlingtoni , though
in characters states of head, mandible, and eye
size, R. curta is more plesiomorphic than R.
darlingtoni. Certainly, it is not possible to draw' a
cladogram from the few characters used as they
would not yield information about any
synapomorphies between the southern species
(R. curta , R. darlingtoni) and the northern
gracilis- complex.

However, when this rough phylogenetic
statement is combined with the distribution
pattern of the species, it clearly reveals a clinal
increase in apomorphy from south to north in
several characters. Thus, the most southern
species, R. curta , is most plesiomorphic in many
respects, with R. darlingtoni a little more

FIG 1 7. Distribution of subspecies of Raphetis gracilis
Moore in northern Queensland.

evolved, whereas R. gracilis of north Queensland
is apomorphic in almost all character states
discussed. Within the gracilis- complex some
minor further clinal development is seen, with a
south-to-north gradient also obvious in several
character states. R. g. gracilis from Atherton
Tableland, for instance, has the plesiomorphic
condition of relatively large eyes and unarmed
elytra. R. g.frerei possesses very depressed eyes,
and R. g. spinosa has relatively convex eyes but
armed elytra. The last two subspecies occur
either further north on Carbine Tableland, or on
the ranges at the eastern margin of Atherton
Tableland.

As noted earlier, the relationships of R. g.
subarmata and R. g. spurgeoni are more difficult
to explain. With respect to eye size and
development of spined elytra, there seems to
exist a cline, but in the opposite direction to the
southern populations, with the most plesiotypic
R. g. spurgeoni in the north, and the most
apotypic R. g. spinosa in the south. R. g.
subarmata is intermediate in many respects.

The above distribution pattern is a scenario that
has proved to be common in ground living
carabids (and other taxa) of the montane rain
forests of northern Queensland (e.g. Baehr,

PSYDRINE GROUND BEETLES

105

TABLE 3. Raphetis spp. measurements.

N

body length
(mm)

ratio length
eye/orbit

ratio width/
length
pronotum

ratio width
base/apex
pronotum

ratio width
pronotum/
head

ratio length/
width elytra

ratio width
elytra/
pronotum

R. curta

3

5.3-5. 7

2.3-2.4

1.20-1.23

1.34-1.40

1.51-1.58

1.31-1.35

1.36-1.43

R. darlingtoni

1

6.1

3.2

1.17

1.15

1.55

1.37

1.35

R. gracilis gracilis

8

7.0-7.8

1. 8-2.0

1.00-1.03

1.08-1.11

1.35-1.41

1.49-1.52

1.49-1.55

R. gracilis frerei

10

7.9-8. 5

1.7-1. 9

1.04-1.07

1.10-1.13

1.46-1.51

1.42-1.47

1.54-1.58

R. gracilis spurgeoni

4

6.6-7.0

2. 1-2.3

1.12-1.16

1.10-1.15

1.45-1.50

1.48-1.50

1.42-1.45

R. gracilis subarmata

5

6.7-7.9

1. 8-2.0

1.08-1.10

1.14-1.18

1.47-1.52

1.45-1.48

1.50-1.58

R. gracilis spinosa

7

6.9-8.0

2.0-2. 1

1.11-1.16

1.18-1.22

1.50-1.55

1.48-1.56

1.42-1.44

1 995b; genus Sitaphe , see above). In several low
vagility groups the species occurring on Atherton
Tableland not only have a wide range but are
usually also rather plesiotypic compared with
those from Bellenden Ker/Bartle Frere ranges.
Carbine Tableland and mountains tops further
north. This pattern is believed to have been
caused by vicariance through separation of
populations at the eastern and northern borders of
a previous larger tableland following its
dissection by deep river valleys and due to retreat
of the rainforests to the mountain tops. Both
events were caused by erosion of the uplifted
tablelands into montane blocks and, at the same
time, by repeated spreading and retreat of the rain
forests during the stadials and interstadials of
glaciation period.

Provided no contrary evidence emerges in
future, this would mean that Raphetis probably
originated somewhere in the subtropical or cool
temperate rain forests of southern Queensland
and later spread northwards to reach eventually
the montane tropical rain forests of northern
Queensland. There it passed through a con-
siderable taxonomic radiation, presumably rather
recently and probably still continuing. This
scenario also agrees well with considerations
about the origin of the Australian Psydrinae in
general that will be developed in my forthcoming
revision of the Amblytelinae (Baehr, in press).

GENERAL DISCUSSION

HABITS AND LIFE HISTORIES. Although
labels do not always give information about the
habits and habitats of the North Queensland
psydrines, fortunately many specimens of all
three genera were collected by hand. The
following notes on their ecology are based on
details kindly contributed by the main collector,
Geoff Monteith, as summarised below.
Regarding all three genera, the first

generalisation that can be made is that all
specimens have been collected in rainforest at
medium to high altitudes and none have been
taken in adjacent eucalypt-dominated open
forests. Secondly, all hand-collected active
individuals have been taken at night during
headlight searching, so we can assume that all are
nocturnal in activity.

The small species of Mecyclothorax live in leaf
litter of upland rain forest, and are collected either
by Berlese extraction of litter or by hand
searching on the ground with a headlight at night.
A few specimens may move a short distance up
the mossy surface of tree trunks where they are
occasionally sampled by pyrethrum knockdown
of that situation. They appear to forage among
litter at night on the ground.

Both Sitaphe and Raphetis also live in upland
rain forest, generally at very high altitude and
most commonly at the absolute summits of the
ranges. Rarely, they are taken in leaf litter, and
their primary habitat seems to be the surface or
even crevices of bark or exposed dead wood.
They rarely occur on living trees, but usually on
dead trees and logs where there are many cracks
and crevices on the surface. Specimens of Sitaphe
usually rest in depressions and grooves on the
wood or bark surface where their strong convex
elytra protects them from attack. In the daytime
they are usually found resting in depressions on
the underside of logs and smaller pieces of wood
lying on the ground. At night they run over the
surface of dead wood and then they can be
detected while hunting with a headlight.

Specimens of Raphetis are much rarer and are
almost never found by hand-collecting in the
daytime. They probably move into tunnels and
chambers inside the dead wood in the daytime,
and their more slender form also suggests such
behaviour. Pyrethrum forces the beetles out from
this situation when the tree trunks and log

106

MEMOIRS OF THE QUEENSLAND MUSEUM

surfaces are fogged in the daytime. At night they
forage on the surface of the dead wood and almost
all the hand-collected specimens are taken at
night when they are doing this, sometimes on logs
and sometimes on standing dead tree trunks.

So the North Queensland psydrines fall into
three rather different habitat categories: Mecyclo-
thorax live in leaf litter on the ground; Sitaphe
and Raphetis forage on the surface of dead wood
with Raphetis retreating inside the wood in the
daytime while Sitaphe conceals itself in crevices
on the surface.

In spite of the above information about habits
of adults, nothing is known about life histories of
any species, especially on their larvae which are
so far unknown.

Although the strange looking body shape of
Sitaphe and Raphetis , especially the narrow head
and the rather elongate, porrect mandibles in both
genera, indicates a specialised feeding method,
there are no observations about diet or feeding
habits of any Sitaphe or Raphetis species. We
only can speculate about feeding and we may
argue that the cychroid structure of head and
mandibles could indicate that they eat snails. The
very smooth, glossy surface in both genera may
also corroborate this opinion.

For the northern species of Mecyclothorax
information about diet is also lacking. Given their
small size feeding on small insects or worms may
be most probable, but no observations are
available.

To confirm speculations about phylogenetic
relations, better knowledge about ecology and
ethology, and especially about life history, would
be helpful.

BIOGEOGRAPHICAL CONSIDERATIONS.
In Sitaphe , Raphetis and the northern species
group of Mecyclothorax , very similar patterns of
distribution can be noted. In the three genera all
taxa are very closely related and most are
restricted to very small ranges that repeatedly
cover a single rain forest block (in the sense of
Baehr 1 995b) or even a single mountain top. This
is a general pattern common in flightless
invertebrates in the montane rain forests of the
Wet Tropics of northern Queensland (Baehr,
1995b, Monteith, 1997, Davies & Lambkin,
2000, Harvey, 2000). In Sitaphe , however, one
species occupies a fairly large range on the
Atherton Tableland, which again is a common
distribution pattern in the Wet Tropics.

At the first glance, the high grade of

morphological similarity of the many taxa should
be evidence of a very recent taxonomic radiation
of rather old stocks in all three mentioned genera.
There has been, however, some discussion in
recent papers about the age of the many closely
related invertebrate taxa occurring in the Wet
Tropics, especially with regard to the montane
species (e.g. Moritz et al., 2000; Russell et al., in
press). In some papers evidence has been
presented, mainly from molecular phylogenetic
analyses (expressed in the percentage of
sequence divergence of 16S rRNA), suggesting
the main speciation events to be of late Tertiary
age, in Pliocene or even late Miocene.

However, it seems to me rather audacious to
translate such percentages of sequence
divergences directly into an absolute time table,
because little is known about different rates of
development of molecular divergences under
different or even rapidly changing environmental
conditions. External morphological differences,
at least can develop very rapidly when ecological
factors are changing.

Hence, as a conclusion, and because molecular
data arc still lacking for the carabid groups
mentioned herein, the high grade of morph-
ological similarity on the background of high
species diversity and a high level of endemism
are still evidence of quite recent — that is
Pleistocene — speciation events that most
probably proceeded by allopatric speciation
caused mainly by vicariance events, which
apparently was a common means of evolution of
diversity in tropical rain forest (Joseph et al.,
1995).

During the last years when much more
scrutinised collecting work was conducted in a
multitude of montane rainforests of eastern and
northern Queensland, a number of unexpected
species of southern origin were recorded far
north of the range of their relatives. Examples
from ground beetles are two species of the
merizodine genus Sloaneana in southern
Queensland (Baehr, 2002b), or the occurrence of
a Tasmanitachoides from a decidedly southerly
species-group on the top of a rainforested
mountain in North Queensland (Baehr, 2001 ), or
the occurrence of about 40 species of the psydrine
genera Amblytelus and Dystrichothorax in
eastern and northeastern Queensland (Baehr, in
press), or even the discovery of an — as yet
undescribed — migadopine species in northern
Queensland (G. Monteith, pers. com.), and,
obviously, also the numerous species of

PSYDRINE GROUND BEETLES

107

Mecyclothorax , Sitaphe, and Raphetis occurring
in the montane rain forests of northeastern
Queensland.

It should be remembered, however, that the
ranges of certain decidedly northern genera also
have been extended to the south due to recent
sampling; as one example the first discovery of a
species of northern Philipis in northern New
South Wales (Baehr, 2002a) should be noted.

These new discoveries show that the traditional
biogeographical division of the Australian biota
is even less rigid than it was believed so far. In
particular the geographic border between the
so-called Bassian and Torresian subregions in
eastern Australia that has been roughly estimated
to follow about the Queensland/New South
Wales border, turns out more and more to be a
fiction. Even the montane regions of North
Queensland include more species that originally
stem from cool- or even cold-adapted southern
groups, than true Torresian faunal elements. If we
accept that there is a clear-cut border at all, then
this border is — so to speak — rather a horizontal
one that extends over a very long distance along
the east coast from northern or even mid-New
South Wales up to northeastern Queensland. It
lies between the warmer and, in parts, drier
lowlands and lower reaches of the mountains and
the cooler and mostly wetter tablelands and tops
of the ranges. Whereas the rain forests, swamps,
ponds and large rivers of the lowlands possess a
true Torresian fauna, the carabid fauna of the
cooler montane tropical rain forests, Nothofagus
forests, fern bogs, and cold streams likely could
be attributed to the Bassian subregion. This
Bassian faunal element along the mountain tops
of eastern Australia superimposes the Torresian
fauna, and it even appears again in the
Nothofagus forests and bogs of the highlands of
New Guinea.

It has been believed that the connection of the
Australian block to the southeast Asian insular
belt during Miocene not only resulted in the
invasion of numerous Oriental species into
Australia, but also that these immigrants
generally were superior over the native fauna and
pushed it back as they advanced to the south (e.g.
Darlington, 1968). Certainly, this picture is not
right, or, at least, it does not apply for the montane
environments of eastern Queensland. On the
contrary, it seems that at least the Pleistocene
glacial period supported the evolution and
taxonomic radiation of Bassian faunal elements

even in the montane rainforests of North
Queensland.

ACKNOWLEDGEMENTS

I am most indebted to Dr G.B. Monteith,
Brisbane, for the generous loan of the material,
for preparing the distribution maps, for revising
English expression in the MS, and for kindly
adding much important information about
collecting circumstances and habits of various
species. Thanks are also due to Mr T.A. Weir,
Canberra, for the kind loan of the types of the
described species.

LITERATURE CITED

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1998. A further new species of the genus
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Spixiana 21: 21-24.

1999. A preliminary survey of the classification of
the Psydrinae (Coleoptera: Carabidae). In, Ball,
G.E.,A. Casale & A. Vigna Taglianti (eds)
Phylogeny and classification of Caraboidea. Atti
di Museo regionale di Scienze naturali de Torino
1998: 359-368.

2000. A new species of the genus Lestignathus
Erichson from Tasmania. With a note on the
Tasmanian species of Mecyclothorax Sharp
(Insecta: Coleoptera: Carabidae: Licininae,
Psydrinae). Records of the South Australian
Museum 33: 123-126.

200 1 . Tasmanitachoides Erwin glabellus n. sp. from
North Queensland, Australia, with a note on
Tasmanitachoides lutus (Darlington) (Insecta,
Coleoptera, Carabidae, Bembidiinac). Animal
Biodiversity and Conservation 24: 1-7.

2002a. New species and additional records of the
tachyine genus Philipis Erwin (Coleoptera:
Carabidae: Bembidiinae). Memoirs of the
Queensland Museum 48: 1-7.

2002b. Two new species of Sloaneana Csiki from
southern Queensland (Coleoptera: Carabidae:
Merizodinae). Memoirs of the Queensland
Museum 48: 9-15.

2000c. Two new species of the genus
Mecyclothorax Sharp from Papua New Guinea

108

MEMOIRS OF THE QUEENSLAND MUSEUM

(Insecta, Coleoptera, Carabidae, Psydrinae).
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In press. The Amblytelini. A tribe of corticolous
ground beetles from Australia. Taxonomy,
phylogeny, biogeography (Insecta, Coleoptera,
Carabidae, Psydrinae). Coleoptera.

BAEHR, M. & LORENZ, W. 1999. A reevaluation of
Loeffleria globicollis Mandl from Borneo
(Insecta, Coleoptera, Carabidae, Psydrinae).
Spixiana 22: 263-267.

BLACKBURN, T. 1889. Further notes on Australian
Coleoptera, with descriptions of new genera and
species. Proceedings of the Linnean Society of
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BOUCHARD. P. 2002. Phylogenetic revision of the
flightless Australian genus Apterotheca Gebien
(Coleoptera: Tcnebrionidae: Coelometopinae).
Invertebrate Systcmatics 16: 449-554.
BRANCUCCI, M. & MONTEITH, G.B. 1996. A
second Terradessus species from Australia
(Coleoptera, Dytiscidae). Entomologica
Basiliensia 19: 585-591.

BRITTON, E.B. 1948. A revision of the Hawaiian
species of Mecyclothorax (Coleoptera:
Carabidae). Occasional Papers of the Bishop
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CASTELNAU, LAPORTE DE, F.L. 1867. Notes on
Australian Coleoptera. (Royal Society of Victoria:
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1 868. Notes on Australian Coleoptera. Transactions
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Coleopterorum Catalogus II, 104: 347-528.
DARLINGTON, P.J. Jr 1961a. Australian carabid
beetles VII. Trichosternus , especially the tropical
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1961b. Australian carabid beetles IX. The tropical
Notonomus. Breviora, Museum of Comparative
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1962. The carabid beetles of New Guinea. Part I.
Cicindelinae, Carabinae, Harpalinae through
Pterostichini. Bulletin of the Museum of
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1971. The carabid beetles of New Guinea. Part IV.
General considerations; analysis and history of
fauna; taxonomic supplement. Bulletin of the
Museum of Comparative Zoologv 142: 129-337.
DAVIES, V.T. & LAMBKIN, C. 2000. Wabua, a new
spider genus (Araneae: Amaurobioidea:
Kababininae) from north Queensland, Australia.
Memoirs of the Queensland Museum 46:
129-147.

DEUVE, T. 1987. Descriptions de deux carabiques
nouveaux de Nouvelle-Caledonie et de Thailande
(Coleoptera, Caraboidea, Psydridae, Trechidae).
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9: 143-146.

HARVEY, M.S. 2000. A review of the Australian
schizomid genus Notozomus (Hubbardiidae).
Memoirs of the Queensland Museum 46:
161-174.

JEANNEL, R. 1944. Un carabique nouveau de la
Nouvelle-Caledonie (Coleoptera). Revue
francaise d'Entomologie 10: 84-85.

JOSEPH, L., MORITZ, C. & HUGALL, A. 1995.
Molecular support for vicariance as a source of
diversity in rainforest. Proceedings of the Royal
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LOUWERENS, C.J. 1949. Carabidae (Col.) from the
Sunda Islands. Wisscnschaftliche Ergebnissc dcr
Sunda-Expedition des Naturhistorischen
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MACLEAY, W.J. 1871. Notes on a collection of insects
from Gayndah. Transactions of the
Entomological Society of New South Wales 2:
79-205.

MANDL, K. 1969. Zwei neue Heptodonta- Arten und
eine neue Carabidae-Gattung (Col.) aus Nord-
Bomeo. Zeitschrift der Arbeitsgemeinschaft der
dsterreichischen Entomologen 2 1 : 51-54.

MONTEITH, GB. 1997. Revision of the Australian flat
bugs of the subfamily Mezirinae (Insecta:
Hcmiptera: Aradidae). Memoirs of the
Queensland Museum 41: 1-169.

MOORE, B.P. 1963. Studies in Australian Carabidae
(Coleoptera) 3. The Psydrinae. Transactions of
the Royal Entomological Society of London 115:
277-290.

1984. Taxonomic notes on some Australasian
Mecyclothorax (Coleoptera: Carabidae:
Psydrini) and descriptions of new species.
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23: 161-166.

1 985. The Carabidae ofNorfolk Island. In Ball, GE.
(ed.) Taxonomy, phylogeny and zoogeography
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memory' of Philip Jackson Darlington, Jr. Series
Entomologia 33: 237-256. (W. Junk).

1992. The Carabidae of Lord Howe Island
(Coleoptera: Carabidae). Pp. 159-173. In
Noonan, GR., Ball, GE. & Stork, N.E. (eds) The
biogeography of ground beetles of mountains
and islands. (Intercept: Andover).

In press. A new, cave-dwelling, species of
Laccocenus Sloane (Coleoptera: Carabidae:
Psydrini) from southern New South Wales. The
Australian Entomologist.

MOORE, B.P., WEIR, T.A. & PYKE, J.E. 1987.
Rhysodidae and Carabidae. Pp. 17-320. In
Zoological Catalogue of Australia, 4. (Australian
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MORITZ, C.. PATTON, J.L., SCHNEIDER, C.J. &
SMITH, T.B. 2000. Diversification of rainforest
faunas: an integrated molecular approach. Annals
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533-563.

PERRAULT, GG 1978. La faune des Carabidae de
Tahiti II genre Mecyclothorax { Sharp). Nouvelle
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1992. Endemism and biogeography among Tahitian
Mecyclothorax species (Coleoptera: Carabidae:
Psydrini). Pp. 201-215. In Noonan, GR., Ball,

PSYDRINE GROUND BEETLES

109

GE. & Stork, N.E. (eds) The biogeography of
ground beetles of mountains and islands.
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RUSSELL, B., MORITZ, C., YEATES, D.K. &
MONTEITH, GB. in press. Biogcography and
evolution of the genus Aellocoris Kormilev
(Hemiptera: Aradidae) in the Queensland Wet
Tropics.

SCHNEIDER, C.J., CUNNINGHAM, M. & MORITZ,
C. 1998. Comparative phylogeography and the
history of endemic vertebrates in the Wet Tropics
rainforest of Australia. Molecular Ecology 7:
487-98.

SHARP, D. 1903. Coleoptera Caraboidea. Coleoptera
pt. II. Pp. 175-292. In Sharp, D. (ed.) Fauna
Hawaiiensis. Vol. Ill, pt. III. (Cambridge,
University Press: Cambridge).

SLOANE, T.G. 1890. Studies in Australian
Entomology. No IV. New genera and species of
Carabidae. Proceedings of the Linnean Society of
New South Wales 5: 641-653.

1898. On Carabidae from West Australia, sent by
Mr. A.M. Lea (with descriptions of new genera
and species, synoptic tables &c.). Proceedings of
the Linnean Society of New South Wales 23:
444-520.

1900. Studies in Australian Entomology. No IX.
New species of Carabidae (with notes on some
previously described species, and synoptic lists
of species). Proceedings of the Linnean Society
of New South Wales 24: 553-584.

1911. Carabidae from Dorrigo, N.S.W. Proceedings
of the Linnean Society of New South Wales 35:
823-843.

1915. Studies in Australian entomology. No XVII.
New genera and species of Carabidae.
(Pamborini, Migadopini, Broscini, Cunepectini,
Nomiini, Pterostichini, Platynini, Oodini,
Harpalini, Lebiini). Proceedings of the Linnean
Society of New South Wales 40: 438-473.

1916. Carabidae from the Upper Williams River,
N.S.W. (Coleoptera). Proceedings of the Linnean
Society of New South Wales 41: 196-208.

1920. The Carabidae of Tasmania. Proceedings of
the Linnean Society of New South Wales 45:
113-178.

STRANEO, S.L. 1941. Sui tipi di Pterostichini
(Coleopt. Carabid.) Australiani della collezione
Castelnau nel Museo Civico di Genova Nota III.
Annali de Museo Civico di Storia Naturale de
Genova 61: 83-94.

FURTHER INFORMATION ON TWO PONTONIINE SHRIMPS FROM ASCIDEAN
HOSTS, DASELLA BRUCEI BERGGREN, 1990 AND PSEUDOPONTONIA MINUTA
(BAKER, 1907) (CRUSTACEA: DECAPODA: PALAEMONIDAE)

A.J. BRUCE

Bruce, A.J. 2003 06 30: Further information on two pontoniine shrimps from ascidian hosts,
Dasella brucei Berggren, 1990 and Pseudopontonia minuta (Baker, 1907) (Crustacea:
Decapoda: Palaemonidae). Memoirs of the Queensland Museum 49(1): 111-1 14. Brisbane.
ISSN 0079-8835.

Further records of two rare and little known pontoniine shrimps, Dasella brucei Berggren,
and Pseudopontonia minuta (Baker) are reported from Queensland and South Australia
respectively. Dasella brucei is previously known only from its lost holotype specimen and
Pseudopontonia minuta has been reported on only three previous occasions. A new host
record is reported for Dasella brucei and the host of Pseudopontonia minuta, the ascidian
Polycarpa flava Kott, is identified for the first time. □ Natantia, Pontoniinae, Dasella
brucei, Pseudopontonia minuta, new ascidian host records, Queensland, South Australia.

A.J. Bruce, Queensland Museum, PO Box 3300, South Brisbane, 4101 Australia (e-mail
abruce@broad.net.au); 20 June 2002.

Pontoniine shrimps of the genus Dasella
Lebour, 1945, are associates of some Indo-West
Pacific ascidian hosts (Bruce, 1993). Only three
species have been described, two from Australia
and one from the western Indian Ocean.

In 1981, Bruce reported a male specimen of
Dasella herdmaniae from Herdmania momus
(Savigny) from Heron Island, Queensland. At
that time D. herdmaniae was the only species of
the genus and otherwise previously known only
from the type material from Tuticorin, India.

Dasella ansoni was described by Bruce ( 1 983a)
from the Arafura Sea in association with Phallusia
depressiuscula Heller. Berggren (1990) reported
on further specimens of D. herdmaniae from
Mozambique and demonstrated the specimen
from Heron Island was not conspecific and
designated the single specimen as a new species,

D. brucei. This specimen, which would have
become the holotype of D. brucei was dispatched
from Heron Island to the Australian Museum on
10 August 1979 but was never received and
cannot now be located in the Australian Museum
collections (P. Berents, 23 May 2000, pers.
comm.) or elsewhere. It is therefore presumed
lost. There have been no subsequent reports of
this species in the literature. Recently further
specimens have become available for study,
considerably extending the range of this species
which is still known only from Queensland waters.

Pseudopontonia minuta (Baker), was one of
the earliest Australian pontoniine shrimps
described and until now was one of the rarest and
least known, with only the type specimens from

‘South Australia’ and two other specimens, both
from New South Wales (Baker, 1907; Bruce,
1972, 1983b). The host animal also remained
unidentified. Further material has been collected
in the MAFRI Benthic Laboratory Survey of Port
Adelaide harbour, which enabled the host to be
identified for the first time.

The specimens are deposited in the collections
of the Queensland Museum, (QMW). CL refers
to the post orbital carapace length. QMGH numbers
refer to the host catalogue numbers. Full details
of D. brucei and P. minuta and synonymies are to
be found in Li (2000) and Davie (2002)

SYSTEMATICS

Sub-phylum CRUSTACEA
Order DECAPODA Latreille, 1802
Family PALAEMONIDAE Rafinesque, 1815
Sub-family PONTONIINAE Kingsley, 1878

Dasella brucei Berggren, 1990
(Fig. 1)

Dasella herdmaniae Bruce, 1981: 50-54, figs 1-2.

Dasella brucei Berggren, 1990: 558. Chace & Bruce, 1993:
78 (key). Muller, 1993: 23, (full synonymy). Li, 2000: 42,
fig. 42, (full synonymy). Davie, 2002: 309.

MATERIAL . Queensland: 1 $ off Murdock Point, North
Queensland, 2 September 1977, 5m, QMW25502. 1 ov. $ ,
Mooloolaba, 4 September 1977, 2.9-22.9m, coll. A.
Rozenfelds. I ov. 9 . Curtin Artificial Reef, Moreton Bay,
stn. 22, 27.07°S 153.22°E,22m,4 April 1995, coll. J. Short,
QMW20466.

HOSTS. (QMW25502) in branchial sac of Herd-
mania grandis (Heller), QMGH768. (Rozenfelds

112

MEMOIRS OF THE QUEENSLAND MUSEUM

specimen and QMW20466) Herdmania momus
(Savigny), QMGH2200, (Ascidiacea).

TYPES. As the original specimen of D. brucei is
lost, the intact female specimen (#1, CL 2.6mm,
W25502) is now designated as a neotype. Type
locality: 14°36’S, 144°54’E.

REMARKS. The Murdock Point specimen is in
good condition, intact, with all pereiopods. The
Mooloolaba specimen is rather incomplete and in
an extremely fragile condition, but agrees well
with the previously published information. A
single pereiopod, probably the third, with the
diagnostic propod and dactyl is preserved.

Dasella brucei is distinguished from D. herd-
maniae by the pair of normal slender tapering
acute distoventral spines on the third ambulatory
propod (Fig. IB), with similar spines along the
ventral border, whereas D. herdmaniae has a pair
of small blunt distally swollen club-shaped
spines distoventrally, with similar spines
ventrally in these positions (Berggren, 1990).

Berggren (1990) commented on the marked
ridge below the level of the mobile hepatic spine
in D. herdmaniae. It may be noted that a similar
horizontal angulation is also present in the
Mooloolaba specimen of D. brucei (Fig. 1 A) but
inconspicuous in the Murdock Point specimen.
The carapace in the former is therefore rather
depressed, with a concavity above the lateral
ridge. The Moreton Bay specimen is the largest,
CL 3.25mm. In all specimens the ventral rostral
tooth is obsolescent.

The fifth thoracic stemite bears a transverse
carina, posterior to the second pereiopod coxae,
with a small median notch. Length of un-
developed ovum, 0.5mm.

The hosts of these shrimps both belong to
Herdmania. The Murdock Point specimen was
found in Herdmania grandis (Heller), an
endemic Australian species, and the Mooloolaba
and Moreton Bay specimens in H. momus
(Savigny), a species of widespread Indo-West
Pacific distribution. Dasella has been found in
association with three genera of ascidian host
Herdmania (D. brucei , D. herdmaniae ), Pyura
(D. herdmaniae) and Phallusia (D. ansoni).

Dasella brucei is known from three localities in
Queensland waters and has not been reported
from outside Queensland. Dasella herdmaniae
has been reported from Mozambique (Berggren,
1990), southern India (Lebour, 1939) and the
Philippines (Chace & Bruce, 1993). It is possible
that the original Heron Island specimen of D.

FIG 1. Dasella brucei Berggren, ovigerous 9, off
Mooloolaba. A, anterior carapace and rostrum,
dorsal; B, distal propod and dactyl of (?) third
pereiopod.

brucei , reported in association with H. momus
(Bruce, 1981) was associated with H. grandis ,
which was considered a synonym of H. momus at
that time, but both species of Herdmania occur on
Heron Island.

DISTRIBUTION. The type locality at Heron Is.,
Murdock Point, Mooloolaba and Moreton Bay.

Pseudopontonia minuta (Baker, 1907)

(Fig. 2)

Pontonia minuta Baker, 1907; 189-190, pi. 24, figs 9-12.
Hale, 1927: 57, fig. 51. Bruce, 1972: 65-74, figs 1-5.
Chace & Bruce, 1993: 62.

Pseudopontonia minuta Bruce, 1992: 1274. Muller, 1993:
128. Li, 2000: 280-281. Davie, 2002: 337-338.

MATERIAL. South Australia: 9, CL 4.0mm, Port
Adelaide, MAFRI Benthic Laboratory Survey, stn 7, 20
December 2002, from pile scrapings, in branchial sac; d,
CL ca. 4mm, idem, in branchial sac. QMW26553.

PONTONIINE SHRIMPS FROM QUEENSLAND ASCIDIANS

113

CEG

Fig. 2. Pseudopontonia minuta (Baker), Port Adelaide, W26553. A, antennule, ventrolateral aspect; B, second
pereiopod chela, fingers; C, third pereiopod, propod and dactyl; D, same, distal propod an ac y , ,
pleopod; F, same, endopod;G, second pleopod; H, same endopod. (A-B, V ; C-H, o ).

HOST. Polycarpaflava Kott, 1 985, ( Ascidiacea).

REMARKS. These specimens agree well with
earlier descriptions and little further description
is necessary. The male has the carapace badly
damaged and largely detached, together with the
eyes and antennae. The antennular peduncle (Fig.
2 A) is robust, the proximal segment stout with a
large acute ventromedial tooth. The lower
flagellum is short, with only four segments; the

upper flagellum is reflexed as in many Pontonia
species. The fingers of the second pereiopod
chelae (Fig. 2B) have strongly curved acute tips,
the cutting edges with two low teeth proximal ly,
the distal edge sharp. The third pereiopod propod
(Fig. 2C) and dactyl (Fig. 2D) are as previously
described. The male first pleopod (Fig. 2E) has
the protopodite slender, about 2.4 x longer than
wide. The exopod is of subequal length, with

114

MEMOIRS OF THE QUEENSLAND MUSEUM

numerous plumose marginal setae. The endopod
(Fig. 2F) is about 0.5 of exopod length, slender,
tapering, about 4.0 x longer than its basal width,
with five short similar simple spines on the
central two fourths of the medial margin, three
short feebly plumose setae laterally with a further
preterminal plumose seta. The second pleopod
protopodite (Fig. 2G) is broader, about 1.5 X
longer than wide, of similar length to the first.
The exopod is similar to the first pleopod. The
endopod (Fig, 2H) is about 0.8 of the exopod
length, with appendices at about 0.5 of the medial
margin length. The appendix interna is long and
slender, about 0.3 of the endopod length, with
only 4-5 terminal cincinnuli. The appendix
masculina is very short and stout, about 0. 12 of
the endopod length, 2.0 X longer than wide, with
a single long feebly plumose terminal spiniform
seta, 3.5 x longer than the corpus, reaching to
beyond the tip of the endopod.

Pseudopontonia minuta is remarkable amongst
pontoniine shrimps, as it is the only species known
with abbreviated larv al development (Bruce, 1 972).
All previous specimens have been females. The
isolated male and female within the ascidian
branchial sac is also surprising as pontoniine
associates of ascidians, and other hosts, are
usually found as heterosexual pairs. In the case of
Parana his tus armatus (H. Milne Edwards, 1 837)
it has been suggested that the high rate of host
infestation with heterosexual pairs of shrimps is
related to an ample source of larvae (Bruce, 2000).
The reverse might apply to the populations of
Pseudopontonia minuta.

The establishment of the identity of the host for
this species removes one obscurity but raises
another. A possible association with nereid
worms, possibly Eunice aphrodite , was
suggested by Bruce (1983b). This was based on a
colour slide which showed a pair of small red
pontoniine shrimps on this host, from Long Reef,
Sydney, New South Wales. The identity of these
shrimps has not yet been established and would
be of special interest as no pontoniine shrimps are
presently known to associate with any annelids.

DISTRIBUTION. The type locality ‘South
Australia’, Port Adelaide, South Australia, and
Meroo Point and Long Reef, New South Wales.

ACKNOWLEDGEMENTS

I am most grateful to Dr P. Mather for the
opportunity to report on these specimens and for
information on the hosts. The preparation of this

study was facilitated by the support of the

Australian Biological Resources Study.

LITERATURE CITED

BAKER, 1907. Notes on South Australian Decapoda
Crustacea. Part V. Transactions of the Royal
Society of South Australia 31: 173-191.

BERGGREN, M. 1990. Dasella herdmaniae (Lebour)
(Decapoda: Natantia: Pontoniinae) from
Mozambique and establishment of a new species,
Dasella brucei. Journal Crustacean Biology
10(3): 554-559.

BRUCE, 1972. Notes on some Indo-Pacific
Pontoniinae, XXIII. A re-description of Pontonia
minuta Baker, 1907, and the occurrence of
abbreviated development in the Pontoniinae
(Decapoda Natantia, Palaemonidae). Crustaceana
23(1): 65-75.

1981. Notes on some Indo-Pacific Pontoniinae,
XXXVII. Additional information on Dasella
herdmaniae (Lebour) (Decapoda Natantia).
Crustaceana 40(1): 50-56.

1983a. A second species of the genus Dasella
Lebour, D. ansoni sp. nov., from the Arafura Sea.
The Beagle, Occasional Papers of the Northern
territory Museum of Arts and Sciences 1(3):
21-29.

1983b. The pontoniine shrimp fauna of Australia.
Australian Museum Memoir 1 8: 1 95-2 18.(1 982).

1992. Designation of two new pontoniine shrimp
genera (Decapoda: Palaemonidae). Journal of
Natural History 26: 1273-1282.

1995. A Synopsis of the Indo-West Pacific Genera
of the Pontoniinae (Crustacea: Decapoda: Palae-
monidae). Theses Zoologicae 25(1994): 1-172.

2000. Biological observations on the commensal
shrimp, Paranchistus armatus (H. Milne
Edwards), (Crustacea: Decapoda: Pontoniinae).
The Beagle, Records of the Museums and Art
Galleries of the Northern Territory 16: 91-96.

CHACE, F.A., Jr, & BRUCE, A.J. 1993. The Caridean
Shrimps (Crustacea: Decapoda) of the Albatross
Philippine Expedition 1907-1910, Part 6:
Superfamily Palaemonoidea. Smithsonian
Contributions to Zoology 543: i-vii, 1-252.

DAVIE, P.J.F. 2002. Crustacea: Malacostraca:
Phyllocarida, Hoplocarida, Eucarida (Part 1). Pp.
xii, 551. In Wells, A. & Houston, W.W.K. (eds)
Zoological Catalogue of Australia, 19. 3 A.
(CSIRO Publishing: Melbourne).

HALE, H.M. 1 927. The Crustacea of South Australia 1 :
pp. 201. (Government Printer: Adelaide).

LEBOUR, M.V. 1939. Decapod Crustacea associated
with the ascidian Herdmania. Proceedings of the
Zoological Society of London 108B: 649-653.

LI, XINZHENG 2000. Catalog of the genera and
species of Pontoniinae Kingsley, 1878.Pp. i-iv,
1-319. (Xueyuan Press: Beijing).

MULLER, H.-G 1993. Catalogue of the Indo-Pacific
Pontoniine Shrimps. P. 148. (Wetzlar).

PERICLIMENES SPECIES (CRUSTACEA: DECAPODA: PONTONIINAE) FROM FAR

NORTH QUEENSLAND

A.J. BRUCE

Brucc, A.J. 2003 06 30: Periclimenes species (Crustacea: Decapoda: Pontoniinae) from Far
North Queensland. Memoirs of the Queensland Museum 49(1): 115-122. Brisbane. ISSN
0079-8835.

Recent collections from inshore waters of Cape York have provided specimens of four
shrimp species of the genus Periclimenes Costa, 1 844, from a single locality, Cape Flattery.

One is a well known species that has been rarely reported from Australia, two are described
as new species and the fourth, damaged, cannot be referred to any described species. □
Decapoda, Caridea, Pontoniinae, Periclimenes, P. adularans, P. paulsoni, new species,

Cape York, Queensland.

A.J. Bruce, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia (e-mail
abruce@broad.net.au); 2 July 2002.

Recent surveys of inshore waters of northern
Queensland for the Introduced Marine Pests
Baseline Survey Projects, by the Marine Biology
and Aquaculture Department of James Cook
University, suggest that much still remains to be
learned about the caridean fauna of this biotope,
which has attracted much less scientific attention
than the nearby Great Barrier Reef. Of the four
species of the pontoniine Periclimenes Costa,
1844, only one can be placed in a described
taxon, which suggests that more extensive or
more detailed sampling might well produce a
further substantial increase in the caridean
diversity known from this region.

Two of the species are described as new species
and a third, represented by a single damaged
specimen, probably also represents a further as
yet undescribed species. The fourth species
collected, Periclimenes grand is, is a well known
and widely distributed Indo-West Pacific species,
but one that has been only infrequently recorded
from Australian waters.

In the descriptions, CL refers to the postorbital
carapace length; QM W to the Queensland Museum
specimen catalogue numbers. Restricted
synonymies only are provided. Full synonymies
are to be found in Li (2000).

SYSTEMATICS

Subphylum CRUSTACEA
Order DECAPODA Latreille, 1803
Family PALAEMONIDAE Rafinesque, 1815
Sub- family PONTONIINAE Kingsley, 1878

Periclimenes grandis (Stimpson, 1860)
(Fig. 1)

Anchistia grandis Stimpson, 1860: 39.

Periclimenes grandis Borradaile, 1898: 382.

Periclimenes (Ancylocaris) grandis Kemp, 1922: 210-214,
figs 58-59, pi. 7, fig 10. „ ,

Periclimenes grandis Li, 2000: 186-187, fig. 235 (full
synonymy).

MATERIAL. 1 ovig. 9, # Sh 369, Cape Flattery; inner
wharf pile, P3, scrapings, 7m, August 200 1 ; 1 9 , # Sh 370,
Cape Flattery, inner wharf pile, P6, 3m, 27 October 2001.

DISTRIBUTION. Australia. Queensland: Magnetic
Island (Bruce, 1977); John Brewer Reef (Bruce,
1987a); Abbot Point (Hoedt, et al., 2000)
Northern Territory; Darwin (Bruce, 1987c); East
Point, Darwin (Bruce, 1988a); Cobourg
Peninsula (Bruce & Coontbes, 1995); Darwin
Harbour: Bullocky Point, Cameron Beach,
Channel Island, Nightcliffe, Dudley Point, Lee
Point, Shell Island, Weed Reef (Bruce &
Coombes, 1997). Western Australia: Hibernia
Reef (Bruce, 1992).

General. Type locality: Oshima, Japan. Also
known from Egypt, Israel, Jibuti, Yemen, Kenya,
Zanzibar, Tanganyika, Mozambique, Comoro
Islands, Madagascar, Seychelle Islands, Sri
Lanka, Burma, Malaya, Singapore, Indonesia,
Vietnam, China, Japan, Papua New Guinea,
Western Australia, Northern Territory,
Queensland, Japan, Caroline Islands, Marshall
Islands, Fijian Islands, Tuvalu and Tuamotu
Islands.

REMARKS. The present specimens (CLs 4.7,
4.0mm) present no special features. The rostral
dentitions are:! + 6/4. 1 + 5/4. The third
ambulatory propod is 1 1 X longer than wide,
with a pair of strong distoventral and 4 ventral
spines. The dactyl is 0.28 of the propod length,
about 4 X longer than its basal width, rather
stouter than as reported by Kemp (1922), who

116

MEMOIRS OF THE QUEENSLAND MUSEUM

notes 6-6.5 X longer. This free-living micro-
predator is probably much commoner in warmer
waters than the relatively sparse Australian
records suggest. It is surprising that no specimens
were found on Heron Island (Bruce, 1981) during
collections from 1975 to 1981. Specimens
however have been collected from Heron Island
(coll. A.H. Banner, 1968). Other specimens have
been seen from Bundegi Reef, Exmouth Gulf and
North West Cape, Western Australia (coll. N.L.
Bruce, 1980).

Periclimenes adularans sp. nov.

(Fig. 2)

MATERIAL. Holotype: 1 ovig. 9 , # Sh 3 1 1 , Cape Flattery
Service Jetty, 27 April 2001, beam trawl, 2m, QM
W26554. Paratypcs: 1 d, # Sh 317, Cape Flattery Service
Jetty, 27 April 2001 , beam trawl, 2m, coll. QM W26555; 1
ovig. 9 , Sh 371, Cape Flattery, Service Jetty, beam trawl,
15m, 27 October 2001, QM 26556.

ETYMOLOGY. Latin adulari , to flatter, after the locality
of capture.

DIAGNOSIS. Small slender shrimp of the holthuisi
species group. Rostrum strongly arched (Fig. 2A),
ventrally sinuous, distally concave, reaching to
near distal margin of intermediate segment of
antennular peduncle, with 13 small acute teeth
dorsally in female, 1 1 in male (Fig. 2B), with
single small subterminal tooth ventrally.
Carapace with epigastric tooth in female, absent
in male; inferior orbital angle (Fig. 2C) strongly
produced, acute, with reflected inner flange;
hepatic spine slender, slightly anterior to the level
of the epigastric tooth in the female; third
abdominal tergite slightly posteriorly produced,
not carinate; first and second pereiopods normal,
similar, slightly unequal, second pereiopod (Fig.
2D) chela (female) (Fig. 2E) 0.9 of CL, with
fingers subequal to palm length, dactylus
unarmed, fixed finger with 2 minute acute teeth
proximally, carpus 0.6 of chela length, subequal
to merus: ischium 1.2 x merus length; third
ambulatory pereiopod (Fig. 2F) with dactyl (Fig.
2G) slender, 7.7 X longer than basal width, 0.28
of propod length, with small slender accessory
tooth at 0.8 of length, closely adpressed, unguis
not clearly demarcated: propod about 19 X
longer than width, uniform. Compared with the
other species of this group, P adularans has a
smaller more slender accessory tooth that is
closely adpressed to the unguis. The dactylus is
distinctly longer and more slender than in the
other species of this group, in which the accessory
tooth is also stouter and more projecting, about

4 X longer than the basal width, and 0.2 of the
propod length (Bruce, 1990).

MEASUREMENTS (mm). Holotype, CL 3.7;
paratypes: (Sh 317), CL 2.7, (Sh 371), CL 3.4
mm; length of ova 0.5.

SYSTEMATIC POSITION. Closely related to
Periclimenes tosaensis Kubo, 1951. Peri-
climenes adularans may be easily distinguished
from P. tosaensis by the accessory tooth on the
ambulatory dactylus as this is absent in P
tosaensis, the onlv species of the holthuisi-g roup
in which it is lacking. This dactylus is otherwise
very similar in these two species, about 7 X
longer than the basal width, about 0.28 of the
propod length. Compared with the other species
of this group, P. adularans has a smaller more
slender accessory tooth that is closely adpressed
to the unguis. The dactylus is distinctly longer
and more slender than in the other species of this
group, in which the accessory tooth is also stouter
and more projecting, about 4 x longer than the
basal width, and 0.2 of the propod length (Bruce,
1990).

COLOURATION AND HOST. Not recorded.
Trawl catches did not indicate any potential hosts.

HABITAT. Silty mud substrate.

REMARKS. The holotype female has a single
second pereiopod. The male lacks its right eye,
one first pereiopod, both second pereiopods and
fourth and fifth pereiopods. The epigastric tooth
also may have been lost artificially. The
ovigerous female paratype lacks the distal half of
the rostrum, and pereiopods 3-5 on the right side.

PONTONIINE SHRIMPS FROM CAPE YORK

117

FIG 2. Periclimenes adularans sp. nov. A, carapace and rostrum, 9 ; B, same, 6 ; C, inferior orbital angle, D,
second pereiopod; E, same, chela; F, third pereiopod, propod and dactyl; G, same, distal propod an ac y . ( ,
& E, holotype; B,C,F & G, paratype).

A key to the then 6 species of the Periclimenes
holthuisi group was provided by Bruce (1990).
Since then two further species have been
described: P. tenuirostris Bruce, 1991, and P
kobayashii Okuno & Nomura, 2002. An updated
key follows:

1 . Ambulatory dactyls simple; R 1 + 7- 1 0/2

P. tosaensis Kubo

Ambulatory dactyls biunguiculate 2

2. Carapace with 2-3 postorbital teeth; R 2-3 + 6-7/2-3. . .

P. aesopius (Bate)

Carapace with 0- 1 postorbital teeth 3

3. Carpus of second pereiopods distinctly longer than chela;

without epigastric tooth; R 5-6/0-2 • • • •

P. longicarpus Bruce & Svoboda

Carpus of second pereiopods shorter than chela .... 4

4. Fingers of second pereiopod chela with series (5-7) of

small acute recurved teeth; propods of ambulatory
pereiopods with short distoventral spines; R 1 4- 5-7/0-2
P. venustus Bruce

Fingers of second pereiopod chelae without series of
small acute recurved teeth; propods of ambulatory
pereiopods with long distoventral spines 5

118

MEMOIRS OF THE QUEENSLAND MUSEUM

5. Second pereiopods with chelae bowed, carpus about 0.6
of palm length; R 1+7-8/ 1-2 . . . P. magnificus Bruce

Second pereiopods with chelae not bowed, carpus
subequal to palm length or longer 6

6. Third abdominal tergite without posterior median carina . 7
Third abdominal tergite with posterior median carina . 8

7. Third ambulatory dactyl 7 or more x longer than basal

width, about 0.28 of propod length, with very slender

adpressed accessory tooth: R 0-1 + 11-13/1

* . . . . P.adularans sp.nov.

Third ambulatory- dactyl about 4 x longer than basal
width, about 0.2 ol propod length, with well developed

projecting accessory tooth; R 1 (-2) + 8-ll/2-4

P. holthuisi Bruce

8. Rostrum feebly arched, almost straight, directed

upwards, exceeding antennular peduncle; first pereiopod
with dactylus longer than palm; R 1 +6-7Z2-3 . ... P.
tenuirostris Bruce ,

Rostrum distinctly arched not exceeding antennular
peduncle; first pereiopod dactylus shorter than palm; R 1
+ 6-8/ 1-3 P kobayashii Okuno & Nomura

Periclimenes paulsoni sp. nov.

(Fig. 3A-P)

MATERIAL. 1 6 holotype, Sh 383, Cape Flattery, inner
wharf pile, scrapings, 7m, August 2001, coll.# FAK, QM
W26557.

ETYMOLOGY. Named in honour of Otto Mikhailovich
PauPson, ( 1 837-86), author of Studies on the Crustacea of
the Red Sea (1875).

DIAGNOSIS. Small sized slender shrimp of the
grandis species group. Rostrum (Fig. 3B) slender,
about 0.8 of CL, well exceeding antennular
peduncle, horizontal, slightly up-curved, with 5
small acute teeth dorsally, 3 ventral teeth, tip
acute, simple. Carapace (Fig. 3 A) with epigastric
tooth, robust marginal antennal spine, inferior
orbital angle (Fig. 3C) obsolete; hepatic spine
slender, slightly anterior in level to the epigastric
tooth; scaphocerite (Fig. 3E) slender, with distal
tooth far exceeding lamella; cornea hemi-
spherical (Fig. 3F), diameter about 0.33 of CL;
fourth thoracic stemite with slender median
process; third abdominal tergite not posteriorly
produced; First pereiopods (Fig. 3G,H) normal,
exceeding scaphocerite by length of chela, carpus
subequal to chela; second pereiopod (Fig. 31)
chela 1 .2 of CL, with fingers (Fig. 3J) about 0.4 of
palm length, unarmed, acute hooked tips, cutting
edges entire, carpus 1.25 of chela length,
subequal to merus and ischium length; third
ambulatory pereiopod (Fig. 3K) with dactyl (Fig.
3L) robust, simple, curved, 5.7 x longer than
basal width, 0.3 of propod length, unguis not
clearly demarcated; propod about 10.5 x longer
than width, subuniform, with 2 slender

distoventral spines, 4 smaller solitary ventral
spines; telson and uropods normal.

MEASUREMENTS (mm). Holotype, CL 1.35,
carapace and rostrum 3.2; second pereiopod
chela 1.9; third pereiopod propod 1 .6.

SYSTEMATIC POSITION. Periclimenes
paulsoni is closely related to P. anacanthus Bruce,
1988b and P. nilandensis Borradaile, 1915. It
may also be closely related to P edwardsii
(Paulson, 1 875) with which species the specimen
was initially identified. Periclimenes edwardsii
and P nilandensis were not included in the
grandis species group by Kemp (1922) as they
lacked distoventral meral spines on the second
pereiopods. Bruce (1987a) revised the species of
this group and included all species with a
conspicuous finger-like median process on the
fourth thoracic stemite, a feature present in P.
paulsoni , P. anacanthus and P. nilandensis. It is
not known if it is present in P edwardsii , but this
seems likely from the close resemblances between
the three species.

Periclimenes paulsoni may be distinguished
from P. anacanthus by the shorter more slender
rostrum, about 1 .2 of CL, as opposed to 1 .6 in
male of P anacanthus , lesser rostral dentition, 1
+ 5/3, as opposed to 1 + 6-9/2-3; the obsolete
inferior orbital angle, well developed, acute in P.
anacanthus ; shorter stouter, 5 segmented fused
ramus of upper antennular flagellum, as opposed
to long slender II segmented ramus; comeal
diameter about 0.3 of CL, as opposed to 0.6; First
pereiopod carpus subequal to chela; second
pereiopod with carpus slightly longer than palm,
subequal to meral length, comparatively short
and stout, as opposed to longer and more slender
in P. anacanthus ; third pereiopod dactyl about 5.7
X longer than basal width, as opposed to 6.5 x ,
propod about 10.5 x longer than width, as
opposed to 1 4.5 x in P. anacanthus.

The pleopods (Fig. 3M,0) of P. paulsoni are
unusual and differ markedly from P. anacanthus.
They are similar on left and right sides. The First
pleopod endopod (Fig. 3N) is simple, tapering
distally, about 4 x longer than the basal width,
with 3 feeble setae medially. In P anacanthus it is
about 7 x longer than the basal width, much
expanded centrally, with numerous spines on the
concave proximo-medial margin, with numerous
Fine marginal setae over the rest of the expanded
portion. The endopod of the male second pleopod
(Fig. 3P) is much reduced in P paulsoni, about
0.6 of the exopod length, with a small appendix
interna at 0.75 of the length. The appendix

PONTONIINE SHRIMPS FROM CAPE YORK

119

FIG. 3. A-P, Periclimenes paulsoni sp. nov. holotype 6 . A, carapace and appendages; B, rostrum; C, interior
orbital angle, dorsal aspect; D, antennule; E, antenna; F, eye, dorsal; G, first pereiopod; H, same, chela; I, second
pereiopod; J, same, chela; K, third pereiopod, propod and dactyl; L, same, distal propod and dactyl; M, first
pleopod; N, same, endopod; O, second pleopod; P, same, endopod. Q, Periclimenes edwardsii (Paulson),
redrawn from Paulson (1875).

120

MEMOIRS OF THE QUEENSLAND MUSEUM

masculina is very well-developed, about 0.6 of
the endopod length, far exceeding the end of the
endopod, with 5 long simple terminal spines and
1 slightly preterminal spine. In P anacanthus the
appendix masculina is much shorter than the
endopod, which is well-developed, about 0.6 of
its length, and has 6 terminal spines, with 4 spines
along the ventral surface.

Periclimenes paulsoni also resembles P.
nilandensis Borradaile, 1915. This species has a
distally broad scaphocerite, not greatly over
reached by the terminal tooth, a well developed
inferior orbital angle not found in P. paulsoni , the
first pereiopod carpus is markedly longer than the
chela, the second pereiopod carpus is markedly
shorter than the merus and the rostral dentition of
1 + 7-8Z3-4, with the first tooth situated on the
carapace.

The present specimen of P. paulsoni was
initially thought to be referrable to P. edwardsii
(Paulson, 1875), a little known species that has
rarely been reported since its original description.
The loss of all Paulson’s material has handi-
capped further description of this species. P.
edwardsii (Fig. 3Q) has a deeper rostrum than P.
paulsoni , with a rostral dentition of 1 + 7/3, with a
bifid tip, an obsolete inferior orbital angle very
similar to P. paulsoni , and a relatively broad
scaphocerite with the lamella exceeding the tip of
the distolateral tooth. Ledoyer ( 1 968) reported on
a number of specimens as P. cf. edwardsi from
Tulear, Madagascar, on the basis of the
assessment by Kemp (1922), and illustrated the

major features. The figure shows a rostral
dentition of 1 + 6/2 and a feebly produced inferior
orbital angle. His material differs from Paulson’s
particularly in the second pereiopod where the
slender carpus, 7 X longer than the distal width,
is subequal to the palm length, about 4 x longer
than wide and 0.6 of the palm length in P
edwardsii s.str.

The key to the species of the expanded
1 Periclimenes grandis species group’ given in
Bruce ( 1 987b) was augmented in Bruce ( 1 988b).
A further augmentation is provided below, to
include P. paulsoni and P. edwardsii , which was
not included in the 1 987a key.

16. Supraorbital spines present 16a

Supraorbital spines absent 17

16a Second pereiopod carpus much longer than palm; R. 1 +

6-9/2-3 P. anacanthus Bruce

Second pereiopod carpus not longer than palm ... 16b

1 6b Inferior orbital angle obsolete; R. 1 + 5/3

P. paulsoni sp.nov.

Inferior orbital angle distinct 16c

16c Rostral lamina slender, second pereiopod carpus

distinctly shorter than merus; R. 1 + 7-8/3-4

P. nilandensis Borradaile

Rostral lamina deep, second pereiopod carpus subequal
to merus; R. 1 + 7/3 P. edwardsii (Paulson)

Couplet 13 of the original key also contained
some unfortunate errors and should read:

13. Epigastric and first three dorsal teeth grouped and
enlarged; ambulatory propods with distal ventral spine

only; R. 1 -6-7/3-4 P. kororensis Bruce

Dorsal teeth similar and evenly distributed; ambulatory

propods ventrally spinulate; R. 1 + 6/5-6

P.platycheles Holthuis

PONTONIINE SHRIMPS FROM CAPE YORK

121

Periclimenes sp. aff. anacanthus Bruce, 1988
(Fig. 4)

MATERIAL. 1 9, Sh 305, Cape Flattery, Service Jetty,
beam trawl, 5m, August 2001, QM W26558.

REMARKS. The single specimen, CL 2.4mm,
with a well-developed median process on the
fourth thoracic stemite, unfortunately lacks both
second pereiopods. The rostrum (Fig. 4A), about
1.25 x the CL, far exceeds the antennular
peduncle and has a dentition of 1 + 9/4, the distal
ventral tooth being minute. The inferior orbital
angle is not acutely produced, almost obsolete.
The first pereiopod has the carpus about 1.35 x
the chela length. The propod of the third
pereiopod (Fig. 4B) is about 0.78 of the CL, 1 8.6
x longer than wide, with a pair of small
distoventral spines and two minute ventral spines
only. The dactyl (Fig. 4C) is about 0.57 of the
propod length, 8.5 x longer than the proximal
depth.

None of the other species of the ‘ grandis group’
that have supraorbital spines have such slender
ambulatory dactyls, except P. anacanthus. This
species has an acutely produced inferior orbital
angle, the first pereiopod carpus about 1.8 x the
chela length, the third pereiopod propod about
14.5 of the CL, with long distoventral spines and
numerous well developed ventral spines, and the
dactylus is about 0.4 of the propod length, 6.5 x
longer than its proximal depth. P. anacanthus is
known from Moreton Bay, Queensland and the
Cobourg Peninsula, Northern Territory.

The Cape Flattery specimen also shows some
similarity to P. digitalis Kemp (1922), which has
much more slender ambulatory dactyls, about 1 4
x longer than the proximal depth and 0.45 of the
length of the propod, which also lacks ventral
spinules. Periclimenes digitalis lacks a supra-
orbital spine, having only a small tubercle in this
position. Periclimenes digitalis has been
reported from the Andaman Islands, Singapore,
Hong Kong, China and Indonesia.

This specimen cannot be referred to any of the
described species of the ‘ grandis species group’
and appears to represent a distinct taxon. Without
second pereiopods it is not suitable for desig-
nation as a holotype specimen and the collection
of further complete specimens must be awaited
for a full description.

ACKNOWLEDGEMENTS

I am most grateful to Dr Kerry Neil for the
opportunity to report on these shrimps. The

specimens were collected by Juan Cruz, John
Ackerman, Phil Osmond and Damian Thomson.
This study was facilitated by support from the
Australian Biological Resources Study.

LITERATURE CITED

BORRADAILE, L.A. 1898. A revision of the
Pontoniidae. Annals and Magazine of Natural
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BRUCE, A.J. 1977. Pontoniine shrimps in the
collections of the Australian Museum. Records of
the Australian Museum 31(2): 39-81.

1981. Pontoniine shrimps of Heron Island. Atoll
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1987a. Metaphrvxus intutus Bruce (Crustacea:
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Northern Territory Museum of Arts & Sciences
3(1): 213.

1987b. Re-descriptions of two little-known Indo-
West Pacific palaemonid shrimps, Periclimenes
cal man i Tattersall and P. delagoae Barnard.
Journal of Natural History 21(6): 1415-1432.

1987c. Notes on some Indo-Pacific Pontoniinae,
XLIV. Periclimenes darwiniensis sp. nov., from
the Northern Territory, Australia (Decapoda,
Caridea). Crustaceana 52(1): 29-39.

1 988a. The shrimp fauna of a small tropical reef, the
East Point Fish Reserve, Darwin. In, Larson,
H.K., Michie, M.G & Hanley, J.R. (eds) Darwin
Harbour, Proceedings of the Workshop on
Research and Management held in Darwin, 2-3
September, 1987. A.N.U. North Australia Research
Unit, Mangrove Monograph 4:226-245.

1988b. A new palaemonid shrimp from the
Zostera- beds of Moreton Bay, Queensland. The
Beagle, Records of the Northern Territory
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1990. A new cnidarian - associated palaemonid
shrimp from Port Essington, Cobourg Peninsula,
Australia. Indo-Malayan Zoology 6(1989):
229-243.

1991. Shallow water Palaemonoid shrimps from
New- Caledonia (Crustacea: Decapoda). Pp.
2 1 -279. In Richer de Forges, B., (ed.) Le Benthos
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Caledonie, 1 . fetudes de Theses. (ORSTOM: Paris).

1 992 Crustacea: Decapoda Caridea. Pp. 1 28- 1 3 1 . In
Russell, B.C. & Hanley, J.R. (eds) The biological
resources and heritage values of the Cartier and
Hibernia Reef systems, Timor Sea. (Northern
Territory Museum: Darwin).

BRUCE, A.J. & COOMBES. K.E. 1995. The
palaemonoid shrimp fauna (Crustacea: Decapoda:
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Territory. The Beagle, Records of the Museums
and Art Galleries of the Northern Territory 12:
101-144.

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1997. An annotated check-list of the caridean shrimps
(Crustacea: Decapoda) of Darwin Harbour, with
descriptions of three new species o fPericlimenes
(Palaemonidae: Pontoniinae). Pp. 301-337. In
Hanley, J.R., Caswell, G, Megirian, D. & Larson,
H.K. (eds) Proceedings of the Sixth International
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HOEDT, F.E., CHOAT, J.H, COLLINS, J. & CRUZ, J.J.
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KEMP, S. 1922. Notes on Crustacea Decapoda in the
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Indian Museum 24: 113-288.

KUBO, I. 1951. Some macrurous decapod Crustacea
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four new species. Journal of the Tokyo University
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LEDOYER, M. 1968. Les Caridea de la frondaison des
herbiers de phanerogames de la region de Tulear
(Republique Malgache). Annales de l’Universite
de Madagascar 6: 65-1 1 5.

LI, XINZHENG 2000. Catalog of the Genera and
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Palaemonidae: Pontoniinae) from the Ryukyu
Islands, Southern Japan. Bulletin of the National
Science Museum, Tokyo, ser. A 28(4): 211-222.

OKUNO. J. & NOMURA, K. 2002. A New Species of
the * Periclimenes aesopius Species Group’
(Decapoda: Palaemonidae: Pontoniinae)
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83-94.

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ADDENDUM

Since the preparation of this article a further species of the Periclimenes holthuisi species
group has been described from the Ryukyu Islands, Japan by Okuno (2002). This species,
Periclimenes sarasvati Okuno, is most closely related to * Periclimenes venustus Bruce
(1990) and may be distinguished from that species by the absence of a bee ocellaire , the
presence of a bilobed distal maxillary endite, the second pereiopods exceeding the
scaphocerite by the proximal part of the palm, with the fingers bearing 2-4 acute recurved
teeth and a rostral dentition of 7-9/1 -2.

SYSTEMATICS AND NEW TAXA OF THE VANNIUS COMPLEX (HEMIPTERA:
MIRIDAE: CYLAPINAE) FROM THE AUSTRALIAN REGION

GERASIMOS CASSIS, MICHAEL D. SCHWARTZ AND TIMOTHY MOULDS

Cassis, G, Schwartz, M.D. & Moulds, T. 2003 06 30: Systematics and new taxa of the
Vannius complex (Hemiptera: Miridae: Cylapinae) from the Australian Region. Memoirs of
the Queensland Museum 49(1): 123-151 . Brisbane. ISSN 0079-8835.

Erection of the tribe Vanniini and its placement in the subfamily Palaucorinae, as proposed
by Gorczyca (1997), is rejected. An informal group, the Vannius complex (Cylapinae:
Cylapini) is recognised, comprised of Afrovannius Gorczyca, Austrovannius gen. nov.,
Paracylapus Carvalho, Vanniopsis Poppius, Vannius Distant, and Vanniusoides Carvalho &
Lorenzato. A ustrovannius nov. gen. is described, including three new species ,A.platnicki sp.
nov., A. scutica sp. nov., and A. xepenehense sp. nov. New species belonging to existing
genera within the Vannius complex arc described, including: Vanniopsis howense sp. nov.,
Vanniusoides asprokara sp. nov. and Vanniusoides melafmns sp. nov. Vanniopsis nifescens
Poppius, 1909 is redescribed and the male genitalia are illustrated for the first time.
Illustrations of male and female genitalia are given for the new species. A cladistic analysis
of the genera of the Vannius complex is presented and the systematic position of the complex
is discussed. □ Hemiptera, Miridae, Cylapinae , Vannius complex, Australian region.

Geras imos Cassis, (e-mail: genyc@austmus.gov.au), Centre for Biodiversity and
Conservation Research, 6 College Street, Sydney 2010, Australia; Michael D. Schwartz,
Research Associate, American Museum of Natural History, New York New York, USA
10024; Timothy Moulds, Centre for Evolutionary Biology and Biodiversity, University of
Adelaide, Adelaide 5005, Australia; 14 March 2003.

The Cylapinae are a basal group of Miridae
(Schuh, 1975) whose systematics and biology are
probably the least known in the family. Schuh
(1995) and Schuh & Slater (1995) rejected the
notion of meaningful monophyletic infrafamilial
groups within the Cylapinae, as suggested by
previous authors such as Carvalho (1952, 1957:
Bothriomirini, Cylapini & Fulviini). Gorczyca
(1997, 1998) has attempted to clarify the
suprageneric classification, although this author
has recently rejected his own inital findings (i.e.
Vannius complex as a tribe within the
Palaucorinae).

Gorczyca (1997) recognised the Vannius com-
plex as a monophyletic tribe of mirids because of
the spatulate parempodia, a condition unique in
basal Miridae. He recognised this complex as
containing Afrovannius Gorczyca, Austrovannius
gen. nov., Paracylapus Carvalho, Vanniopsis
Poppius, Vannius Distant and Vanniusoides
Carvalho & Lorenzato.

This work reports on an examination of new
material from the Australian region, with new
species described from the Australian mainland.
Lord Howe Island, Fiji, New Caledonia and
Vanuatu. This study also includes a broader
overview of the systematics of the Vannius
complex, with a description of three genera
(Austrovannius, Vanniopsis and Vanniusoides)

from the Australian region. In addition, the
ingroup relationships of the Vannius complex are
analysed, and their position within the Cylapinae
is discussed.

The Cylapinae, as with most mirid subfamilies,
are poorly documented for Australia. Cassis &
Gross (1995) listed only nine genera and 11
species, yet existing collections indicate that the
cylapine fauna of Australia, may be an order of
magnitude more species rich. This work is part of
a series on the systematics of the Miridae of the
Australian region, and the results herein are the
first record of the Vannius complex from
Australia.

This paper is based on a study of existing
collections from the tropical parts of the
Australian region, including the southwest
Pacific. A smaller collection of other specimens
from New Caledonia has been examined but not
included in this paper. This material represents at
least another ten new species of Austrovannius,
and it is apparent that material examined from
each mountain top is an endemic species. This
material will be the subject of a second paper, as
we await the collection of additional specimens.

MATERIALS AND METHODS

This study was based on about 100 specimens,
borrowed from the Bernice P. Bishop Museum

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MEMOIRS OF THE QUEENSLAND MUSEUM

(BPBM), the Queensland Museum (QM), the
South Australian Museum (SAMA), and the
University of Queensland Insect Collection
(UQIC). Dr Roger Kitching (Griffith University)
provided one specimen. The holotypes of Austro-
vannius platnicki sp. nov. and A. xepenehense sp.
nov. are lodged in the Museum National
d’Histoire Naturelle, Paris (MNHP). Material
was also collected by us during fieldwork on
Lord Howe Island and New Caledonia, and is
housed at the Australian Museum (AM). Geographic
coordinates for specimens without such inform-
ation were calculated using GIS techniques, and
are given in brackets in the Materials section of
the species descriptions.

All measurements are maximum lengths and
the range for each species is given in millimeters.
Abbreviations for morphometric characters are:
BL = body length; HW = head width across eyes;
IOD = interocular distance; HL = head length; PL
= pronotal length; PW pronotal width at humeral
angles; All = length of second antennal segment;
and, LL = labial length.

Scanning electron micrographs were prepared
using techniques outlined in Bolte (1996) and
Swearingen et al. (1997).

Colour images were captured using a Nikon
DX1 digital camera fitted with an Infinity K2
long distance microscope and a CF-3 objective.
Illumination was supplied by the Microptic
ML- 1000 fiber optics system with the images
relayed directly to Nikon Capture 2 software.

The data were constructed and analysed using
Winclada (Nixon, 1999) and NONA (GolobofF,
1999). The characters are unweighted and all
multistate characters are unordered. The
following three taxa were used as outgroups:
Bothriomiris lugubris Poppius, Peritropisca
bituberculata Carvalho and Lorenzato and an
undetermined species of Cylapocerus Carvalho.

BIOLOGY. The biology of the basal suprageneric
groups (Isometopinae, Cylapinae, Psallopinae)
of the Miridae are subject to debate because of a
lack of empirical evidence (Schuh, 1975, Cassis
& Gross, 1 995; Wheeler, 200 1 ). Many researchers
(e.g. Schuh, 1976; Carvalho & Lorenzato, 1978)
have argued that Cylapinae are stenotopic in their
habits, being associated with the mycelia and
fruiting bodies of higher fungi, in habitats such as
fallen logs and under bark. There are some
observations that support the notion that
cylapines are mycetophagous (Schuh, 1975).
Most cylapines (aside from Bothriomirini and
most members of the Vannius complex) are

characterised by an extremely elongate labium,
which is suggestive of fungal feeding. Unlike in
the Aradidae, which are well documented fungal
feeders, the labium of cylapines does not possess
coiled stylets, and there have been no reports on
the mechanism of cylapine feeding. Wheeler
(2001 ) suggested that some cyalpines, particularly
Fulviini, are predaceous.

There is very limited knowledge of the biology
of species of the Vannius complex. We found a
new species of Vanniopsis from Lord Howe
Island, V howense sp. nov., in lowland rainforest,
associated with leaf mould of fallen palm leaves
(Howea fosteriana Becc.). This microhabitat was
characterised by dense matts of fungal mycelia
and high moisture content. V howense was found
in association with other heteropteran taxa;
Acaraptera Usinger & Matsuda (Aradidae), and
two species of Reduviidae: Emesinae ( Emesopsis
sp. and Atisne derelictus Wygodzinsky).
Similarly, Vanniopsis rufescens Poppius and the
new species of Austrovannius have been
collected amongst leaf litter in rainforests. These
data provide indirect support that the Vannius
complex, like other cylapines are associated with
fungi. Other species of Vanniopsis are known
from vegetation, including sweeping ferns. Chris
Reid (pers. comm.) observed that V. howense was
occasionally collected on ferns. Vanniopsis
crobylos has also been collected from a low plant
(Elatostema sp. [Urticaceae]). Together, the
above data suggest that Vanniopsis species are
epigaeic but frequent low vegetation during the
day.

Austrovannius species are also primarily
ground-dwelling, collected from either sieved
litter or by pyrethrum fogging of fallen logs.
Nothing is known of the biology of the species of
Vanniusoides that are described in this work.
Gorczyca & Konstantinov (2001) reported that
Vanniusoides elypeatus Gorczyca & Konstantinov
is found in a riparian habitat amongst stones.

There is also limited biological information for
species of the Vannius complex from the
Afrotropical and Neotropical regions. Gorczyca
(1996, 1997) and Gorczyca & Konstantinov
(200 1 ) reported that Paracylapus lestoni is found
on fungi, and species of Vannius are known from
fruit and flowers.

HOMOLOGY AND TERMINOLOGY. A range
of external morphological and genitalic
characters are used to differentiate genera and
species within the Vannius complex. Homology
and terminology for non-genitalic characters are

SYSTEMATICS OF THE VANNIUS COMPLEX

125

consistent with those described in Cassis (1995
and references therein) and Cassis & Moulds
(1995). The homology and terminology of the
male genitalia follow Kelton (1959) for gross
features, and Stonedahl (1988) for specific
attributes of the endosoma. The homology of the
male genitalia proposed by Kerzhner &
Konstantinov (1999) is not employed in this
work. We use the term endosoma to refer to the
aedeagus distal to the secondary gonopore. The
descriptions of the female genitalia arc based on
Slater ( 1 950) and Davis ( 1 955). Modifications to
terminology for the female genitalia by Schwartz
& Footitt (1998) are used.

Colouration. The colour of the body, particularly
the dorsum is critical in differentiating species in
the Vdnnius complex. Aside from Vanniopsis
crobylos and V rufescens , most taxa are determined
by colour patterns of the dorsum, particularly of
the pronotum and forewings. These patterns have
a generalised form, which are partially charac-
teristic the Vannius complex, and are composed
of a pallid background, interspersed with
species-specific red markings (Fig. 3). We refer
to these patterns as disruptive colouration. In
addition, there are associated red markings of
lateral aspects of the head and the thoracic pleura.
The colour patterning of the second antennal
segment is valuable in discriminating genera and
species, with All either banded (Fig. 3C) or
concolorous (Fig. 3A,B).

Texture. The body of most taxa of the Vannius
complex is smooth, aside from two species of
Vanniopsis (V crobylos and V rufescens) which
have a weakly rugose dorsum.

Vestiture. The vestiture of the Vannius complex is
not significant taxonomically. In general, the
setae are simple, usually bristle-like, and of
sparse to moderate distribution (Fig. 3D).

Head. The head is vertical in orientation (Fig.
1A), with the clypeus, and mandibular and
maxillary plates cone-like, with the bucculae
small and arcuate. The gular is consequently
greatly reduced in size. Gorczyca (1997; figs
11-16) illustrated differences of the head between
genera. There are some species differences found
in Vanniusoides , where the clypeus and man-
dibular plates are apically rounded; cf. V
clypeatus (see Gorczyca & Konstantinov, 2001;
fig. 4) and other Vanniusoides species. The most
significant character examined was the presence
of the longitudinal suture found on the vertex
(and ffons partially), which was either absent
(Austrovannius and Vanniusoides) (e.g. Figs 3 A,

6) or present ( Vanniopsis) (Fig. 2A). In most taxa
of the Vannius complex the antennae are much
longer than the body and thread-like (e.g. Figs
3A,B,D, 6, 13). Antennal length is useful in
differentiating species of Vanniopsis , with V
crobylos and V rufescens (Fig. 3C) having
relatively short antennae (as long as the body),
whereas V. howense have the antennae
significantly longer than the body (Fig. 13). The
length of the first antennal segment is critical in
determining Vanniopsis , having AI much longer
than the head (Figs 3 A-C, 1 3). In other taxa of the
Vannius complex, AI is subequal in length to the
head (Figs 3D, 6, 8). The length of the labium is
not as significant, with most taxa having the
labium reaching between the meso and
metacoxae, aside from Austrovannius , which has
an extremely elongate labium, reaching between
abdominal sternum VII and the genital segments.

Pronotum. The flattened broad collar (Figs 3 A-C,
2A, 13) is diagnostic for the Vannius complex.
The collar is almost vestigial in Austrovannius
(Fig. I A), however, this is likely correlated to
wing reduction. It is noteworthy that Vanniopsis
howense also has staphylinoid wings (Fig. 13),
but possesses a broad flattened collar. Other
pronotal characters of significance include the
campanulate pronotum (Fig. 3 A-C), the undif-
ferentiated disc (without calli) and the bisinuate
posterior margin (Fig. 3A-C).

Thoracic Pleura. Thoracic pleura are largely
uninformative, aside from the metathoracic
spiracle and peritreme of the external efferent
system of the metathoracic glands. The meta-
thoracic spiracle is either concealed (Fig. IB) or
exposed (Fig. 2C). The peritreme is either
medially (Figs 1B-D, 2C) placed on the
metepistemum, or adjacent to the anterior margin
of the metepistemum, as in the remainder of the
Cylapini (Cassis, 1995, fig. 128).

Forewings. Wing polymorphism is diagnostic at
the genus and species levels in the Vannius
complex. Staphylinoidy (sensu Schuh & Slater
1995) is diagnostic for Austrovannius (Figs 3D,
6, 8), Vanniopsis howense (Fig. 13) and
Vanniusoides brevis. All other taxa of the Vannius
complex are macropterous.

Legs. Legs are elongate in the Vannius complex,
and are exaggerated in Austrovannius (Figs 6, 8),
and some species of Vanniopsis (Fig. 13) and
Vanniusoides. Tarsi are uniformly two-
segmented in the Vannius complex. The pretarsus
has spatulate parempodia (Fig. 1E,F), which is
diagnostic and synapomorphic for the Vannius

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SYSTEMATICS OF THE VANNIUS COMPLEX

127

complex. The absence (Fig. 1 E) or presence (Fig.
2F) of subapical teeth is diagnostic at the generic
level, and in some cases the number of teeth
varies (Gorczyca, 1 997).

Male Genitalia. The pygophore is diagnostic for
certain genera. In most taxa the genital opening
of the pygophore is dorsal in orientation (Fig.
1H), or rarely terminal (Fig. 2E). The ventral
surface of the pygophore is impressed along the
midline in Austrovannius (Fig. 1G). The
structures of the aedeagus (Figs 5C, 7C, 9C, 12C,
14C, 15C, 18C) are of primary significance in
determining genera and species in the Vannius
complex. The presence of basal sclerites (Figs
5C, 7C,9C, 12C, 14C, 15C, 18C) and the number
and location of lobal sclerites (Figs 7C, 9C) of the
endosoma are variable among genera, although
their form is consistent within species. The
secondary gonopore is either indistinguishable
( Vannius and Vanniopsis) (Figs 12C, 14C, 15C,
1 8C) or strongly sclerotised and cup-shaped, with
the aperture dentate ( Afrovannius , Austro-
vannius , Paracylapus , and Vanniusoides) (Figs
5C, 7C, 9C).

Female Genitalia. The sclerotised rings (Figs
10A,C, 16A,C,E 19A,C) and associated ventral
and dorsal labiate plates, as well as the
inter-ramal sclerite and associated inter-ramal
lobes (Figs 10B,D, 16B,D,F, 19B,D) are useful
for distinguishing both genera and species of the
Vannius complex. In Austrovannius the
sclerotised rings (Figs 1 0A,C) and posterior wall
(Figs 10B,D) and their associated components
have a complexity of structure not found in either
Vanniopsis and Vanniusoides. The species of the
latter two genera have thin, simple, flattened
rings (Figs 16A,C,E, 19A,C) placed between
separate ventral labiate plates and wide,
sometimes scroll-like, dorsal labiate plates. In
contrast, species of Austrovannius have large,
ornate, asymmetrical rings (Fig. IOC) between a
convoluted, entire ventral plate and strongly
sclerotised, but narrower, dorsal plates. Similarly
the posterior wall of Austrovannius species are
complete (Fig. 10B,D), spanning the base of the
first valvifers, with spinose and bulbous
inter-ramal lobes. Species of Vanniopsis and
Vanniusoides (Figs 16B,D,F 19B,D), in

comparison, have slight, bilaterally separated
inter-ramal sclerite with lobes.

VANNIUS COMPLEX

DIAGNOSIS. The Vannius complex is
recognised by the following combination of
characters: head vertical (Fig. 1A); elongate
thread-like antennae (Fig. 3D); pronotal collar
broad and flat (Fig. 13); elongate metafemora
(Fig. 3B); tarsi two-segmented (Figs 6, 8, 13);
spatulate parempodia (Figs IF, 2F); pretarsus
lacking pulvilli or pseudopulvilli (Fig. IE); and,
endosoma with basal sclerites (strap-like and/or
digitiform) (Fig. 5).

DESCRIPTION. Colouration. Body either
stramineous to mostly red, if pale often with
disruptive colouration (Fig. 3), with red markings
on head, pronotum, thoracic pleura and
forewings. Antennae often concolorous (Fig.
3B), stramineous to brown, sometimes with
reddish highlighting, less commonly with All
banded (Fig. 3B). Legs often pale, sometimes
femora with red banding (Fig. 3B). Abdomen
either stramineous to red, often concolorous, less
commonly dark, sometimes with paler regions
ventral ly.

Texture. Body smooth; impunctate; dorsum
sometimes weakly rugose.

Vestiture. Body with sparse to moderate
distribution of decumbent to semi-erect, pale to
dark, soft to stiff (bristle-like) setae (Fig. 3D)
Structure. Macropterous (Fig. 3A-C) or
staphylinoid (Figs 3D, 6, 8, 13); elongate to
elongate-ovoid. Head subtriangular (Fig. 2A) to
oval; transverse (Fig. 3 A); vertical (Figs 1 A, 2B);
clypeus not visible from above, posteroventrally
directed, usually flat, sometimes weakly swollen,
sometimes with postclypeal processes; vertex
flat to weakly convex (Fig. 2A), sometimes with
weak depressions medially, often with shallow
longitudinal sulcus (Fig. 2A); bucculae short,
arcuate; mandibular plate greatly enlarged,
triangular (Figs 1A, 2B); eyes enlarged,
contiguous with anterior margin of pronotum
(Figs 1A, 2A,B), not significantly extending
beyond plane of head (Fig. 1A); ocelli absent
(Fig. 2A). Antennae elongate, thread-like, at least
as long as the body (Fig. 3D); AI weakly swollen.

FIG. 1. Scanning electron micrographs. A-C, E-G Austrovannius xepenehense, D and H Austrovannius scutica.
A, head, lateral view; B, thoracic pleura, metathoracic glands, lateral view; C, peritreme and evaporative area,
D, peritreme and evaporative area; E, strongly recurved tarsi, ventral view; F, tarsi, lateral view, (J, pygophore,
ventral view; H, pygophore dorsal view. EA = evaporative area, LP = left paramere, P - peritreme, R1 - right
paramere, VR = ventral ridge.

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MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 2. Scanning electron micrographs. Vanniopsis howense. A, head and pronotum, dorsal view; B, head and
pronotum, lateral view; C, thoracic pleura, lateral view; D, forewing, dorsal view; E, pygophorc, dorsal view; F,
pretarsus, ventral view. EES = external efferent system of metathoracic glands, LP = left paramere, ME =
mesepimeron, MT = metepistemum, RP = right paramere, S = metathoracic spiracle, SAT = subapical tooth, SP
= spatulate parempodia.

either subequal in length to head length (Fig. 3D)
or significantly longer (Figs 3A-C, 13), either
cylindrical or bilaterally compressed and
outwardly arcuate (Figs 3B, 13); All cylindrical,
thin, elongate, longer than posterior width of
pronotum (Figs 3, 6, 8, 13); AIII-IV extremely
thin (Fig. 3D), cylindrical, elongate. Labrum
subtriangular, shorter than first labial segment.

Labium usually extending between meso and
metacoxae, rarely elongate, reaching genital
segments; LI longer than bucculae; LI1 often
short. Pronotum transverse; often campanulate
(Fig. 3A-C) with broad flattened collar, or in
wing shortened species collar usually vestigial
(Figs 6, 8), with pronotum ring-like and short;
callosite region absent; posterior margin of

SYSTEMATICS OF THE VANNIUS COMPLEX

129

FIG 3. Habitus photographs. A, Vanniusoides asprokara; B, Vanniusoides melafrons\ C, Vanniopsis rufescens, D,
Austrovannius xepenehense. Scale bar= 1.0mm.

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MEMOIRS OF THE QUEENSLAND MUSEUM

pronotum either bisinuate (Fig. 3A-C) or
rectilinear (Figs 6, 8). Hemelytra: macropterous
species with wings weakly to strongly deflexed at
cuneal fracture; cuneal fracture small to
moderately large; cuneus elongate and usually
narrow; two membrane cells (Fig. 3A,B).
Proepistemum anteriorly directed (Figs 1 A, 2B);
proepimeron flat or medially depressed.
Mesepimeron broadly fused with mesostemum
(Fig. 2C); metathoracic spiracle exposed (Fig.
2C) or concealed (Fig. IB); evaporative bodies
on ventral angle of mesepimeron (Figs 1B-D,
2C). Metepisternum subrectangulate; well
developed external efferent system of meta-
thoracic glands, occupying about 1/3 of sclerite
(Figs 1 B, 2C); evaporative areas contiguous with
mesepimeron; peritreme elliptical, raised
posteriorly (Fig. 1C,D), positioned anteriorly or
medially (Figs IB, 2C) on sclerite, sometimes
more broadly raised, spout-like. Legs elongate
(Figs 3, 6, 8, 13). Metafemora greatly elongate,
tapered distal ly (Fig. 3B, 13). Tarsi two-
segmented (Figs 6, 8, 13). Pretarsi with weakly

(Fig. 2F) to strongly arcuate (Fig. 1 E) claws, with
(Fig. 2F) or without (Fig. IE) subapical teeth;
spatulate parempoida; lacking pul villi or
pseudopulvilli (Fig. IE). Male genitalia:
pygophore conical (Figs IG, 2E), sometimes with
pygophoral processes; genital opening oval, dorsal
(Fig. 1H) or terminal (Fig. 2E) in orientation;
parameres roughly equivalent in size (Fig.
5A,B); sometimes only weakly asymmetrical;
left paramere always largest, usually C-shaped
(Fig. 5A) to weakly arcuate (Fig. 7A), with apex
of shaft hooked (Fig. 5A), sometimes with basal
(Figs 7A) or subapical (Fig. 9A) processes;
aedeagus with membraneous endosoma (e.g.
Figs 5C, 7C 14C, 15C), always with sclerotised
strap-like (often looped) basal sclerites; often
with digitiform basal sclerites (Figs 5C, 7C, 9C),
sometimes with multiple digitiform lobal
sclerites (Figs 7C, 9C), less commonly with
sclerotised, fan-like lobal sclerites (Figs 5C),
with apical margin serrate; secondary gonopore
either undifferentiated (Figs 12C, 14C, 15C,
18C) or with prominent ring-like process with

SYSTEMATICS OF THE VANNIUS COMPLEX

131

dentate aperture (Figs 5C, 7C, 9C); ductus
seminis short; phallotheca short to moderately
large. Female genitalia: sclerotised rings (Figs
10A,C, 16A,C,E, 19A,C) relatively flattened or
slightly folded and sometimes with small lobes
on outside margin of ring; ventral labiate plate
entire or separate, wider than width of ring(s);
dorsal labiate plate reflexed mesially dorsal to
rings or with scroll-like lateral margins; some-
times sclerotised rings and associated structures
asymmetrical; posterior wall (Figs 10B,D,
16B,D,F, 19B,D) with intcr-ramal sclerite entire
or bilaterally separated; sometimes with one to
three pairs of bilaterally separated inter-ramal
lobes; dorsal lobe and lateral lobes absent.

DISTRIBUTION. The Vannius complex is
circumtropical, with centres of endemism in
Madagascar, island archipelagos of the south-
west Pacific (Fiji, New Caledonia, Vanuatu), the
wet tropics of northern Queensland, and the
northern Neotropical region. All of the genera are
restricted to a major zoogeographical region. Ten
species and three genera (Austrovannius,
Vanniopsis and Vatmiusoides) are endemic to the
Australian region (Figs 4, 11, 17).

REMARKS. Gorczyca (1997) erected a new
tribe of Miridae, the Vanniini, for Afrovannius
Gorczyca, Paracylapus Carvalho, Vanniopsis
Poppius, Vannius Distant and Vatmiusoides
Carvalho & Lorenzato, which have spatulate
parempodia and toothed claws. He removed
these genera from the Cylapinae and placed them
w ith the annectant taxon, Palaucoris Carvalho, a
genus for which Carvalho (1956) established a
subfamily with unspecified affinities. Gorczyca
(1997) thereby established the following class-
ification: Palaucorinae: Palaucorini {Palaucoris)
and Vanniini {Vannius complex). In contrast,
Schuh (1975) regarded the palaucorines as a
subtribe of Bryocorinae, and as sister-group to
the Eccritotarsina, within the tribe Eccritotarsini.

Gorczyca (1998) questioned his original
arrangement, comparing the characters of the
Vanniini with selected taxa of Cylapini
( Cylapomorpha migratoria (Distant) and
Phylocylapus lutheri Poppius). He recognised
‘great similarities’ of the body, antennae, tarsi
and colour pattern. However, Gorczyca did not
clearly restore the Vanniini within the Cylapinae,
despite implicitly maintaining their tribal status.
Moreover, he argued that there were alternative
interpretations, based on potential homoplasy of
the observed characters, including the parempodia.

We consider the arrangement of Gorczyca
(1997) to be unsupported based on our
reappraisal. Toothed-claws occur in Cylapinae,
Psallopinae and some Isometopinae (Schuh,
1975, 1976; Schuh & Schwartz, 1984), and is
therefore too general to be considered a synapo-
morphy of Palaucorini + Vanniini. Furthermore,
the claw (with deeply notched subapex) of
Palaucoris , is more elaborate than in the Vannius
complex, and the putative homology is at best
conjectural.

Spatulate parempodia as a synapomorphy for
Palaucorinae sensu Gorczyca also needs
re-analysis. Spatulate parempodia occur within
other higher taxa, such as the Deraeocorinae:
Termatophylini (e.g. Arygrotelaenus Reuter &
Poppius — see Cassis, 1995; figs 89, 95), and
Phylinae (e.g. Arafuramiris Schuh — see Schuh,
1984: fig. 670) but these latter exemplars are not
indicative of suprageneric relationships. How-
ever, the presence of spatulate parempodia in
Palaucorinae sensu Gorczyca does appear to be
invariant and their homology cannot be falsified
at present.

Gorczyca (1997) listed a number of characters
additional to the aforementioned pretarsal
characters, as a basis for his concept of the
Palaucorinae. These include the vertical head and
the short labium. The former character occurs in
many mirid groups, but most significantly in
many of the taxa placed within the Cylapini sensu
Carvalho. This suggests that the Vannius
complex is related to taxa within the Cylapini.
Moreover, Schuh (1976) showed that Bryocorinae:
Eccritotarsini, including Palaucoris , possess a
vertical head.

The length of the labium has limited phylo-
genetic value. In most Miridae the labium extends
between the middle and hind coxae, and are
distinct in comparison to the elongate condition
found in most Cylapinae sensu Carvalho. Gorczyca
(1997) reported that the Palaucorini and Vanniini
possess a short labium. Our observations suggest
that the length of the labium varies considerably
in the Vannius complex, varying from the
mesocoxae to the gential segments. In contrast,
the labium in Palaucoris does not extend beyond
the middle of the mesocoxae.

In summary, we believe that there is not ample
support for the conception of the Palaucorinae as
a subfamily comprising Palaucoris and the
Vannius complex. There is considerable homo-
plasy exhibited in the characters described by
Gorczyca ( 1 997, 1 998). Pending a phylogenetic

132

MEMOIRS OF THE QUEENSLAND MUSEUM

analysis of a larger sample of cylapine taxa we
propose that the Vannius complex be maintained
in the Cylapinae as incertae sedis and that its
relationship with Palaucoris be rejected.

CHECKLIST OF VANNIUS COMPLEX

Austrovannius gen. nov.
platnicki sp. nov.
scutica sp. nov.
xepenehensesp. nov.
Afrovannius Gorczyca, 1997
annulicomis (Poppius, 1 909)
halinae Gorczyca, 1997
mahensis (Distant, 1913)
schmitzi (Gorczyca, 1 996)
Paracylapus Carvalho, 1952
insularis Carvalho, 1952
lestoni (Gorczyca, 1996)
Vanniopsis Poppius, 1909
crobylos sp. nov.
howense sp. nov.
rufescens Poppius, 1909

New Caledonia
Australia (north Qld)
New Caledonia

Madagascar
Sierra Leone
Seychelles
Madagascar

Madagascar

Ghana

Vanuatu
Australia (Lord Howe I)
New Caledonia,
Vanuatu

Vannius Distant, 1 883
crassicornis Poppius, 1 909
oculatus Carvalho, 1955
podager Bergroth, 1 922
rubrovittatus Distant, 1 883

Bolivia
Costa Rica
Brazil
Colombia,
Guatemala, Panama
Vanniusoides Carvalho & Lorenzato, 1978

asprokara sp. nov. F ij i

brevis (Poppius, 1 909) New Guinea

clypeatus Gorczyca & Konstantinov, 2001

Solomon Islands

melafrons sp. nov. Australia (north Qld)

KEY TO THE GENERA OF THE
VANNIUS COMPLEX

1. Labium extending to genital segments; males with
longitudinal ridge on ventral surface of pygophore (Fig.

1G) Austrovannius gen. nov.

Labium extending at most to fourth abdominal segment;
males without ventral ridge on ventral surface of

pygophore 2

2. Head with longitudinal shallow sulcus (Fig. 2A) . ... 3

Head entire, without sulcus (Fig. 6) 4

3. AI significantly longer than head (Figs 3A,B, 13)

Vanniopsis

AI equal to or shorter than length of head (Figs 3D, 6, 8)
Vannius

4. Two subapical teeth on tarsal claws; peritreme of external
efferent system of metathoracic glands medially

oriented on metepistemum Paracylapus

One subapical tooth on each tarsal claw (Fig. 2F);
peritreme of external efferent system of metathoracic
glands anteriorly oriented on metepistemum 5

5. Dorsum with distinct red-orange markings and narrow
black markings on head, clavus and hemelytra; femora

enlarged basally Afrovannius

Dorsum with distinct red-orange markings only; femora
narrow Vanniusoides

Austrovannius gen. nov.

(Figs 1, 3D, 4-10)

ETYMOLOGY For its restriction to the Australian region
and its membership of the Vannius complex.

TYPE SPECIES. Austrovannius scutica sp. nov., by
original designation.

DIAGNOSIS. Staphylinoid (Figs 6, 8); pronotal
collar almost obsolete, at most as an impressed
line (Figs 6, 8); labium reaching between
abdominal sternum VII and genital segments;
pronotum reduced and flattened (Figs 6, 8); claws
strongly arcuate (Fig. 1 E); subapical teeth absent
(Fig. IE); pygophore with longitudinal medial
ridge on ventral surface (Fig. 1G); endosoma
with basal and lobal sclerites (Figs 5C, 1C, 9C);
sclerotised rings (Fig. 10A,C) asymmetrical,
large, and with marginal processes; and, posterior
wall (Fig. 1 0B,D) with inter-ramal lobes.

DESCRIPTION. Colouration. Body reddish to
brown, sometimes with disrupted red markings
on dorsum (Figs 3D, 8), lateral aspects of head
and thoracic pleura. Metafemora sometimes with
red banding. Abdominal venter stramineous to
red-fuscous.

Vestiture. Body with sparse to moderate density
of soft to stiff (bristle-like) semi-erect to
decumbent, simple setae (Fig. 3D).

Texture. Body smooth.

Structure. Small; staphylinoid (Figs 3D, 6, 8);
ovoid. Head: vertex flat to weakly convex (Fig.
1A), sometimes with submedial weak depress-
ions. Antennae thread-like, elongate, much
longer than body, at least 1 .5 x longer (Fig. 3D);
AI equal to head length (Figs 6, 8), weakly
expanded; All elongate (Figs 6, 8), cylindrical,
narrow, a little broader than AIII-A1V; AIII-AIV
elongate, very narrow. Labium elongate, extend-
ing between abdominal sterna VII and genital
segments. Pronotum (Figs 6, 8) transverse,
subrectangular, ring-like, small, flat; collar
vestigial, at most as thin crease; remainder of
pronotum uniform, not subdivided into callosite
region and disc; posterior margin rectilinear.
Mesoscutum not visible (Figs 6, 8). Scutellum
triangular, transverse, small, shorter than
pronotum, flat (Figs 6, 8). Proepimeron weakly
depressed medially (Fig. 1A). Mesepimeron:
metathoracic spiracle (Fig. 1B-D) not exposed;
ventral angle of mesepimeron with evaporative
bodies. External efferent system of metathoracic
glands (Fig. 1 B-D) occupying a little more than a
third of metepistemum; peritreme elliptical,
moderately tumid and raised, positioned

SYSTEMATICS OF THE VANNIUS COMPLEX

133

medially on metepistemum, not reaching lateral
margin of evaporative area. Metafemora greatly
elongate, tapered distally. Pretarsal claws (Fig.
1E,F) strongly arcuate, subapical teeth absent.
Male genitalia: pygophore subconical (Fig.
1G,H), with longitudinal medial ridge on ventral
surface; genital opening large, ovoid, dorsally
oriented; sometimes with flange-like processes
dorsoanteriorly to paramere insertions; para-
meres simple; left paramere (Figs 5A, 7A, 9A)
C-shaped to subtriangular, usually elongate,
sometimes with basal or subapical process; right
paramere (Figs 5B, 7B, 9B) either short or
elongate; phallotheca large, subconical;
aedeagus (Figs 5C, 7C, 9C) with membraneous
endosoma, most often with multiple lobal
sclerites, rarely without lobal sclerites, most
lobal sclerites digitiform, tapered apically, often
with fan-like lobal sclerites, with apices serrate;
strap-like basal sclerites present, often with
additional digitiform processes; secondary gono-
pore large, sclerotised, with dentate aperture.
Female genitalia: sclerotised rings (Fig. 10A,C)
asymmetrical, moderately large to large, folded
with pointed processes on lateral margins;
ventral labiate plate entire, convoluted; dorsal
labiate plate well-sclerotised not as wide laterally
as ventral labiate plate; posterior wall (Fig.
10B,D) with inter-ramal sclerite entire and
anterior surface with one to three pairs of bilateral
inter-ramal lobes.

DISTRIBUTION AND HABITAT. Austro -
vannius is restricted to the southwest Pacific (Fig.
4), and is known from New Caledonia, the
Loyalty Islands, and tropical Queensland
(Bellenden Ker Range). The three described
species are known from leaf litter in rainforests.

REMARKS. Austrovannius is the most
distinctive genus of the Vannius complex, and has
a number of notable autapomorphies, including,
the short ring-like pronotum (collar vestigial)
(Figs 6, 8), ridge-like ventral midline of the
pygophore (Fig. 1G), the elongate labium (almost
or reaching genital segments), the strongly
arcuate tarsal claws, and the elaborate male and
female genitalia. The species are all very small,
less than 2mm in length, and have an ovoid body,
often with disruptive colouration (Fig. 3D). The
male genitalia are the most intricate in the
Vannius complex with the endosoma bearing
both basal and lobal sclerites (Figs 5C, 7C, 9C).
In addition, the secondary gonopore is strongly
sclerotised with a dentate aperture.

As mentioned above the genus contains many
undescribed species and are the subject of
another paper.

KEY TO THE SPECIES OF
AUSTROVANNIUS

1. Pronotum and forewings usually concolorous (Fig. 6),

cither brown or reddish 2

Pronotum and forewings stramineous with patterned red

markings (Figs 3D, 8) 3

2. Pronotum and forewings brown; left paramere (Fig. 5A),

without basal process apex hooked: endosoma (Fig. 5C)
with two sickle-shaped spiculi attached below secondary
gonopore, and two apical fan-like lobal sclerites; New

Caledonia platnicki

Pronotum and forewings stramineous-red; left paramere
(Fig. 7A) with basal process, apex not hooked;
endosoma (Fig. 7C) without sickle-shaped spiculi or
fan-like lobal sclerites; North Queensland . . scutica

3. Metafemora mostly brown with apex stramineous;

endosoma (Fig. 5C) with two basal spiculi, without lobal
sclerites; New Caledonia platnicki

Metafemora mostly stramineous with submedial and
subapical red bands (Figs 3D, 8); endosoma (Fig. 9C)
with digitiform and fan-like lobal sclerites, without basal
spiculi; Loyalty Islands xepenehense

Austrovannius platnicki sp. nov.

(Figs 4, 5)

ETYMOLOGY. In honour of Dr Norman Platnick, who
was one of the collectors of the type series.

MATERIAL. NEW CALEDONIA. Holotype, <5, Mount
Koghis, 22°irS 166 0 3TE, 500m, 23-30 May 1987. R
Raven and N Platnick. rainforest pitfalls (MNHP);
Paratypes: 6 <5 <5, same data as holotype (AM, QM);
M29 9 Mount Koghis, 22°irS 166°0rE, 500m, 22
November 2000, GB Monteith rainforest sieved litter
(QM); lc539 9, Mount Koghis, 22° ITS 166 o 0rE,500m.
22 November 2000, GB Monteith, pyrethrum trunks and
logs, 9931 (QM).

DIAGNOSIS. Pronotum and forewings most
often stramineous-brown; labium uniformly
stramineous, reaching abdominal sternum VII;
thoracic pleura dark brown; metafemora dark
brown; tibiae stramineous; left paramere (Fig.
5A) sickle-shaped, without basal process, shaft
broadly and evenly arcuate; endosoma (Fig. 5C)
without lobal sclerities, with pair of bifurcate
basal sclerites, two basal arcuate spiculi and two
apical fan-like plates.

DESCRIPTION. Colouration. Body mostly
stramineous brown, rarely with red markings on
dorsum. Head stramineous to yellow-brown;
vertex with an obscure brown (rarely red) sub-
triangular marking; mandibular and maxillary
plates, bucculae, genae and gular mostly
stramineous, rarely mostly red, with fuscous

134

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Austrovannius platnicki. A, left paramere; B,
right paramere; C, aedeagus. BS = basal sclerites,
BSP = basal spiculum, DLS = digitiform lobal
sclerite, FLS = fan-like lobal sclerite, M =
membaneous sac, P = phallotheca, SG = secondary
gonopore. Scale bar = 0. 10mm.

highlighting. Labium stramineous to stramineous-
brown. Antennae mostly stramineous. All with
an apical whitish annulation. Pronotum
stramineous-brown, laterally darker, sometimes
laterally red. Thoracic pleura stramineous-brown,
sometimes with ventral margin of propleura
paler, rarely more uniformly red with fuscous
highlighting. Scutellum stramineous, rarely
anterior margin with red tinge. Forewings mostly
stramineous-brown, rarely with disruptive
reddish markings. Legs mostly stramineous-brown;
coxae brown; fore and mesofemora stramineous;
hind femora darker brown, with apices
stramineous; tibiae and tarsi stramineous.

Abdomen mostly brown, sometimes pygophore
paler ventral ly.

Structure. Labium reaching abdominal sternum
VII. Male genitalia: left paramere (Fig. 5A)
sickle-shaped, without basal process, shaft
broadly and evenly arcuate; right paramere (Fig.
5B) arcuate, subequal in size to left paramere,
apex weakly angulate; endosoma (Fig. 5C) with
two major membraneous lobes, two basal arcuate
spiculi, one with apex acute, other with apex
serrate, secondary gonopore sclerotised, with
aperture dentate, with pair of bifurcate digitiform
basal sclerites, with two apical fan-like lobal
sclerites, without digitiform lobal sclerites.
Female genitalia not examined.

Measurements. 3 cJ 6 BL 1 .49- 1 .92, H W 0.6 1 -0 62
IOD 0.28-0.30, HL 0.34-0.37, PL 0.15-0.17, PW
0.64-0.66, All 0.98-1.06, LL 0.87-0.96; 19 BL
1.88, HW 0.63, IOD 0.30, HL 0.36, PL 0.17 PW
0.66, All 1.01, LL 1.10.

DISTRIBUTION AND HABITAT. Mid-altitude
rainforest in southern New Caledonia, from a
single location (Mt Koghis) (Fig. 4). The species
is epigaeic, with most specimens collected in
pitfall traps or from sieved litter. A series was
collected by pyrethrum fogging, from trunks and
logs.

REMARKS. Austrovannius platnicki is unlike
the other species of the genus, in lacking
digitiform lobal sclerites. The endosoma (Fig.
5C) is complex however, in possessing bifurcate
basal sclerites, apical fan-like lobal sclerites and
basal spiculi. The genitalia are most like those of
A. xepenehense , in possessing fan-like lobal
sclerites and basal processes, but the latter is
differentiated by having lobal sclerites (cf. Figs
5C and 9C). The parameres of these species are
also alike, with both having C-shaped parameres
(cf. Figs 5A.B and 9A,B).

These above species are often distinguished
externally by colour pattern differences, with
most specimens of A. platnicki uniformly brown,
and A. xepenehense with red markings on the
dorsum. A single series of A. platnicki (reference
number 993 1 ) has red markings, similar to those
found in A. xepenehense , but the male genitalia
are clearly conspecific with the other Mt Koghis
specimens of A. platnicki. Precedence is given to
the value of the male genitalia in species level
decision-making, because of the complexity of
the structures of the endosoma. The species
description of A. platnicki is primarily based on
the ‘Raven and Platnick’ material because there

SYSTEM ATICS OF THE VANNIUS COMPLEX 135

FIG 6. Habitus of Austrovannius scutica.

are more specimens and they are in superior
condition. The three series were collected at
different periods (May and November), which
suggests that this species is multivoltine and
colour polymorphism may be attributable to
generational factors. The female specimens were
not examined because there is some doubt about
their conspecificity.

Austrovannius scutica sp. nov.

(Figs 1D-H, 4, 6, 7, 10A,B)

ETYMOLOGY. Latin scutica , referring to the extremely
long whip-like antennae.

MATERIAL. QUEENSLAND. Holotype, <5, Bellenden
Ker Range, 1km S of Cable Tower 6, 17°9’36"S
145 D 3r48”E, 500m, 17-24 October 1981, 500 metres,
Earthwatch and Queensland Museum Survey, rainforest
sieved litter, QM Berlesate No. 319, GB Monteith (QM;
Registration no. QM T 1086 14). Paratypcs: 2 6, same data
as holotype, QM Berlesate Nos 3 1 5 and 320 (QM); 2 6 6 ,
Bellenden Ker Range, Cableway base station, [17°9 , 36"S
145 0 32’24'’E], 100m, 17 October-9 November 1981,
Earthwatch and Queensland Museum survey, rainforest
pitfall trap, GB Monteith (AM; QM).

DIAGNOSIS. This species is distinguished by
the following combination of characters:
pronotum and forewings red; first labial segment
red, LII-IV stramineous; thoracic pleura mostly
uniformly red, sometimes more reddish-fuscous;
left paramere (Figs 7A) subtriangular, with basal
process, apex hooked; right paramere (Fig. 7B)
subtriangular with apex hooked; endosoma (Fig.
1C) with seven lobal sclerities; and, posterior
wall (Fig. 1 0B) with one pair of widely separated
dorsal inter-ramal lobes.

DESCRIPTION. Colouration. Body (Fig. 6)
mostly red to reddish-brown with stramineous
patches on head, ventral surface of body mostly

FIG. 7. Austrovannius scutica. A, left paramere, lateral
view; B, left paramere, mesial view; C, right
paramere; D, aedeagus. Scale bar = 0. 1 0mm.

red, sometimes with stramineous or fuscous high-
lighting. Head stramineous to yellow-brown;
posterior margin of vertex occasionally more
enbrowned; mandibular plate yellow-brown to
fuscous-red; maxillary plate, bucculae, genae
and gular most often red. Labium: LI red; LII-IV
stramineous. Antennae mostly stramineous; AI
sometimes with small red spots basally; AII-AIV
sometimes weakly enbrowned. Pronotum: red to
reddish-fuscous. Thoracic pleura red or
reddish-fuscous. Scutellum stramineous-red to
stramineous-fuscous. Forewings mostly
stramineous-red, rarely reddish-fuscous. Legs

136

MEMOIRS OF THE QUEENSLAND MUSEUM

mostly stramineous; coxae stramineous-red; fore
and mesofemora stramineous; metafemora
red-brown with stramineous annulations on
distal third and apex; tibiae and tarsi stramineous.
Abdomen mostly red; male pygophore
sometimes stramineous-red ventrally.

Structure. Habitus (Fig. 6). Labium reaching
genital segments. External efferent system of
metathoracic glands (Fig. ID). Pretarsus (Fig.
1E ; F). Male genitalia: pygophore (Fig. 1G,H);
left paramere (Fig. 7 A) small, subtriangular. with
acute hook-like basal process, apex of shaft
hooked; right paramere (Fig. 7B) small, elongate,
also with apical and basal hook-like processes;
endosoma (Fig. 1C) enlarged, with multilobed
membranous sac, one lobe of sac with four
elongate lobal sclerites, without basal arcuate
spiculi or apical fan-like lobal sclerites. Female
genitalia: (Fig. 10A,B) sclerotised rings and
labiate plates (Fig. 10A) moderately large, rings
attenuated mesially, widely separated, folded,
and adhering ventral plate anteriorly; ventral
labiate plate entire, anterior margin thickened;
dorsal labiate plate obscure; inter-ramai sclerite
of posterior wall (Fig. 10B) with ventral margin
attenuated, flanking minutely spinose mesial
region, with two widely separated, bilaterally
sculpted inter-ramai lobes dorsally.

Measurements. 56 6 BL 1.20-1.40, HW 0.55-0.56,
IOD 0.24-0.27, HL 0.27-0.34, PL 0.15-0.18, PW
0.54-0.61, All 1.18-1.29, LL 0.92-1.10; 1$ BL
1.58, HW 0.57, IOD 0.26, HL 0.37, PL 0.16, PW
0.63, All 1.15, LL 1.16.

DISTRIBUTION AND HABITAT. Mid-altitude
rainforest in the wet tropics of Queensland
(Bellcnden Ker Range) (Fig. 4). Epigaeic, with
specimens collected either by sieving litter, or in
pitfall and flight intercept traps. A single
specimen was collected by pyrethrum fogging.

REMARKS. Austrovannius scutica is distinguished
from A. platnicki and A. xepenehense by
differences in the shape of the parameres (cf. Figs
5A,B, 7A,B and 9A.B) and structure and size of
the endosoma (cf. Figs 5C and 9C). Unlike its
congeners, A. scutica , has a distinct left paramere
(Figs 7 A); with a basal process and the shaft not
broadly arcuate. The endosoma (Fig. 7C) of the
latter species is diagnostic, and is small, lacks
arcuate spiculi and apical fan-like lobal sclerites,
and has seven elongate digitform lobal sclerites.
A. scutica is also more uniformly red in colour.

FIG. 8. Habitus of Austrovannius xepenehense.

Austrovannius xepenehense sp. nov.

(Figs 3D, 4, 8,9, 10C,D)

ETYMOLOGY. Types from Xepenehe.

MATERIAL. LOYALTY ISLANDS: Holotype, <J, Lifou
Island, Xepenehe, 20°47’S 167°11'E, 20m, 6 December
2000, GB Monteith, rainforest sieved litter (MNHP).
Paratypes: 4 <3 6 1 9 , same data as holotype (AM; QM).

DIAGNOSIS. Dorsum stramineous with disruptive
red markings (Figs 3D, 8); first antennal segment
with reddish highlighting; propleura with two red
transverse bands; forewings with ten red mark-
ings; metafemora stramineous with submedial
and subapical red bands; left paramere (Fig. 9 A)
with subapical triangular process; endosoma
(Fig. 9C) with four digitiform and two fan-like
lobal sclerites, and basal sclerites, without basal
spiculi; sclerotised rings (Fig. 10C) large, folded,
with conspicuous lateral lobes; posterior wall
(Fig. 10D) with three bilaterally paired
inter-ramai lobes on anterior surface of complete
inter-ramai sclerite .

DESCRIPTION. Colouration. Body yellow-
brown to stramineous, with red markings on
dorsum (Fig. 3D, 8). Head mostly stramineous;
vertex with reddish markings; frons and genae
with red markings, often bounding antennifers;
postclypeus with reddish tinge; genae posteriad
of maxillary plate with red spot. Labrum mostly
stramineous, sometimes with reddish high-
lighting. Labium: LI mostly red, remainder
stramineous to stramineous-brown, with apex of
LIV with fuscous highlighting. Antennae: AI
stramineous with reddish highlighting (Fig. 3D);

SYSTEMATICS OF THE VANNIUS COMPLEX

137

FIG 9. Austrovannius xepenehense. A, left paramere;

B, right paramere; C, aedeagus. Scale bar = 0. 1 0mm.

AII-A1V dark brown, apex of All with whitish
apical band. Pronotum mostly stramineous with
reddish markings (Fig. 3D); collar with medial
red transverse marking; disc laterally and
medioposteriorly with red markings. Scutellum
most often stramineous, sometimes with reddish
highlighting. Forewings with 10 red markings
(Fig. 3D); each wing with pair sub-basally, pair
submedially, single spot apical ly. Proepistemum
fuscous-red; proepimeron with two transverse
bands of fuscous-red. External efferent system of
metathoracic glands stramineous, remainder of
metepistemum red to fuscous-red. Legs mostly

FIG 10. Female genitalia. Austrovannius scutica A,
sclerotised rings; B, posterior wall; Austrovannius
xepenehense ; C, sclerotised rings; D, posterior wall.
IRL = inter-ramal lobe, IRS = inter-ramal sclente, SR
= sclerotised ring, VLP = ventral labiate plate. Scale
bar = 0.25mm.

stramineous to light-brown, sometimes with red
highlighting; metafemora with submedial and
subapical red bands (Fig. 3D); tibiae often
stramineous-red.

Structure. Habitus (Figs 3D, 8). Labium reaching
abdominal sternum VII. Male genitalia:
pygophore with small flange-like processes
anterodorsally to paramere insertions; left
paramere (Fig. 9A) C-shaped, with subapical
triangular process, shaft evenly arcuate with apex
hook-like; right paramere (Fig. 9B) sublinear
with apex angulate; endosoma (Fig. 9C) bilobed,
each subtended by basal processes, one a
multifurcate process with four arms, the other
with an elongate digitform process; four elongate
digitform and one fan-like (with serrated apex)
lobal sclerites. Female genitalia: sclerotised rings
(Fig. 10C) large, folded, narrowly separated,
attenuated mesially, anterior margin asym-
metrically convoluted and attached to dorsal
labiate plate; posterior wall (Fig. 1 0C) with three
bilaterally paired inter-ramal lobes on anterior
surface of complete inter-ramal sclerite, dorsal
pair pointed, prominently projecting above
dorsal margin of inter-ramal sclerite equal to
height of sclerite, middle pair pointed, short,
ventral pair blunt with sculptured surface.

Measurements. 4<J 6 BL 1 .35- 1 .50, HW 0.58-0.60,
IOD 0.29-0.30, HL 0.31-0.32, PL 0.15-0.17, PW
0.63-0.65, All 1.10-1.20, LL 0.98-1.03; 19 BL
1.75, HW 0.65, IOD 0.30, HL 0.33, PL 0.19, PW
0.68, All 1.00, LL 1.05.

138

MEMOIRS OF THE QUEENSLAND MUSEUM

9

o

Vanuatu

vi

Fiji

9b

o.

K

New Caledonia \\

; 5>

fcs *

O

Vanniopsis
• crobylos
■ howense

■

Lord Howe Island

•

Norfolk Island

▲ rufescens

FIG 1 1 . Distribution map of Vanniopsis.

DISTRIBUTION AND HABITAT. From two
lowland rainforest sites on Lifou Island (Loyalty
Islands) (Fig. 4), amongst leaf litter, and on tree
trunks and fallen logs. These specimens have
been collected near sea level.

REMARKS. The species is noteworthy for its
disruptive colouration (Fig. 3D) and distinctive
male aedeagus (Fig. 9C). The endosoma
resembles A. platnicki in possessing prominent
basal processes and apical fan-like lobal sclerites
(ct. Figs 5C and 9C). It differs from it by having
digitiform lobal sclerites, this latter condition
also occuring in A. scutica. The left paramercs of
A. xepenehense and A. platnicki are alike in being
C-shaped, but the former species has a small
subapical triangular process (cf. Figs 5 A and 9 A).

Vanniopsis Poppius, 1909

Vanniopsis Poppius, 1909: 17 (sp. nov.); Carvalho, 1957: 33
(catalogue); Schuh. 1995: 39 (catalogue); Gorczyca, 1997:
519, 540-542, figs 8 and 16 (description).

TYPE SPECIES. Vanniopsis rufescens Poppius, 1909, by
original designation.

DIAGNOSIS. Head subtriangular (Figs 2A, 13);
firms prominent (extending beyond eyes); AI
longer than head (Figs 3C, 13); AI compressed
and arcuate, much longer than head, with dense
distribution of bristle-like setae; All banded (Fig.
3C); pronotal collar flat and broad (Fig. 3C, 13);
tarsal claws weakly arcuate (Fig. 2F); subapical
teeth present (Fig. 2F); endosoma (Figs 12C,
14C, 15C) membraneous, usually with paired
looped strap-like basal sclerites, secondary
gonopore undifferentiated; phallotheca short;
parameres strongly asymmetrical; left paramere
(Figs 12 A, 14A, 15 A) large, C-shaped, tapered
apically; right paramere (Fig. 16A,C,E) smaller
than left paramere, sublinear, apex of shaft
tapered; sclerotised rings (Figs 12A, 14A, 15 A)
widely separated, ovoid, flattened mesially; and,
posterior wall (Fig. 1 6B,D,F) simple, with small,
paired inter-ramal sclerites, without inter-ramal
lobes.

DESCRIPTION. Colouration. Head and
pronotum mostly brown, with forewings either
reddish, or stramineous to brown with reddish to
orange-red markings (Figs 3C, 13). Head often

SYSTEMATICS OF THE VANNIUS COMPLEX

139

brown, with posterior margins of vertex with
reddish markings; frons often pale yellow to light
brown; clypeus either pale yellow to red;
bucculae, mandibular and maxillary plates, and
gcnae either pale yellow or red. Antennae: AI
either mostly stramineous with reddish markings
or red; All banded (Fig. 3C), mostly brown with
basal, medial and apical yellow bands; AIII
mostly brown with apical yellow band; AIV
brown. Labium mostly stramineous to
stramineous-orange, with LI sometimes with red
highlighting, LIV sometimes with fuscous
highlighting. Thoracic pleura mostly red,
sometimes propleura more brown. Legs mostly
stramineous; metafemora with subapical reddish
marking. Pronotum mostly brown, sometimes
with longitudinal medial stramineous line.
Scutellum mostly brown, sometimes apex
stramineous. Forewings (Fig. 3C) either mostly
red to orange-red with stramineous to hyaline
regions on corium or in staphylinoid species
brown with four reddish markings; cuneus red or
orange-red with apex red; membrane fiimose.
Abdominal venter red to orange-red.

Vestiture. Dorsum usually with short dark
semi-erect, bristle-like setae. Frons sometimes
with dense distribution of elongate semi-erect
setae (Figs 3C, 13).

Texture. Mostly smooth, with forewings often
weakly rugose.

Structure. Macropterous (Fig. 3C) or staph-
ylinoid (Fig. 13); elongate to elongate-ovoid.
Head: (Fig. 2A,B) vertex weakly convex to flat,
with shallow longitudinal sulcus. Antennae
elongate, thread-like, subequal (Fig. 3C) to or
longer than body (Fig. 13); AI (Figs 3 C, 13)
thickened, bilaterally compressed, arcuate,
longer than head length; AI longer than AIII.
Labium extending to metacoxae. Pronotum
transverse; campanulate or rectangulate; collar
broad, flat. Mesoscutum exposed or concealed.
Scutellum flat to weakly convex, subequal in
length to pronotum. Hemelytra: corium broad,
moderately convex; cuneus elongate. Thoracic
pleura: (Fig. 2C) proepimeron flat to weakly
depressed medially; metathoracic spiracle ex-
posed, bounded by evaporative bodies; external
efferent system of metathoracic glands
moderately developed, occupying about l/3rd of
segment; peritreme tumid, orientated anteriorly
to medially, not reaching dorsal margin of
evaporative area, sometimes peritreme strongly
raised and tumid, and external efferent system
spout-like. Legs: metafemur elongate (Fig. 3C)
and tapered distally; tarsal claws weakly arcuate;

claws with subapical teeth. Male genitalia: left
paramere (Figs 12A, 14A, 15A) large, C -shaped,
tapered apically; right paramere (Figs 14B, 15B)
smaller, sublinear, apex of shaft tapered;
endosoma (Figs 12C, 14C, 15C) membraneous,
with or without sclerotised spinose fields, without
lobal sclerites, usually with paired looped strap-
like basal sclerites supporting membrane; sec-
ondary gonopore undifferentiated; phallotheca
short. Female genitalia: sclerotised rings (Figs
1 6A,C,E) weakly differentiated, widely
separated, ovoid, flattened mesial ly; posterior
wall (Figs 16B,D,F) simple, with small, paired
separated narrow, lunate-shaped inter-ramal
sclerites, without inter-ramal lobes.

DISTRIBUTION. New Caledonia, Vanuatu and
Lord Howe Island (NSW, Australia) (Fig. 11).

REMARKS. Vanniopsis is best differentiated by
its first antennal segment and the aedeagus. The
most distinctive feature of this genus is the
arcuate, elongate (much longer than head) and
bilaterally compressed first antennal segment
(Figs 3C, 13), which also has dense bristle-like
decumbent setae. In other taxa of the Vannius
complex, the first antennal segment is a little
larger or subequal to the head length, and is never
compressed and arcuate, or has the vestiture
described above.

There are superficial reasons for erecting a new
genus for V. howense , because of its morph-
ological distinctiveness; namely, the staphylinoid
body and correlated reductions of the pronotum
and scutellum, as well as the elongate antennae
(Fig. 13). However, this species is clearly
congeneric with its macropterous relatives, based
on the aforementioned characters (first antennal
segment and aedeagus).

KEY TO THE SPECIES OF VANNIOPSIS

1. Macropterous (Fig. 3C); antennae subequal in length to

the body; genital opening of male pygophore dorsal in

orientation * 2

Staphylinoid (Figs 2D, 13); antennae significantly longer
than body; genital opening of male pygophore terminal
in orientation (Fig. 2E); Lord Howe Island . . howense

2. Frons densely setate; clypeus, mandibular and maxillary

plates stramineous; left paramere with subapical flange
(Fig. 12A); three endosomal basal sclerites, not looped
(Fig. 12C); Vanuatu crohylos

Frons with sparse distribution of short decumbent setae;
clypeus, mandibular and maxillary plates red; two pairs
of looped endosomal basal sclerites (Fig. 15C); New
Caledonia, Vanuatu rufescens

140

MEMOIRS OF THE QUEENSLAND MUSEUM

Vanniopsis crobylos sp. nov.

(Figs 11, 12, 16A,B)

ETYMOLOGY. Greek hvbylos, referring to the tuft of
hair on the head.

MATERIAL. VANUATU. Holotype, 6 Malekula
I[sland], South West Bay, [16°3(TS 167°26’E], 300-350m,
2 October 1971, P. Cochereau Roy[al] Socjiety] Percy
Sladen Expedition, beating trees and sweeping grasses,
forest (SAM A). Paratypes. 9 same data as holotype
(SAMA); 19, Espiritu Santo I[sland] (SW), below
Namatasopa, [15°3rs 166°49'E], 250m, 1 September
1957, JL Gressitt (BPBM); 19, Elate I[sland] (NW),
Limestone Plateau N of Maat. [17.6833°S 168.25°E],
100m, 19 August 1957, JL Gressitt (BPBM); 1 9, Santo
I[sland], lakabone [Iakobone] N of Port Olry, [15°0rS
167°03’E], 40-200m. 21-22 October 1981, JL Gressitt on
Elastostemma [sic] (BPBM)

DIAGNOSIS. Macropterous; forewings rugose;
frons densely hirsute; vertex with shallow
longitudinal sulcus; AI densely hirsute; collar
present; All banded; pronotum campanulate;
posterior margin of pronotum bisinuate; meso-
scutum exposed; peritreme raised, anteriorly
oriented; endosoma with three basal sclerites,
membrane reduced (Fig. 12C); and, left paramere
(Fig. 12A) with subapical flange.

DESCRIPTION. Colouration . Body mostly pale
red, with stramineous to stramineous-orange
markings. Head brown, with posterior margin of
vertex with red markings; clypeus, mandibular
and maxillary plates, genae and gular stram-
ineous, sometimes genae posterior to eyes red.
Antennae: A I red, sometimes stramineous-red;
All banded, dark brown with medial and apical
stramineous bands, with base minutely stram-
ineous; All I mostly brown with apical yellow
band; AIV brown. Labium stramineous to
yellow-orange, apex of IV sometimes enbrowned.
Pronotum brown. Thoracic pleura red.
Mesoscutum brown to red-brown. Scutellum
stramineous-brown to orange-brown. Hemelytra
mostly red, sometimes with exocorium broadly
stramineous and translucent; cuneus mostly red,
with apex and medial angle stramineous; mem-
brane furnose, veins red. Legs mostly stramineous;
coxae stramineous to red-stramineous; meta-
femora with subapical red marking. Abdomen
mostly red; ovipositor stramineous to stramineous-
brown.

Texture. Pronotum and hemelytra rugose.

Vestkure. Dorsum with moderate distribution of
simple, fine, erect, elongate, dark setae. Frons
with a dense tuft of elongate bristle-like setae.

FIG 1 2. Vanniopsis crobylos. A, left paramere; B, right
paramere; C, aedeagus. Scale bar = 0.10mm.

Structure. Macropterous; elongate. Antennae
subequal in length to body. Labium extending to
apices of metacoxae. Pronotum campanulate;
collar broad, anterior margin rectilinear;
posterior margin bisinuate. Mesoscutum
exposed. Scutellum as long as wide. External
efferent system of metathoracic glands spout-
like; peritreme anteriorly oriented. Hemelytra:
strongly depressed at corial fracture; cuneus
narrow and elongate. Male genitalia: pygophore
subconical, genital opening large, ovoid, dorsally
directed; left paramere (Fig. 12A) small,
elongate, subrectangular, broadly expanded
subapically, flangelike with hook-like apex; right
paramere (Fig. 12B) small, subrectangular,
lateral margin moderately emarginate, acute
apical process coplanar with remainder of shaft;
endosoma (Fig. 12C) with three strap-like basal
sclerites supporting thin membranous sac,
secondary gonopore undifferentiated. Female
genitalia: sclerotised rings (Fig. 1 6A) moderately

SYSTEMATICS OF THE VANNIUS COMPLEX

141

small, semi-elliptical, mesial margin flattened;
ventral labiate plate separated, convex anteriorly,
flattened mesially; dorsal labiate plate scroll-like
laterally; posterior wall (Fig. 16B) with
bilaterally paired, thin inter-ramal sclerites on
dorsal margin.

Measurements. 1 6 BL 3.82, HW 0.68, IOD 0.26,
HL0.38, PL 0.50, PW 0.95; 4$ 9 BL 3.74-4.01,
HW 0.62-0.66, IOD 0.24-0.26, HL 0.37-0.41, PL
0.50-0.60. PW 1.00-1.07, All 1.15-1.24, LL
0.90-1.24.

DISTRIBUTION AND HABITAT. From three
islands of Vanuatu (Fig. 11). A single specimen
has been collected on Elatostema sp. [as Elasto-
stemma] (Urticaceae). The two specimens from
Malekula Island were taken from vegetation.
These data suggest that V crobylos are
‘up-on-plants’, and occupy a different habitat
than V howense, a known epigaeic species.

REMARKS. V crobylos and V rufescens are
morphologically alike, with both species being
macropterous and primarily red in colour. The
former species can be recognised readily by the
hirsute frons and the pale ventral lateral aspect of
the head. The male genitalia are significantly
different, with the aedeagus of V crobylos
reduced, whereas the endosoma of V rufescens
(cf. Figs 12C and 15C) is an enlarged multi-
furcate membranous sac, that has paired looped
basal sclerites and spinose fields. The latter
endosomal condition and the parameres are most
similar to those found in V howense (cf. Figs 1 2 A
and 14 A).

Vanniopsis howense sp. nov.

(Figs 2, 11, 13, 14, 16C,D)

ETYMOLOGY. For its geographical location.

MATERIAL. LORD HOWE ISLAND. Holotype, 6 , base
of Intermediate Hill, 31°33’S 159°4’48"E, 6 December
2000, G Cassis (AM). Paratypes: 46 6 2 9 9 11 juveniles,
same data as holotype; 1 6 2 9 9 2 juveniles, Goat House
track at creek bed, 31°33'S 159°4 , E, 5 m, 8 December
2000. G Cassis, ex fallen leaves of Howea fosteriana
(AM); 6 juveniles, base of ‘Round Face’ Mt Lidgbird, Far
Flats, 31 34’9"S 159V35"E, 27 November 2000, CBCR
survey, ex leaf litter. Broad Megaphyllous Closed
sclcrophyll forest - Howea belmoryana habitat, site
LHI36L (AM); 2 juveniles, N bank of Rocky Run Creek at
junction of W costal trail to Boat Harbour, 31 33’19"S
159 5 , 33"E, 21 November 2000, CBCR survey, ex leaf
litter. Broad Megaphyllous Closed sclcrophyll forest
Pandamis habitat site LHI24L (AM); 2 juveniles, S end of
Salmon Beach, vicinity of Little Island, 31 34’8 M S
1 594’28 M E, 27 November 2000, CBCR survey, ex leaf
litter, Broad Megaphyllous Closed sclerophyll forest -

FIG. 13. Habitus of Vanniopsis howense.

Howea fosteriana habitat site LHIS35L(AM)* 1 juvenile,
Mount Gower walking track. 31 35’12”S 159 4’35"E, 28
November 2000, CBCR survey, ex leaf litter, dosed
gnarled Mossy Forest - Bubbia/Dracophyllum , site
LHI50L (AM); 26 6 5 9 9 , behind Leanda-Lei, [3 1°53’S
1 59°07'E], 45 m, 6 November 1979, GB Monteith, ex leaf
litter, calcareous soil Broad Megaphyllous Closed
sclerophyll forest - Howea fosteriana habitat (QM).

DIAGNOSIS. Staphylinoid (Figs 2D, 13);
forewings stramineous with red disruptive
colouration; frons sparsely setate; antennae
significantly longer than body (Fig. 13); pro-
notum (Fig. 2A) rectangulate, posterior margin
rectilinear; pronotal collar moderately broad,
anterior margin convex; mesoscutum concealed;
legs greatly elongate; abdomen fuscous and
polished; left paramere (Fig. 14A) without
processes or flange; right paramere (Fig. 14B)
truncate subapically with acute apex.

DESCRIPTION. Colouration. Body mostly
stramineous-brown to brown, with banded
antennae, pale legs, and fuscous abdomen. Head
brown to fuscous, sometimes with obscure
circularto linear stramineous markings on medial
aspect of vertex, sometimes extending to inner
margins of eyes. Clypeus, bucculae, mandibular

142

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 14. Vanniopsis howense. A, left paramere; B, right
paramere; C, aedeagus. Scale bar = 0.10mm.

and maxillary plates, genae and gular red to
fuscous-red. Antennae: AI stramineous-red; All
mostly brown, with stramineous annulations
basally, medially and apically, occasionally with
reddish highlighting; AIII brown with apex
stramineous. Pronotum brown to fuscous, most
often with medial longitudinal stramineous
marking; posterior margin occasionally with red
highlighting. Scutellum fuscous, commonly with
medial longitudinal stramineous marking,
contiguous with pronotal medial marking,
posterior quarter to third stramineous. Forewings
stramineous to fuscous, with broad red markings
medially and on medial margins. Legs stram-
ineous; sometimes basal third of coxae fuscous;
mcfafemora with subapical to apical red marking.
Thoracic pleura and abdomen usually fuscous to
red-fuscous. Abdomen fuscous-red to fuscous.

Texture. Dorsum smooth; abdomen polished.

Vestiture. Dorsum with moderate density of
simple stout dark semi-erect setae. Frons without
dense tuft of setae.

Structure. Habitus (Fig. 13). Staphylinoid (Figs
2D, 13). Pronotum (Fig. 2B) subrectangular;
posterior margin rectilinear; pronotal collar
moderately broad, anterior margin convex.
Mesoscutum concealed. Labium reaching apices
of metacoxae. Legs greatly elongate. Male
genitalia: pygophore (Fig. 2E) subconical,
genital opening large, ovoid, terminally oriented;
left paramere (Fig. 14A) small, elongate,
moderately elongate with hook-like apical
process; right paramere (Fig. 14B) short truncate
subapically with acute apical process coplanar
with remainder of shaft; endosoma (Fig. 14C)
with paired looped basal sclerites supporting
membrane, secondary gonopore indis-
tinguishable. Female genitalia: sclerotised rings
(Fig. 16C) moderately large, semi-elliptical,
mesial margin flattened; ventral labiate plate
separated, pointed anteriorly flattened mesially;
dorsal labiate plate folded dorsal to rings;
posterior wall (Fig. 16D) with two bilateral pairs
of thin inter-ramal sclerites.

Measurements. 5 6 6 BL 2.23-2.6 1 , HW 0.55-0.66,
IOD 0.27-0.30, HL 0.38-0.41, PL 0.39-0.43, PW
0.63-0.70, All 1.46-1.72, LL 1.12-1.26; 59 9 BL
2.49-3.08, HW 0.64-0.69, IOD 0.30-0.31, HL
0.40-0.46, PL 0.41-0.47, PW 0.75-0.81, All
1.41-1.56, LL 1.25-1.32.

DISTRIBUTION AND HABITAT. Primarily in
lowland rainforest on Lord Howe Island (Fig.

1 1 ), in a range of vegetation classes (see Pickard,
1983 for floristics). A few specimens were also
encountered in high altitude sites, in closed wet
forests, including the Bubbia/Dracophyllum
vegetation type of Mt Gower. One of us [GC]
collected a large series of V howense in Kentia
palm habitat ( Howea fosteriana ), where it was
found amongst fallen palm litter, in association
with other heteropterans (Atisne sp. [ReduviidaeJ,
Acaraptera sp. [Aradidae] and rhyparochromid
species). Their microhabitat was characterised
by rotting leaf mould and abundant fungal
mycelia.

REMARKS. Vanniopsis howense is distinct
within the genus. Much of its distinctiveness is
attributable to modifications that are commonly
correlated with wing shortening, namely the
compact body, simple pronotum (Fig. 2 A) (short
and subrectangular, narrower collar) and

SYSTEMATICS OF THE VANNIUS COMPLEX

143

scutellum (short and transverse). Like much of
the epigaeic insect fauna of Lord Howe Island,
this species is flightless and there are no known
macropterous specimens.

Unlike V crobylos and V rufescens , this
species has extremely elongate antennae (much
longer than the body) and legs (Fig. 13).
However, the state of the first antennal segment
(compressed, arcuate, elongate and densely
setate) (Fig. 13), is sufficient for its inclusion
within Vanniopsis , as this is a putative synapo-
morphy for the genus. In addition, the simple
male endosoma (lacking lobal sclerites and un-
differentiated secondary gonopore) (Fig. 1 4C), is
shared by the three constituent species of the
genus. It should be noted however, that this
endosomal type is also found in some species of
Vanniusoides.

Vanniopsis rufescens Poppius, 1909
(Figs 3C, 11, 15, 16E,F)

Vanniopsis rufescens Poppius, 1909: 17 (sp. nov.); Bergroth,
1920: 72 (list): Carvalho. 1952: 50 (type species):
Carvalho, 1957: 33 (catalogue); Schuh, 1995: 39
(catalogue); Gorczyca, 1997: 540 (description)

MATERIAL. NEW CALEDONIA: lc3, Mt Koghifs],
22 Q 1FS 166°0rE, 400m, 12-14 November 1986. RL
Brown black light trap (BPBM); 1 8 , Mt Koghi[s], 22° 1 1 ’S
166°0rE, 15 February 1963, NLH Krauss (BPBM); 1 9,
7km S of Koh, 21°32’57 M S 165°50’00"E,31 January 1963,
CM Yoshimoto (BPBM); Id 29 9 1 larva, 20°40’S
1 65°1 3’E, Ponandou intake, 1 00 m, 25 November 200 1 ,
GB Monteith, pyrethrum tree trunks (QM): VANUATU :

1 6 , Erromanga Island, Port Narevin, 1 8°27’S 1 69°03’E
sea level, 25 August 1 979, ferns along streams ( BPBM).

DIAGNOSIS. Macropterous (Fig. 3C); dorsum
weakly rugose; frons sparsely hirsute; pronotum
(Fig. 3C) campanulate, posterior margin
bisinuate; pronotal collar broad, anterior margin
rectilinear; mcsoscutum exposed; peritreme
moderately spout-like; two pairs of looped basal
sclerites: left paramere (Fig. 1 5 A) with basal and
subapical processes; and, right paramere (Fig.
15B) with sub-basal process.

DESCRIPTION. Colouration. Body mostly pale
red, sometimes with stramineous to stramineous-
orange markings (Fig. 3C). Head stramineous-
brown to brown, with posterior margin of vertex
with red markings; frons and medial aspect of
clypeus stramineous; lateral margins of clypeus,
mandibular and maxillary plates, and genae red
to red-orange; gular orange. Antennae: AI red;
All banded, dark brown with medial and apical
stramineous bands, with base minutely
stramineous; AIII mostly brown with apical

stramineous band; AIV brown. Labium mostly
orange-brown; LI most olten with red
highlighting or more uniformly red; apex of LIV
enbrowned. Thoracic pleura mostly red, often
with propleura more brown to orange-brown with
red highlighting. Mesoscutum and scutellum red
to red-brown, sometimes with orange high-
lighting. Hemelytra mostly red, sometimes with
lateral margins of clavus and exocorium more
orange-red, or more variolate (Fig. 3C); medial
angle of endocorium posteriad of claval
commissure sometimes stramineous; cuneus
orange-red, sometimes subapically red, some-
times apex and medial angle stramineous;
membrane fumose; membrane veins red. Legs
mostly stramineous to stramineous-orange; fore-
tibiae sometimes with red highlighting distally;
apical 1/4 of metafemora with red highlighting.
Abdomen mostly red basal ly, more orange
laterally and posteriorly.

Texture. Dorsum weakly rugulose.

144

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 16. Female genitalia. Vanniopsis crobylos. A, sclerotised rings; B,
posterior wall; Vanniopsis howense ; C, sclerotised rings; D, posterior wall;
Vanniopsis rufescens ; E, sclerotised rings; F, posterior wall. Scale bars, A-D
~ 0.25mm ; E-F = 0.20mm.

Vestiture. Head, pronotum and hemelytra with
moderate to sparse distribution of elongate,
simple, dark, erect, setae.

Structure. Habitus (Fig. 3C); Macropterous;
elongate. Antennae subequal in length to body
(Fig. 3C). Labium extending to apices of meta-
coxae. Pronotum (Fig. 3C) campanulate; collar
broad, anterior margin rectilinear; posterior margin
bisinuate. Mesoscutum exposed. Scutellum as
long as wide. External efferent system of meta-
thoracic glands spout-like; peritreme anteriorly
oriented. Hemelytra: strongly depressed at corial
fracture; cuneus narrow and elongate. Male
genitalia: pygophore subconical; genital opening
moderately large, subovoid, dorsally oriented;
left paramere (Fig. 15 A) small, elongate sensory
lobe slightly produced, subapical dorsal margin
with small tubercle, hook-like process apically;
right paramere (Fig. 15B) small, sensory lobe
slight expanded, subapical region slightly
expanded dorsally, pointed apical process
coplanar with remainder of shaft; endosoma (Fig,
15C) with two looped basal sclerites supporting
membranes, also areas with spinose fields;
secondary gonopore indistinguishable. Female
genitalia: sclerotised rings (Fig. 1 6E) moderately
large, widely separated, semi-elliptical, mesial
margin flattened; ventral labiate plate separated,
somewhat truncate anteriorly, flattened mesially,
lateral margin excavated; dorsal labiate plate
asymmetrically scroll-like laterally, right side
larger than left; posterior wall (Fig. 16F) with
bilaterally paired, thin inter-ramal sclerites on
ventral margin.

Measurements. 36 6 BL 3.60-3.75, HW 0.61-0.62,
IOD 0.22-0.23, HL 0.35-0.39, PL 0.48-0.50, PW

0.88-0.94, All 1.26-1.28, LL
1.02-1.08; 3 2$ BL

3.63-3.82, HW 0.60-0.63,
IOD 0.22-0.23, HL 0.32- 0.36,
PL 0.48-0.50, PW 0.90-0.98,
All 1.07-1.20, LL 1.05-1.15.

DISTRIBUTION AND
HABITAT. Vanuatu and
New Caledonia (Fig. 11),
with the majority of
specimens collected from
the latter. This species is
known from the Northern
(Espiritu Santo Island) and
Southern districts (Erro-
manga Island) of Vanuatu,
which spans much of the
latitudinal range of the
archipelago. In contrast, V. crobylos is restricted
to islands of the Central and Northern districts of
the same archipelago. V rufescens is known from
sea level to mid-altitude (400m) localities.

This species has been collected on ferns in a
riparian habitat and from tree trunks. As with V
crobylos , these data suggest that V. rufescens is
not epigaeic.

REMARKS. V, rufescens was described by
Poppius (1909) from a single specimen collected
from Espiritu Santo Island (Vanuatu). According
to Gorczyca (1997), the specimen is badly
damaged, with the abdomen missing. We have
not been able to examine the holotype, however
Gorczyca’s description refers to the lateral and
ventral aspects of the head as red in colour. This is
the condition found in the specimens we have
identified as V rufescens , which distinguishes it
from our new species, V crobylos , which has
these components of the head stramineous in
colour.

These two species are alike but can be
differentiated by the aforementioned head colour
differences and the autapomorphic head vestiture
in V. crobylos. These differences are supported by
significant differences in the parameres (cf. Figs
12A,B and 15A,B) and aedeagus (cf. Figs 12C
and 15C).

The sympatry of these two species in the
Vanuatu archipelago (Espiritu Santo Island) is
noteworthy. V rufescens , aside from the type, is
known from more southern latitudes (Southern
district of Vanuatu, Lifou and New Caledonia),
whereas V crobylos is restricted to more northern
districts of Vanuatu. This extrinsic information

SYSTEMATICS OF THE VANNIUS COMPLEX

145

casts some doubt on the separation of these species,
however their morphologies are conclusive.

Vanniusoides Carvalho &

Lorenzato, 1978

Vanniusoides Carvalho & Lorenzato, 1978: 128 (gen. nov.);
Schuh, 1995: 39 (catalogue); Gorczyca, 1996:337, 340
(note); 1997: 520, 537 (description); Gorczyca &
Konstantinov, 2001: 107-1 10 (description).

TYPE SPECIES. Vannius brevis Poppius, 1909, by
original designation.

DIAGNOSIS. Body smooth; pallid with dis-
ruptive red colouration; frons weakly surpassing
eyes dorsally; pronotal collar moderately broad,
laterally restricted; antennae longer than body;
first antennal segment a little longer than head
length (Fig. 3A,B); first antennal segment
weakly arcuate, not compressed nor densely
setate; pronotum (Fig. 3A,B) campanulate;
posterior margin of pronotum bisinuate;
mesoscutum exposed; peritreme anteriorly
oriented, spout-like; metafemora fiarrow and
elongate; tarsal claws with subapical tooth;
endosoma (Fig. 1 8C) without lobal sclerites, with
pair of arcuate basal sclerites; left paramere (Fig.
18 A) strongly arcuate; female with spinose
posterior wall (Fig. 19B,D).

DESCRIPTION. Colouration. Body (Fig. 3A,B)
mostly stramineous, dorsum with disrupted
red-orange markings. Antennae not banded.
Labium: LI mostly red; L1I1-1V stramineous.
Thoracic pleura mostly red or fuscous-red. Tibiae
and femora either mostly stramineous and
concolorous or stramineous and banded (red).
Hemelytral membrane clear. Abdomen mostly
stramineous with red markings.

Texture. Dorsum smooth.

Vestiture. Dorsum with moderate distribution of
simple, decumbent, elongate setae.

Structure. Macropterous (Fig. 3A,B); elongate.
Head: vertex flat without longitudinal medial
sulcus (Figs 3A,B); eyes enlarged, occupying
much of the lateral aspect of the head apical ly
acute or rounded; frons weakly expressed beyond
eyes; mandibular plates enlarged, sometimes
flattened, anteriorly directed. Antennae: (Fig.
3A,B) significantly longer than body,
thread-like; AI moderately swollen, weakly
arcuate, cylindrical in cross-section (not
compressed), a little longer than head; AII-A1V
elongate, thin. Labium extending to apices of
metacoxae. Prontoum (Fig. 3A,B) transverse,
campanulate; posterior margin bisinuate;
pronotal collar flat, narrow, strongly narrowed

laterally. Mesoscutum visible. Hemelytra (Fig.
3A,B) elongate, extending well beyond abdomen,
depressed beyond cuneal fracture; cuneal fracture
weakly developed; cuneus elongate, narrow.
Proepisternum weakly depressed medially.
Metathoracic spiracle exposed, bounded by
evaporative bodies. External efferent system:
peritreme anteriorly oriented, tumid, spout-like.
Legs elongate; metafemora elongate (Figs
3A,B), narrow, tapered distally. Tarsal claws
moderately arcuate, each with subapical tooth.
Male genitalia: parameres elongate, subequal in
length; left paramere (Fig. 18A) with expanded
sensory lobe; right paramere (Fig. 1 8B) elongate,
sublinear, apex attenuated; phallotheca short;
endosoma (Fig. 18C) either a membranous sac
with a pair of small basal sclerites or strongly
sclerotised basal sclerite with small terminal mem-
branous sac, secondary gonopore undifferentiated.
Female genitalia: sclerotised rings (Fig. 19A,C)
flattened, somewhat ovate, thin; ventral labiate
plate widely separated; dorsal labiate plate
mostly in one plane; posterior wall (Fig. 19B,D)
mostly membranous, with thin bilateral
inter-ramal sclerites; without inter-ramal lobes.

DISTRIBUTION. Northeastern Papua New
Guinea, the Solomon Islands, Fiji Islands and
Australia (north Queensland) (Fig. 17).

REMARKS. Vanniusoides does not have any
characters that uniquely distinguish it, and is at
present defined polythetically. The male genitalia
are somewhat simplified, with the endosoma
(Fig. 18C) without lobal sclerites, and composed
of strap-like basal sclerites, with the secondary
gonopore undifferentiated. In V melafrons , the
endosoma (Fig. 18C) is tube-like (basal sclerites
somewhat fused) and appears to be modified
from the simple condition found in V. clypeatus ,
where the endosoma is a narrow membraneous
sac supported by strap-like basal processes. This
latter character state is also found in Vanniopsis
crobylos , and to some extent in Vanniopsis
howense. The female genitalia are not
sufficiently distinctive to separate Vanniusoides
and Vanniopsis. These genera are best separated
by external characters. In Vanniopsis , the first
antennal segment is compressed and densely
setate, character states that do not occur in
Vanniusoides. Moreover, the vertex of the latter is
not sulcate, in comparison to most other genera of
the Vannius complex (including Vanniopsis),
aside from Austrovannius.

146

MEMOIRS OF THE QUEENSLAND MUSEUM

KEY TO THE SPECIES OF VANNIUSOIDES

1 . Clypeus with distal tubercle brevis

Clypeus without tubercle 2

2. Frons and clypeus fuscous to fiiscous-red; propleura

fuscous to fuscous-red .... melafrons sp.nov.
Head uniformly stramineous, at most with minor red
highlighting; propleura fuscous to fuscous-red or with
reddish longitudinal band . . ♦ * 3

3. Lateral margins ofpronotum with red markings (Fig. 3 A);
clypeus narrow and parallel sided, distally truncate;

propleura stramineous asprokara sp.nov.

Lateral margins of pronotum entirely stramineous;
clypeus distinctly broadened at base, distally rounded;
propleura stramineous with transverse reddish band
clypeatus

Vanniusoides asprokara sp. nov.

(Figs 3 A, 17, 19A,B)

ETYMOLOGY. Greek aspros (= white) and kara (=
head), in reference to the stramineous head of this species.

MATERIAL. FIJI. Holotype, 9, Thawathi, Ovalau,
[17°38’S 178°49’E], 600-800 ft. [183-244m], 16 July
[19]38, EC Zimmerman (BPBM). Paratypes: 9, Draiba
Trail, Ovalau, [17°42’S 178°48’E], 600-800 ft.
[183-244m], 9 July [19]38, EC Zimmerman (AM); 9, nr.

Vuma, Ovalau, [17°40’S 178°49’59"E], 700 ft. [213m], 14
July [19]38, EC Zimmerman (BPBM).

DIAGNOSIS. Head uniformly stramineous;
pronotum and hemelytra with distinct pattern of
red markings (Figs 3A,B); apex of clypeus
truncate, not expanded, without distal tubercle;
metafemora stramineous with subproximal,
submedial and apical red bands; tibiae uniformly
stramineous.

DESCRIPTION. Colouration. Body (Fig. 3A)
mostly stramineous with disruptive red markings
on dorsum. Head: mostly stramineous,
sometimes areas adjacent to antennal insertions
with reddish highlighting. Antennae: AI mostly
red, sometimes stramineous-red; All stram-
ineous, sometimes with obscure red tinge;
AIII-IV stramineous-brown. Pronotum mostly
stramineous, with lateral margins and medio-
posterior region red; collar mostly stramineous,
laterally red, rarely with medial red spot.
Mesoscutum and scutellum mostly stramineous,
with continuous medial red spot. Hemelytra
stramineous with red markings; clavus

SYSTEM ATICS OF THE VANNIUS COMPLEX 147

stramineous, with broad red longitudinal
marking on anterior half medially; endocorium
with two large medial red spots, subapex of
embolium with narrow red marking; cuneus
mostly stramineous with medial red spot. Legs
mostly stramineous; coxae sometimes with red
highlighting; femora stramineous with submedial
and apical red banding; tibiae stramineous,
sometimes with red stippling. Abdomen mostly
stramineous, with lateral margins and subgenital
regions red.

Structure. Habitus (Fig. 3A). Clypeus parallel-
sided, apex truncate, without tubercles. Labium
reaching metacoxae. Female genitalia with
sclerotised rings (Fig. 19A) obscure, flattened,
ovoid, thin, widely separated; ventral labiate
plate widely separated: dorsal labiate plate
undifferentiated; posterior wall (Fig. 19B)
extremely membranous with bilateral inter-ramal
sclerites in middle of membrane .

Males unknown.

Measurements. 39$ BL 3.84-4.20, HW 0.6 1-0.65,
IOD 0.24-0.25. HL 0.28-0.35, PL 0.34-0.37, PW
1.01-1.06, All 1.90-2.08, LL 1.08-1.20.

DISTRIBUTION AND HABITAT. On Ovalau in
the Fijian Islands (Fig. 1 7). The habitat is unknown.

REMARKS. Vdnniusoides asprokara is dis-
tinguished from other species of Vanniusoides by
the pattern of red markings on the dorsum (Fig.
3A); in particular the red lateral margins of the
pronotum are unique to this species. In addition,
this species has the propleura uniformly stram-
ineous and the metafemora have red bands. The
female genitalia are also distinct (cf. Fig. 19D).

Vanniusoides nielafrons sp. nov.

(Figs 3B, 17, 18, 19C,D)

ETYMOLOGY. Greek melas (= black) and Latin frons (=
forehead) referring to the dark front of the head.

MATERIAL. QUEENSLAND: Holotype, 6\ Cape
Tribulation, \6°0iyS 145°26.3’E, 28 March 2000, RL
Kitching canopy fog (QM T 1086 15). Paratypes 9,
Kuranda, [16°49’S 145 D 38’E]. 13 March 1956, JLGressitt
(BPBM ); 5, Cooper Creek, 10 ml [16km] N of Daintrce
Riv[er], [16°08’S 145°27'E], 2 May 1970, GB Monteith
(UQIC).

DIAGNOSIS. Frons and clypeus fuscous to
fuscous-red; clypeus distally truncate, without
tubercle; pronotum and hemelytra with distinct
red patterning; metafemora stramineous with
subproximal, submedial and apical red bands;
tibiae with two submedial red bands; parameres

(Figs 18A,B) elongate; endosoma (Fig. 18C)
strongly sclerotised with serrate apical flange.

DESCRIPTION. Colouration. Body mostly
stramineous with disruptive red markings on
dorsum (Fig. 3B). Head: vertex stramineous,
remainder of head red to fuscous-red (Fig. 3B).
Antennae: AI red; All stramineous, sometimes
with red highlighting; AIII-AIV stramineous to
stramineous-brown. Pronotum mostly
stramineous with pair of submedial caudal red
spot (Fig. 3B). Thoracic pleura fuscous to
fuscous red, sometimes metepisternum more
red-brown. Mesoscutum and scutellum mostly
stramineous with pair of sublateral red spots.
Hemelytra mostly stramineous, with distinctive
red patterning (Fig. 3B); clavus with inner
margins and apices red; endocorium with broad
red band adjacent to clavus, red spots on
endocorial angle, subapex of embolium and
posteriad of R+M. Legs mostly stramineous;
coxae either mostly brown with apices

148

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 19. Female genitalia. Vanniusoides asprokara. A, sclerotised
rings; B, posterior wall; Vanniusoides melafrons ; C, sclerotised
rings; D, posterior wall. Scale bar = 0.25mm.

REMARKS. The fuscous front of
the head differentiates this species
from V asprokara and V clypeatus ,
both of which have pale heads.
Moreover, the hemelytra have more
red markings (cf. Fig. 3A,B). The
male genitalia of V melafrons and V
clypeatus are the only species
investigated to date, and their
morphologies differ, with the former
species having a tube-like endosoma
(Fig. 18C).

Vanniusoides clypeatus

Gorczyca & Konstantinov, 200 1
(Fig. 17)

stramineous or more uniformly pale; metafemora
stramineous with subproximal, submedial and
apical red bands; tibiae with two submedial red
bands. Abdomen mostly stramineous with lateral
margins and subgenital regions red.

Structure. Habitus (Fig. 3B). Clypeus parallel-
sided, apex truncate, without tubercles. Labium
extending just beyond apices of metacoxae. Male
genitalia: left paramere (Fig. 18A) elongate with
expanded sensory lobe and acute apex; right
paramere (Fig. 18B) elongate, abruptly con-
stricted subapically, apex pointed; endosoma
(Fig. 18C) tube-like, formed by pair of strongly
sclerotised basal sclerites, one strap with terminal
serrate flange, membranous sac terminal. Female
genitalia: sclerotised rings (Fig. 19C) flattened,
obscure, circular, thin, widely separated; ventral
labiate plate widely separated
with strong spinules; dorsal
labiate plate basically flatten-
ed, spinose, simple, adhering
to floor of genital chamber;
posterior wall (Fig. 1 9D) thin,
membranous with strong
spinules mesially and bilateral
inter-ramal sclerites dorsally .

Measurements. 1 d (Holotype)

BL 3.2. HW 0.61, IOD 0.17,

HL 0.37, PL 0.40, PW 1.01,

All 1.74, LL 1.25; 22 9 BL
4.10-4.30, HW 0.60-0.62,

IOD 0.17-0.19, HL 0.34-0.37,

PL 0.33-0.34. PW 1.08, All
1.76-1.80, LL 1.34-1.40.

Vanniusoides clypeatus Gorczyca & Konstantinov 2001: 108
(sp. nov.).

MATERIAL. SOLOMON ISLANDS: Id, New Geoigia
Island, Munda, 0-1 00m, XI- 1980, N.L.H. Krauss coll.
(BPBM).

REMARKS. This species is distinctive because
of the highly autapomorphic condition of the
clypeus and mandibular plates, which are both
rounded apically. The male genitalia are identical
to those illustrated by Gorczyca & Konstantinov
(2001). These authors have provided a detailed
description of the species and we have opted
against providing a redescription.

Peritroptsca bituberculata
3 4

FIG. 20. Cladogram of genera of Vannius complex. Synapomorphies - black
DISTRIBUTION AND boxes; homoplasies = grey boxes. Superscript numerals = character
HABITAT. Rainforests of the number; subscript numerals = character state. Numerals in circles = node
wet tropics of northeast number. Length = 30 steps. Consistency index = 0.70. Retention index =
Queensland (Fig. 17). 0.70.

SYSTEMATICS OF THE VANNIUS COMPLEX

149

TABLE 1. Characters and character states of the
Vannius complex and outgroups.

1. Head orientation: horizontal (0); dorsoventral (1).

2. Frons and clypeus: bilobed (0); coplanar (1).

3. Vertex: without medial sulcus (0); with shallow sulcus (1);
with deep sulcus (2).

4. Antennal segment I: shorter or subequal to head length (0);
significantly longer than head length (1).

5. Antennal segment II: concolorous (0); banded (1).

6. Pronotal collar: absent (0); rounded and thin (1); flat and
broad (2).

7. Dorsum commonly with disrupted red colouration: no (0);
yes (1).

8. Metafemora: moderately elongate (0); greatly elongate ( 1 ).

9. Pretarsal claws: without subapical tooth (0); with subapical
tooth (1).

10. Parempodia: setiform (0); spatulate (1).

11. Secondary gonopore: indistinguishable (0);
well-sclerotised, with spinose aperture (1).

12. Endosoma: without basal sclerites (0); with basal sclcrites

( 1 ).

13. Endosoma: without lobal sclerites (0); with lobal sclerites

( 1 ).

14. Basal sclerite: free from base of endosoma, not fused (0);
extending to apex of endosoma, fused and somewhat bent
( 1 ).

15. Sclerotised rings: thin, not strongly bent, without
projections from lateral and mesial margins (0); thick,
somewhat strongly bent, with projections from lateral and
mesial margins (1).

16. Ventral labiate plate: separated medially (0); joined on
anterior margin (1).

17. Dorsal labiate plate: not expanded lateral of sclerotised
rings (0); expanded lateral of sclerotised rings, lateral most
margins scroll-like (l).

18. Intcr-ramal sclerite: divided, structure formed of two thin
sclerites (0); completely spanning base of first valvifer ( 1 ).

19. Inter-ramal lobes: absent (0); present (1).

PHYLOGENETICS

These results are based on an analysis of the six
genera of the Vannius complex and three
outgroups within the Cylapinae, including
exemplars of the Bothriomirini ( Bothriomiris
lugubris ), Fulviini ( Peritropisca bitubercuhta)
and Cylapini (Cylapocerus). The characters and
character states are given in Table 1 and the data
matrix in Table 2. A single cladogram of
minimum length (30 steps) was obtained with a
consistency index of 0.70 and a retention index of
0.67. The discussion of sister groupings is based
on the synapomorphies shown in Fig. 20.

Node 1 - The Cylapini are defined in this
analysis by the dorsoventral head (1-1),
secondary gonopore sclerotised with a dentate
aperture (11-1), and the endosoma with basal

sclerites (12-1). Neither of these characters are
exclusive to the Miridae, but are not found in the
exemplars of the other recognised suprageneric
groups of Cylapinae (Bothriomirini and Fulviini)
that were examined. Some representatives of the
Fulviini (e.g. Kelton, 1959) are known to possess
basal sclerites, and this is not considered to be a
reliable synapomorphy for the Cylapini, although
the basal sclerites of the latter tribe are often
strap-like. Further investigation is required to
determine if the ‘basal sclerites’ character can be
differentiated to identify additional synapo-
morphies. The secondary gonopore varies within
the ingroup and this character is homoplasic in
this analysis.

Node 2 - The Vannius complex is defined by
the following synapomorphies: gular and the
frons and the clypeus in the same plane (2-1);
pronotal collar broad and flat (6-1); dorsum with
disrupted colouration (7-1); metafemora greatly
elongate (8- 1 ); and, parempodia spatulate (10-1).
The derived states of characters 7 and 8 are also
found in other members of the Cylapinae, and
require further investigation beyond the out-
groups that were examined. The coplanar frons +
clypeus is also deserving of more investigation,
particularly for taxa such as Cylapus Say and its
relatives, but is unlikely to be upheld as a
synapomorphy for the Vannius complex. The
spatulate parempodia remain the most conclusive
synapomorphy for the Vannius complex, and is
unknown for all other examined cylapines. As
noted above, the condition also occurs in
Palaucoris , a few taxa of Termatophylini
(Deraeocorinae) and Phylinae, but these are
undoubtedly independent derivations. The broad
and flat pronotal collar is also considered to be a
credible synapomorphy, as the pronotal collar in
the Cylapini is at most narrow and more rounded.

Node 3 - In this analysis, Austrovannius is
sister-taxon to the remainder of the lannius
complex. The latter subset is defined by the
banded antennae (5-1) and tarsal claws with
subapical teeth (9-1). Both of these characters
exhibit homoplasy, with Vannius lacking
subapical teeth (5-0) and Vanniusoides
possessing concolorous antennae (9-0).

Node 4 - Paracylapus is the next most basal
taxon of the Vannius complex. T he remaining
genera are united by a single homoplasic
character; secondary gonopore indistinguishable
( 11 - 0 ).

Node 5 -Afrovannius is sister-taxon to the rest
of the Vannius complex. Vannius + (Vanniopsis +

150

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Character matrix of three outgroups ( Bothriomiris lugubris , Peritropisca bituberculata and
Cyalpocerus sp.) and six genera of the Vannius complex against 19 characters. Missing data = ?

Character

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

Bothriomiris lugubris

0

0

0

0

0

0

0

0

0

?

0

1

7

?

?

?

1

1

0

Peritropisca bituberculata

0

0

1

0

0

0

0

0

0

0

0

1

7

?

?

7

0

0

1

Cylapocerus

1

0

2

1

0

0

0

?

0

1

1

0

0

?

?

?

7

?

1

Afrovannius

1

1

1

0

l

1

1

1

1

0

1

0

0

?

7

?

7

7

2

Austrovannius

1

1

0

0

0

1

1

0

1

1

1

1

0

1

1

0

1

1

2

-

Paracylapus

1

1

1

0

1

1

1

1

1

1

1

0

0

7

7

?

7

7

2

Vanniopsis

1

1

1

1

1

1

1

1

1

0

1

0

?

0

0

1

0

0

2

Vannius

1

1

1

0

1

1

1

0

1

0

1

0

1

?

?

7

?

7

2

Vanniusoides

1

1

0

1

0

1

1

1

1

0

1

0

1

0

0

1

0

0

2

Vanniusoides) are a clade on the basis of one
synapomorphy; basal sclerites extending to apex
of endosoma (14-1). This represents the most
supported in-group clade within the Vannius
complex. These three Eastern Hemisphere
genera are also saliently alike, and it is predicted
that additional synapomorphies will be found for
this clade.

Node 6 - Vanniopsis and Vanniusoides are
united on the basis of a single homoplasic
character; first antennal segment significantly
longer than the head (4- 1 ). In addition, these taxa
also exhibit similar male genitalia, with the
endosoma having basal sclerites and lacking
lobal sclerites.

ACKNOWLEDGEMENTS

The following curators are thanked for their
provision of specimens: Gordon Nishida
(BPBM); Geoff Monteith (QM), Jan Forrest and
Gordon Gross (SAMA); and, Margaret
Schneider and Greg Daniels (UQ1C). Roger
Kitching (Griffith University) also provided
material from north Queensland. Geoff Monteith
is particularly thanked for providing much of the
material and taking a special interest in collecting
cylapines in wet rainforests of north Queensland
and New Caledonia. His material has allowed for
a broader understanding of Cylapinae
systematics. Ian Hutton assisted with the
collection of Vanniopsis howense from Lord
Howe Island. The Lord Howe Island Board is
thanked for allowing collecting on the island and
access to their research facility. Rossana Silveira
assisted with many aspects of the project,
including the illustrations of the female genitalia.
Gareth Carter assisted with the distribution maps.
Matthew Bulbert took the habitus figures of
Figure 3. Hannah Finlay inked the sketches of the

male genitalia. Heloise Gibb provided the habitus
illustrations. Sue Lindsay was responsible for the
scanning electron microscopy. Chris Reid is
particularly thanked for his support of our work,
including collections of new material and reading
of the manuscript. Dan Bickel and Winston
Ponder also provided advice on the biogeography
of the Australian region. The Australian Museum
Trust is thanked for its support of biodiversity
and taxonomic research. This work is partly
funded by an allocation from the New South
Wales State Biodiversity Strategy and the
Australian Biological Resources Study.

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107-110.

KELTON, L. 1959. Male genitalia as taxonomic
characters in the Miridae (Hemiptera). Canadian
Entomologist (Supplement) 11: 1-72.

KERZHNER. I.M. & KONSTANTINOV, F.V. 1999.
Structure of the aedeagus in Miridae (Heteroptera)
and its bearing to suprageneric classification. Acta
Societas Zoologicae Bohemicae 63: 117-137.

NIXON, K.C. 1999. WinClada ver. 1.0000 (Published
by the author: Ithaca, NY, USA).

POPPIUS, B. 1909. Zur Kenntnis dcr Miriden-
Unterfamilie Cylapina Reut. Acta Societatis
Scientiarum Fennicae 37(4): 1-46.

SCHUH, R.T. 1975. The structure, distribution, and
taxonomic importance of trichobothria in the
Miridae (Hemiptera). American Museum
Novitates 2585: 1-26.

1976. Pretarsal streuture in the Miridae (Hemiptera)
with a cladistic analysis of relationships within
the family. American Museum Novitates 2601:
1-39,

1984. Revision of the Phylinae (Hemiptera,
Miridae) of the Indo-Pacific. Bulletin of the
American Museum of Natural History 177(1):
1-476.

1 995. Plant bugs of the world (Insecta: Heteroptera:
Miridae): systematic catalog, distributions, host
list, and bibliography. New York: The New York
Entomological Society, i-xii, 1-1329.

SCHUH, R.T. & SCHWARTZ, M.D. 1984. Carvalhoma
(Hemiptera: Miridae) new subfamily placement.
Journal of the New York Entomological Society
92: 48-52.

SCHWARTZ, M.D. & FOOTTIT, R.G 1998. Revision
of the Nearctic species of the genus Lygus Hahn,
with a review of the Palaearctic species
(Heteroptera: Miridae). Memoirs on Entomology,
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(Hemiptera). Iowa State College Journal of
Science 25: 1-81.

STONEDAHL, GM. 1988. Revision of the mirine
genus Phytocoris Fallen (Heteroptera: Miridae)
for western North America. Bulletin of the
American Museum of Natural History 198: 1-88.

SWEARINGEN, M., HEADRICK, D. & BELLOWS,
T. 1997. Comparison of fixation and drying
procedures for scanning electron microscopy
among insect body types. Proceedings of the
Entomological Society of Washington 99(3):
513-522.

WHEELER, A.G 2001. Biology of the plant bugs
(Hemiptera: Miridae). Pests, predators,
opportunists. (Cornell University Press: Ithaca).

NEW RECORDS OF PALICID CRABS (CRUSTACEA: BRACHYURA: PALICIDAE)

FROM AUSTRALIA

P. CASTRO AND P.J.F. DAVIE

Castro, P. & Davie, P. 2003 06 30: New records of palicid crabs (Crustacea: Brachyura,
Palicidae) from Australia. Memoirs of the Queensland Museum 49(1): 153-157. Brisbane.
ISSN 0079-8835.

Eight species of palicid crabs have been identified from Australia. Parapalicus ambonensis,
Paliculus kyusyuensis, Exopalicus maculatus and a species close or identical to Miropalicus
vietnamensis are recorded for the first time in Australia. There are also first records ol
Neopalicus jukesii from Western Australia and Crossotonotus spinipes from the Northern
Territory and the Cocos (Keeling) Islands. Thirteen species of Palicidae are now known from
Australian waters. □ Palicid crabs, new species, Australia.

P. Castro. Biological Sciences Department, California State Polytechnic University,
Pomona, CA 91768, USA; P.J.F. Davie, Queensland Museum, POBox 3300, South Brisbane
4101, Australia; 5 November 2002.

Little is known about the biology of palicid
crabs. Most species are inhabitants of soft
sediments in relatively deep water, while some
are found in shallow water associated with coarse
sediments near coral reefs or with rocky bottoms.
All seem to ingest sediment particles. The
taxonomy and biogeography of the 43 known
species of Indo-west Pacific palicids has been
reviewed by Castro (2000).

Castro (2000) recorded 9 palicid species from
Australian waters. Material from several Australian
museums adds four additional species, increas-
ing the number of Australian species to 13. In
addition to the 8 species listed here, 5 were
previously recorded from Australia by Castro
(2000): Pseudopalicus investigatoris (Alcock,
1 900), P. macromeles Castro, 2000, P. ohauensis
(Rathbun, 1906), P. serripes (Alcock &
Anderson, 1895) and Pleurophricus cristatipes
A. Milne Edwards, 1873.

Abbreviations used: AM. Australian Museum,
Sydney; MMUS, MacLeay Museum, University
of Sydney; QM, Queensland Museum, Brisbane;
WAM, Western Australian Museum, Perth; CL,
carapace length; CW carapace width.

Subfamily PALICINAE Bouvier, 1898

Parapalicus Moosa & Serene, 1981
Parapalicus ambonensis Moosa & Serene, 1981
(Fig. 1A)

Parapalicus ambonensis Moosa & Serene. 1981: 29, figs 2a,
3a, pi. 1, fig. D; Castro, 2000: 489, figs 16, 19a, 58.

MATERIAL. QM W 16970, 19, off Mission Beach,
NEQLD, Australia, 17°53’ S, 146°5r E, 140-142m,
20.01.1986, CSIRO, R.V. ‘Soela’.

REMARKS. This is the first record of the species
in Australia. It has been collected from mostly
muddy bottoms of the Andaman Sea coast of
Thailand, Banda and Coral Seas and off the island
of Futuna, southwestern Pacific Ocean (Castro,
2000). Bathymetric range: 80-440m.

Miropalicus Castro, 2000
Miropalicus cf. vietnamensis (Zarenkov, 1968)
(Fig. IB)

Palicus vietnamensis Zarenkov, 1968: 762, fig. 2A-F.
Miropalicus vietnamensis ; Castro, 2000: 522-525, tigs 29,
30a, 59, 60f (full synonymy and references).

MATERIAL. QMW25170. Incomplete specimen. North
West Shelf, Western Australia, 19°59.1’S 117°49.0’E,
beam trawl, 43m, CSIRO, RV Soela , stn 3D1BT, 25 June
1983.

REMARKS. The Australian specimen consists
of only the anterior border of the carapace of a
very small, most probably juvenile, specimen
(distance between outer anterior lobes of 0.^
mm). The two orbits, the right supraorbital and
suborbital borders and the right basal antennal
segment were present. The pterygostomial lobe
was missing on both sides. Characteristic of
Miropalicus are the wide orbits, large eyes with
dorsoventrally flattened cornea, very short
supraorbital lobes and relatively short suborbital
lobes, long and conspicuous postorbital angles
with pointed tip directed outward, and a
rectangular, slender basal antennal segment that
lacks a distal expansion. Unlike the only species
so far included in the genus, M. vietnamensis , the
shallow supraorbital lobes are very wide and
have a straight edge, not narrow with rounded tips
(Castro, 2000: fig. 22a). The suborbital border

154

MEMOIRS OF THE QUEENSLAND MUSEUM

consists of a wide, oblique
inner lobe unlike the
slightly triangular lobe of
M. vietnamensis (Castro,
2000: fig. 22b) and a
narrow, slightly pointed
outer lobe that is similar to
that of M. vietnamensis. It
is possible, however, that
the simpler arrangement of
both supraorbital and
suborbital lobes is only a
characteristic of juveniles.
There are three small
frontal lobes instead of the
two in M. vietnamensis , a
rare situation observed in
other palicids and that
perhaps results from
damage and subsequent
regeneration.

The Australian specimen
was collected from 43m
depth, whereas M.
vietnamensis is known
from 239-647m from the
South China Sea to the
Loyalty Islands, southern
Coral Sea (Castro, 2000).

Paliculus Castro, 2000

Paliculus kyusyuensis

(Yokoya, 1933)

(Fig. 1C)

Palicus kyusyuensis Yokoya,
1933: 206,217, fig. 70.
Paliculus kvusyuensis; Castro,
2000: 527-530, figs 30b, 31,
56 (full synonymy and
references).

FIG 1 . A, Parapalicus ambonensis Moosa & Serene, 1981, 9 (QM W 1 6970); B,
Miropalicus cf. vietnamensis (Zarenkov, 1968), incomplete specimen
(QMW25 1 70); C, Paliculus kyusyuensis (Yokoya, 1 933), 9 (QM W 1 54 1 5); D,
Exopalicus maculalus (Edmondson, 1930), 6 (QMW25171).

MATERIAL. QMW15417,

39,1 juv, 9 , off Great Barrier
Reef, Queensland, 17°21.77’S
146°48.52’E, epibenthic sledge,

296-302m, Cidaris I, stn 42-2,

FRV Franklin , 15.05.1986;

QMW15418. 89, juv. 9 (same
data its W 1 54 1 7); QMW 15415,

9, off Great Barrier Reef,

17°34.58’S 146°53.2rE, epibenthic sledge, 458-500m,
Cidaris I, stn 43-2, FRV Franklin , 15.05.1986;
QMW15416, 9, off Great Barrier Reef, 17°55.38’S
147°00.96 , E, beam trawl, 295-309m, Cidaris I, stn 46-3,
FRV Franklin, 16.05.1986.

REMARKS. The conspicuous, salient tubercles
along the posterior border of the carapace show

noticeable differences in size and shape, varying
from short and broad to narrow and more acute.
Dorsally the carapace has short, rounded tubercles,
as in specimens studied by Castro (2000). These
tubercles, however, were not shown in the
illustration (Castro, 2000: fig. 23a).

The species is known from sandy bottoms at
depths of 30-7 10m in relatively few locations in

NEW PALICID CRABS FROM AUSTRALIA

155

the Indian Ocean (Madagascar) and from Japan
to Wallis Island in the southwestern Pacific (Castro,
2000). This is the first record from Australia.

Exopalicus Castro, 2000
Exopalicus maculatus (Edmondson, 1930)
(Figs ID, 2)

Palicus maculatus Edmondson, 1930: 15, figs 6a-g, pi. 1, fig. C.
Exopalicus maculatus ; Castro, 2000: 546-548, figs 37d, 38,
56, 61b (full synonymy and references).

MATERIAL. QMW25171, d. Lady Elliott I., Bunker
Group, Queensland, 24°07’S 152 0 43’E, coral reef near
lighthouse, night dive, P. Davie & D. Potter, 14 Aug. 1 985.

REMARKS. The presence of Exopalicus
maculatus in eastern Australia confirms its wide
distribution, having been known only from the
Hawaiian and Marshall islands in the Pacific and
Reunion in the western Indian Ocean (Castro,
2000). It has been collected in Guam (Castro,
unpub 1. data). As in most of the material from
Hawaii and Reunion, the Queensland specimen
was collected from a hard substrate in shallow
water at night.

The live Australian specimen showed two
large pairs of orange spots on a cream to light-
yellow carapace (Fig. 2). There were irregular,
red-brown markings along the anterior and
posterior borders of the carapace. The legs were
banded red-brown.

Neopalicus Moosa & Serene, 1981
Neopalicus jukesii (White, 1847)

Cymopolia jukesii White, 1 847: 338, pi. 2, fig. 1 .

Neopalicus jukesii: Castro, 2000: 554-558 , figs 39b, 40b-c,
41c, 49, 61c (full synonymy and references).

MATERIAL. North West Shelf, Western Australia, CSIRO,
RV Soda (64 stations between 19°03.0\S, 1I9°02.4 , E and
20°01.2'S, U6°57.6 , E, 36-82m depth, epibenthic sledge or
beam trawl, from 8.12.1982 to 30.10.1983): QMW25I23, 2
juv. 9 ; QMW25 134, d; QMW25 1 1 0, 2 9 ; QMW25 126, <5;
QMW25144, 2d; QMW25138, juv. 9: QMW25139, 2d;
QMW25116, d; QMW25146, d; QMW25I15, 9;
QMW25I24, juv. 9; QMW25133, d; QMW25127, 9;
QM W25 1 56, 2 9 ; QMW25 1 60, unsexedjuv.; QMW25I25,
9; QMW25I54, 29; QMW25132, d; QMW25166, 9;
QM W25 1 29, juv. 9; QMW25155. 2 juv. 9;QMW25162,
d; QMW25111, d 9; QMW25147, d incomplete
specimen: QMW25 1 14, 3 9 ; QMW25 1 1 7, 7d 3 9 juv. 9 ;
QMW25141, d 2 unsexed juv.: QMW25135, juv. 9;
QMW25140, 9;QMW2513l,juv. 9;QMW25106, d 29;
QMW25158, d 29; QMW25145, d; QMW25163, 6 3
juv.; QMW25120. juv. 9; QMW25130, juv. 9;
QM W25 1 57, 3 9 juv. 9 2 unsexedjuv.; QMW25 1 59, d 2 9 ;
QMW25148, d 5 unsexedjuv., 2 incomplete specimens;
QMW25165,juv. d 29; QMW25152, damaged specimen;
QM W25 1 67, 2 9 ; QMW25 1 28, 3 9 ; QMW25 1 08, 2 d 39 5
juv. 9; QMW25164, d 2 juv.; QMW25161, d 2 juv. 9 15

FIG 2. Exopalicus maculatus (Edmondson. 1930), d
(QM W25 171) showing pattern of live colouration.

unsexed juv.; QMW25118, 9; QMW25168, 2d 29;
QMW25153, 2d juv. 9; QMW25169, 59; QMW25109,
9;QMW25112, djuv. 9:QMW25119, lOd 269 3juv. 9;
QMW25151,4d 9 juv. 9 32 unsexedjuv.; QMW25 149, d
3 juv. 9 6 unsexedjuv., 2 damaged specimens; QMW25 1 07,
4d 39 juv. 9; QMW25137, 2 juv. 9 3 unsexedjuv., 2
incomplete specimens; QMW25143, 2d juv. 9;
QMW25122, 9: QMW25121, I6d 59 4 juv. 9;
QMW25136, 7d 9 4 juv. 9 3 incomplete specimens;
QMW25142, 3 unsexedjuv.; QMW25113, d 9 juv. 9;
QM25150, 9 juv. d juv. 9 damaged specimen.
WAM-C27188, 9 Dampier Archipelago, 1.4 miles ESE of
Tish Point, Rosemary I., 20°30.48 , S 1 16°36.53’E, rake-box
dredge, 9- 10m, sandy mud 26.07.1999; QMW 17431, 29,
Gulf of Carpentaria, 15°57.6'S 1 38°4L8T, beam trawl, 25m,
FRY Southern Surveyor ; stn 94, 1 1 . 1 2. 1 99 1 ; AMP 1 0524, d ,
Hayman I., Whitsunday Group, Queensland, 20°03’S
148°53’E, 9m, F.A. McNeill Jan. 1934; AMPI9525, d 29,
Whitsunday Group, Black L, near Langford, Nov. 1969.

REMARKS. The large number of juveniles
collected (as small as CL 0. 1 8mm, CW 0.27mm)
permits their description for the first time.
Anterior lobes are more salient than in the larger
individuals. Anterolateral teeth are similarly
more pointed and salient, the most anterior ones
being the largest; the third and most posterior
teeth are smallest and are absent in the smallest
individuals. The carapace is nearly smooth.

N. jukesii ranges across the Indian Ocean to the
western Pacific (southern Japan to the C oral Sea)
in mostly coarse sand at depths of 10- 1 46m.

Palicoides Moosa & Serene, 1981
Palicoides whitei (Miers, 1 884)

vmopolia whitei Miers, 1 884: 55 1 , pi. 49, figs C, c.
Palicoides whitei ; Castro, 2000: 565-568. figs 42b. 43b-c, 50,
61 e (synonymy and references).

MATERIAL. MMUS-C2139, d. Damley I., Queensland,
9°35’S 143°46’E, W.J. MacLeay, HMS Chevert,
7-8 08.1875; QMW9896, juv. 9, 15km NW Lizard I.,
Queensland, 14°35.3’S \45°23% 27m, mud and shell
substrate. Commonwealth Northern Prawn Survey, stn
IB/ 14, FV Markwell Explorer , Sept. 1979.

156

MEMOIRS OF THE QUEENSLAND MUSEUM

REMARKS. The species is known across the
Indian Ocean to the western Pacific (southern
Japan to the Coral Sea) (Castro, 2000). It inhabits
coarse sand at depths of 7-70m.

Subfamily CROSSOTONOTINAE Moosa &
Serene, 1981

Crossotonotus A. Milne Edwards, 1873
Crossotonotus compressipes A. Milne
Edwards, 1873

Crossotonotus compressipes A. Milne Edwards, 1873: 259;
Castro, 2000: 571-574, figs 44, 51 (full synonymy and
references).

MATERIAL. MMUS-C2138, <3, Damley 1., Queensland.
9°35 , S 143°46’E, W.J. MacLeay, H.M.S. Chevert ,
7-8.08.1875; MMUS-C2137, 9 (data as for
MMUS-C2138); QMW12488, juv. 9 , Dugong I., Torres
Strait, 10°3rS 143°04’E, reef flat, low tide, Queensland
Fisheries Service, 17.07.1974.

REMARKS. The dorsal surface of the female
(QMW 12488: CL 8.6mm, CW 9.4mm), although
preserved for almost 26 years showed a dark-
brown, irregular pattern on a light background.

C. compressipes , although rarely collected, is
known from southern Japan to Samoa (Castro,
2000). It appears to be restricted to hard bottoms
in shallow water.

Crossotonotus spinipes (De Man, 1888)

Pleurophricus spinipes De Man, 1888: 344, pi. 15, figs 1,
la-c.

Crossotonotus spinipes ; Castro, 2000: 574-578, figs 45, 46,
51, 61 f (full synonymy and references).

Crossotonotus brevimanus ; Morgan. 1992: 47.

Manella brevimana ; Springthorpc & Lowry, 1994: 94.

MATERIAL. WAM-C 19675, <5, juv. 6 , SW end of
Horsburgh I., Cocos (Keeling) Islands, to 30m, GJ.
Morgan, 16.02.1989; WAM-C 197573, juv. 9 , Cocos
(Keeling) Islands, C.W. Biyce & F.E. Wells, 17.02.1989;
WAM-C20595, 6 juv. 9 , SW of Descartes I., Kimberley
region. Western Australia, coral reef, GJ. Morgan, KIRE
1991, stn 22, 19.08.1991; QMW24154, juv. 9 , Gove
Peninsula, Northern Territory, 12°10’S 136°5(TE, N.
Coleman, Feb. 1993; AM-P 19434, juv. 9 , Mandora Point,
Darwin, Northern Territory, 12°27’S 1 30°50’E, under rock
at low tide, N. Coleman, 23. 1 1 . 1 972; QM W2245 1 , juv. 9 ,
NE end of East Herald Cay, Coringa-Herald Nature
Reserve, Queensland, 16°56'S 149°irE, lagoon, 24m,
Royal Geographic Society of Queensland Herald Cay
Expedition, P. Davie & M. Preker, 27.06.1997;
QMW25I72, juv. < 3 , Masthead I., Queensland, 23°32’S
151°44’E, pontoon, night dive, 9m, P. Davie & D. Potter,
11.02.1986; QMW 1505, d, Mud I., Moreton Bay,
Queensland, 23°32’S 150°5TE, V.F. Collin, 16.07.1942;
QMW21S95. juv. 9 , Lamont Reef. Capricorn Group,
23°36’S 152°03’E, 10m, N. Coleman, Sept. 1992.

REMARKS. The carapace of a very large male
(QM1505: CL 27.6mm, CW 33.8mm) was
atypical for the species. The anterior lobes were
not bent upward and the tips of the anterolateral
teeth were less pointed than is usual.

The species is widely distributed in the Indo-
west Pacific except the southeastern Pacific
(Castro, 2000). It is restricted to hard bottoms and
known from the intertidal to 146m.

ACKNOWLEDGEMENTS

We are most grateful to P. Berents and S. Ayong
(AM), J. Short (QM), D. Jones and M. Hewitt
(WAM) and S. Norrington (MMUS) for their
assistance in providing us with the material.

LITERATURE CITED

ALCOCK, A. & ANDERSON, A.R.S. 1895. Natural
history notes from H.M. Indian Marine Survey
Steamer 'Investigator,' Commander C.F. Oldham,
R.N., commanding. Series II., No 17. List of the
shore and shallow-water Brachyura collected
during the season 1893-1894. Journal of the
Asiatic Society of Bengal 63: 197-209.
ALCOCK, A. 1900. Materials for a Carcinological
Fauna of India. No. 6. The Brachyura Catametopa,
or Grapsoidea. Journal of the Asiatic Society of
Bengal 69: 279-456

BOUVIER, E.L. 1 898. Observations on the crabs of the
family Dorippidae. Annals and Magazine of
Natural History' (7)1 : 103-105
CASTRO, P. 2000. Crustacea Decapoda: A revision of
the Indo-west Pacific species of palicid crabs
(Brachyura Palicidae Bouvier, 1898). Pp.
437-610, figs 1-61. In Crosnier, A. (ed.) Resultats
des campagnes MUSORSTOM, Vol. 21.
Memoire Museum nationale dTIistoire naturelle
Paris 184: 1-813.

DE MAN, J.G 1 888. Bericht liber die von Herm Dr. J.
Brock im indischen Archipel gesammelten
Decapoden und Stomatopoden. Arch.
Naturgesch. 53( 1 ): 2 1 5-600.

EDMONDSON, C.H. 1930. New' Hawaiian Crustacea.
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MIERS, E.J. 1884. Crustacea. Report on the Zoological
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during the Voyage of H.M.S. k Alerf 1881-2. Part
I. The collections from Melanesia. Part II.
Collections from the western Indian Ocean. Pp.
178-322, 513-575. (British Museum: London).
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Musee de M.C. Godeffroy. Journal of the
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MOOSA, M.K. & SERENE, R., 1981. Observations on
the Indo-West-Pacific Palicidae (Crustacea:
Decapoda) with descriptions of two new
subfamilies, four new genera and six new species.
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157

MORGAN, G.J. 1992. Decapod crustaceans. In
Morgan, GJ. (ed.) Survey of the Aquatic Fauna of
the Kimberley Islands and Reefs, Western
Australia. Report of the WAM Kimberley Island
and Reef Expedition, Aug. 1991. (Western
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RATHBUN, M.J. 1906. The Brachyura and Macrura of
the Hawaiian Islands. Bulletin of the U.S.
Fisheries Commission 23(3): 827-930.

SPRINGTHORPE, R.T. & LOWRY, J.K. 1994.
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Reports of the Australian Museum 11: 1-134.

WHITE, A. 1 847. Descriptions of a new genus and five
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Surveying Voyage of H.M.S. Fly ... in Torres
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Prir 73(3): 57-62.

1

NEW SPECIES OF CHLAMYDOPSIS (HISTERIDAE: CHLAMYDOPSINAE), WITH A
REVIEW AND PHYLOGENETIC ANALYSIS OF ALL KNOWN SPECIES

MICHAEL S. CATERINO

Caterino, M.S. 2003 06 30: New species of Chlamydopsis (Histeridae: Chlamydopsinae),
with a review and phylogenetic analysis of all known species. Memoirs of the Queensland
Museum 49(1): 159-235. Brisbane. ISSN 0079-8835.

Forty new species of Chlamydopsis are described and the 29 previously described species are
reviewed. One of the new species is the second known New Guinean Chlamydopsis. The
remainder are Australian, with the highest diversity from Queensland. Six species groups are
proposed and three species are left unplaced. A phylogenetic analysis of adult morphology
provides support for several of these groups, and offers clues to the placement of the
enigmatic species. This study revealed numerous species to be sexually dimorphic. The
cladistic analysis indicates that some of these dimorphisms have arisen independently in
several lineages. This study synonymises C. excavata Lea and C. puncticollis Oke with C.
reticulata Lea, and C. atra Lea with C variolosa Lea. The following new species are
described: C. rana sp. nov., C. antennata sp. nov., C. trichonota sp. nov., C. dimorpha sp. nov.,
C. monteithi sp. nov., C. setifera sp. nov., C. lawrencei sp. nov., C. convergens sp. nov., C.
coronis sp. nov., C. erupta sp. nov., C. transversa sp. nov., C kununurra sp. nov., C. acutricha
sp. nov., C. myrmecophila sp. nov., C. mallee sp. nov., C. pecki sp. nov., C. degallieri sp. nov.,
C. jayawijaya sp. nov., C. lucifer sp. nov., C. bataviae sp. nov., C. bumetta sp. nov., C.
zborowskii sp. nov., C. pluriseta sp. nov., C. contorta sp. nov., C. pilosipes sp. nov., C. nielseni
sp. nov., C. australis sp. nov., C. lepida sp. nov., C. convexa sp. nov., C. dispersa sp. nov., C.
weiri sp. nov., C. crowcrofti sp. nov., C. macmillani sp. nov., C. nullarbor sp. nov., C. rotunda
sp. nov., C. carinota sp. nov., C. storeyi sp. nov., C. matthewsi sp. nov., C. mareeba sp. nov., and
C.parallelus sp. nov. □ Coleoptera, Histeridae, Chlamydopsis, myrmecophily, tactile mimicry.

Michael S. Caterino, Santa Barbara Museum of Natural History. 2559 Puesta del Sol Road,
Santa Barbara, CA 93105 USA, (e-mail: mcaterino@sbnature2.org); 7 November 2002.

The Chlamydopsinae is a remarkable group of
histerid beetles. They apparently all live in the
colonies of social insects, mainly ants, and
exhibit a tremendous diversity of morphological
specialisations for this lifestyle. The group’s
distribution is centred in Australia, although
several lineages have spread and diversified
throughout southeastern Asia, reaching India in
the west, Japan to the north, and Fiji to the east
(Caterino, 2000; Degallier, 1984; Nishikawa,
1 995, 1 996). Until recently the group has received
little study. However, recent collecting efforts,
particularly those incorporating flight interception
traps, have revealed a wealth of unrecognised
diversity. Here 40 new species are described, the
taxonomic status of previously described species
is reviewed, and the phylogenetic relationships
among all known Chlamydopsis Westwood, are
investigated. A recently discovered new species
of Chlamydopsis from New Caledonia is not
described here but is included in the phylogenetic
analysis.

Something of a quandry is faced in presenting
the species. While one of the goals of the phylo-
genetic analysis is to provide some framework

for an intrageneric classification, it is clear from
the outset that complete phylogenetic resolution
will not be obtained from existing data. Too many
species are known from only a single sex or even
a single specimen, and many important data are
therefore missing. For the purposes of facilitating
identification and placement of additional
material, a series of species groups is established.
Some of these appear likely to be monophyletic,
whereas putative synapomorphies of others are
clearly weak. The phylogenetic analysis
presented below will provide some preliminary'
indications of the relative values of morph-
ological characters in the group. However,
establishing a solidly phylogenetic classification
will require much additional material and study.

CONVENTIONS

All species accounts include diagnoses and
type data. For groups of closely related species,
generally only one is fully described, with the
remainder diagnosed from the first. In each
treatment of a previously described species type
locality, type repository, whether or not the type
has been examined by the author (where it has

160

MEMOIRS OF THE QUEENSLAND MUSEUM

been, label data are quoted), and additional
records, with either a repository or literature
reference for each record, are presented. Nontype
records are listed by source to distinguish
specimen records from unverified literature
records. Host ant names are as given by original
sources. See Table 1 for equivalents in current
formicid nomenclature.

A number of body dimensions and proportions
are useful for species recognition. Following
histerid conventions, total body length (L) is
measured from the anterior margin of the
pronotum to the posterior margin of the elytra,
while width (W) is taken at the widest point,
invariably near the elytral humeri. Measurements
were made of the holotype where possible, are
grouped at the beginning of each description (or
diagnosis if no description is presented) to
facilitate comparisons, and are abbreviated as
follows: L (mm - dorsal length along midline); W
(mm - width across humeri); E/PnL (ratio -
elytral length/pronotal length); E/PnW (ratio -
elytral width/pronotal width); Pn W/L (ratio -
pronotum width/length); E LAV (ratio - elytra
length/width); Pr/Py (ratio - Propygidium
Iength/'pygidium length); Sterna - pro, meso,
meta (mm - lengths along midline); Tibiae -
pro,meso,meta (mm - straight line length from
base to apex, ignoring curvature). Some
measurements are missing due to inaccessibility
of material or relevant body parts. Accepted
terminology is lacking for many unique
chlamydopsine features. Terms adopted in this
paper seem largely self-explanatory with one
exception. The depression surrounding the
prothoracic leg is margined by a stria originating
at the apex of the prosternal keel, curving
obliquely toward the anterolateral prosternal
corner, thence curving variously toward the
posterolateral prosternal corner. Regardless
significant variation in exact orientation and
degree of impression (which may render it more
carinalike than striate) this is termed the
circumcoxal stria.

Repositories are abbreviated as follows:
Australian Museum, Sydney (AMS); Australian
National Insect Collection. Canberra (ANIC);
Henry and Anne Howden Collection (HAHC);
Queensland Department of Primary Industries,
Mareeba (DPIM); Museum of Comparative
Zoology, Harvard University (MCZ); Michael
Caterino Collection (MSCC); Museum Victoria,
Melbourne (MVM); The Natural History
Museum, London (NHM); Queensland Museum,
Brisbane (QMB); South Australian Museum,

TABLE 1. Several host names have changed since
their chlamydopsine association was first reported.
Although some ambiguities have been encountered,
the following equivalences appear valid (following
Shattuck & Barnett, 2001). Multiple valid names
indicate that the original species has been split into
several. Those potentially sympatric with the
beetle(s) are listed. Names not listed apparently
remain valid as originally given.

Published Name

Valid Current Name

| Chalcoponcra metallica

Rhylidoponera metallica (Smith)

Ectatomma metallicum

Rhytidoponera metallica (Smith)

Rhvtidoponera convexa var.
violacea

R. violacea (Forel)

Euponera lutea

Pachvcondvla lutea (Mayr)

Iridomyrmex delectus

I. purpureus (Smith), I. sanguineus
Forel. or /. viridiaeneus Viehmeyer

Aphaenogaster longiceps

A. 1. longiceps (Smith)

Notoncus foreli

N. ectatommoides (Forel)

Meranoplus hirsutus

M. minor Forel

[formerly M. hirsutus minor]

Adelaide (SAM); Staatliches Museum fur
Naturkunde, Stuttgart (SMNS); United States
National Museum, Washington (USNM); Western
Australian Museum, Perth (WAM). fc QMT’
registration numbers are given for holotypes
deposited in QMB.

Within material lists, holotype data are quoted
exactly but for other material states and various
geographical features are abbreviated as follows:
Australian Capital Territory (ACT); Queensland
(Qld); Northeastern Queensland (NEQ);
Southeastern Queensland (SEQ); New South
Wales (NSW); Victoria (Vic); South Australia
(SA); Western Australia (WA); Northern
Territory (NT); rainforest (RF). Collectors are
abbreviated as follows: J. Brown (JB); A. Calder
(AC); D.J. Cook (DC); S. De Faveri (SD); K.
Halfpapp (KH); J. Hasenpusch (JH); H. Janetzki
(HJ); J.F. Lawrence (JL); L. Miller (LM); G.B.
Monteith (GM); E.S. Nielsen (EN); E. Schmidt
(ES); S. Shattuck (SS); R. Storey (RS); G.I.
Thompson (GT); L. Umback (LU); M.S. Upton
(MU); T. Weir (TW); P. Zborowski (PZ).
Collection methods abbreviated as: Flight
Interception trap (FIT); Malaise trap (MT);
pitfall trap (PT). Collection dates are given as
day.month.year, with month represented by
lower case roman numerals (e.g., l .vi.2002).

Chlamydopsis Westwood, 1869

Chlamydopsis Westwood, 1869: 317.

Byzenia King, 1869: 74 (type species: Byzenia formicicola
King); Blackburn, 1891: 92.

REVIEW OF CHLAMYDOPSIS

161

TYPE SPECIES. Chlamydopsis striatella Westwood,
1869: 318; designated by Lewis, 1903: 428.

REMARKS. Species of Chlamydopsis are very
diverse in morphology, and the genus cannot be
defined at present by any definite synapo-
morphies. It may well prove to be paraphyletic
with respect to at least one or two other genera.
The species currently contained in Chlamydopsis
all share a visible scutellum and an upturned
anterior pronotal margin, at least above the
antennal cavities, if not along the entire margin.
None are entirely flattened dorsally (like
Ectatommiphila Lea) and none have the
strikingly elongate trichome setae seen in
Eucurtia Mjoberg.

STRIA TIPENNIS GROUP

The striatipennis group is founded primarily
on trichome morphology. All share a well
developed humeral trichome in which the outer
and upper surface are continuously rounded, with
the trichome open only mesally and along the
lower anterior edge. The humeri are variously
enlarged, often with the anterior comers broad
and angular, apparently hollow within (the lateral
surface being somewhat translucent), with a very
short, inconspicuous anterior superficial stria,
and lined along the inner, upper edge with a
single longitudinal or oblique row of setae, which
extends to near the anterior elytral margin. A
protuberance arises from the elytral disk beneath
the anteromesal comer of the trichome, which
meets the dorsal fringe, or projects above it
forming an erect, rounded lamina (Figs 1A-G,
281). Generally a separate ‘whorl’ of setae can
also be seen within the mesal opening of the
trichome, just beneath the longitudinal fringe.
The anterior prostemal margin is striate but
ungrooved. The metatibiae are elongate and/or
widened in many. The central portion of the
anterior pronotal margin is usually separated by a
notch from the anterolateral portions, with the
notch continued behind the lateral portion by a
stria or groove. The species C. rana sp. nov., C.
antennata sp. nov., and C. trichonota sp. nov. are
placed here somewhat tentatively, based on
trichome structure. However, the prostemal
marginal stria of these three is well impressed,
and appears more groovelike than in the rest of
the group. Whatever their relationships to the
group, these three are all easily recognised as the
only Chlamydopsis with lateral pronotal
trichomes (Fig. 1E-G)

Chlamydopsis striatipennis Lea, 1919
(Figs 1A, 2A, 3, 28A, 291)

Chlamydopsis striatipennis Lea, 1919: 177; Type locality:

Vic: Lome; repository: SAM; not examined.

RECORDS. NHM: Vic: Warburton, Fem Tree Gully. Lea
(1925): Vic: Lakes Entrance, Oct., ‘small black
Iridomyrmex ’; Vic: Beaconsficld, with Ectatomma
metallicum. Oke (1923): Whittlesea; Warburton: Femtree
Gully; Belgrave; Emerald [all Vic], ANIC: ACT: 35.19S
148,5 IE, Wombat Ck„ 6km NE of Piccadilly Circus,
750m, i. 1 985; NSW: Mt Keira, Wollongong, ii.1981;
NSW: Lorien W.R. 3km N LansdowneHaree, 22.xi.l987,
ex r/f margin, wet scler. forest FIT. DPIM: NSW: 3km N.
Lansdowne via Taree, 25.1.1987, MT, rainforest maigin.
WAM: NSW: Wollongong, 1938.

DIAGNOSIS. L: 2.62; W: 0.87; E/Pn L: 2.0; E/Pn
W: 1 .62; Pn W/L: 1 .50; E L/W: 0.82; Pr/Py : 1.18;
Sterna: 0.69,0.19, 0.93; Tibiae: 1.18, 1.25, 1.62.
Chlamydopsis striatipennis appears to comprise
one of the more distinctive, temperate offshoots
of a varied complex of populations centred in the
more tropical parts of eastern Australia. Thus far
samples have not been sufficient to fully resolve
species limits within this complex, though
certainly more than one species is present. The
populations which constitute C. striatipennis s.
str. have the humeri strongly angular (produced
laterally almost perpendicular to the pronotal
margin), and have the elytra setose mainly on the
anterior surface of the trichome and along the
apical margin. They are also slightly larger in
body size than average for the complex (among
examined samples). However, humeral shape
and pilosity vary tremendously across this
complex, and it might eventually be desirable to
define the species more broadly.

REMARKS. In addition to the records presented
above (pertaining solely to C. striatipennis s. str.)
specimens considered members of the
'‘striatipennis complex' have been collected
throughout Queensland, as far north as 12°S.
Specimens from southeastern Queensland
(27°20’S, 152°48’E, Stony Ck) are particularly
close to ‘typical’ striatipennis. At the same time a
few additional specimens from Vic (Bonang,
Warburton) and NSW (Lansdowne; Wilson River
Reserve) do not conform to the strict definition of
C. striatipennis , as recognised here. A
particularly distinctive form is known from
several upper elevation localities (> 1 1 00m) in the
vicinity of the Hugh Nelson Range (Qld:
17°27’S, 145°29’E). However, even among the
few localities represented, there is variation that
should be better represented before an additional

162

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1 . Dorsal views of Chlamydopsis spp. A, C. striatipennis. B, C. leai. C, C. compressipes. D, C. pallida. E, C.
rana. F, C. antennata. G, C. trichonota. H, C. reticulata (d). I, C. reticulata (9).

REVIEW OF CHLAMYDOPSIS

163

species is described. It is also possible that some
of the remaining named species in this species
group are derived from within the complex,
particularly C. leai and C. compressipes.

Chlamvdopsis leai Oke, 1923
(Figs IB, 2B, 3)

Chlamydopsis leai Oke, 1923: 155; Lectotype 9, hereby
designated: Belgrave, Vic., 13.12.1 920, C. Oke/
Chlamydopsis leai Oke Type/ Presented by C.G. Oke/ 895
Type, MVM. Paralectotype: Belgrave, Vic., July 1921,
C.Oke 2.7.21/ Chlamydopsis leai , Oke, Co-Type/ 896
Paratypc; in MVM; examined, 2000.

DIAGNOSIS. This species is a not-very-
distinctive member of the striatipennis complex.
The most distinctive character is its fairly broad,
arcuate meso- and metatibiae (shared with C.
compressipes , below). The upper inner edge of
the trichome is slightly more oblique than typical,
incising the humeral elevation posteriorly,
although this is true of C. striatipennis s. str. as
well. The species needs to be included in any
broader study of variation in this complex.

REMARKS. Reported from nests of Iridomyrmex
sp. (Oke, 1923). This species is known only from
the type series.

Chlamvdopsis compressipes Lea, 1919
(Figs 1C, 2C, 3)

Chlamydopsis compressipes Lea, 1919; Type 9:
compressipes [handwritten] Lea, Type, Mt Tambourine/
C/2086 / Type/ 10678, Chlamydopsis compressipes Lea,
Queensland, QMB; examined, 2000.

RECORDS. QMB: SEQ: Mt Glorious, i-iii. 1982 and
ix-x.1990; CMN: Qld: Mt Glorious, 27°20’S,152°49’E,
3-9.X.1998, N. Power, MT.

DIAGNOSIS. This species is most easily recog-
nised by its longitudinal laminae projecting up
above the inner edges of the humeral trichomes.
This is seen to a much lesser degree as well.
However, in the latter species the laminae are
oblique and do not project above the elevated
humeri in C pallida. The humeri arc also much
narrower, and the mediobasal elytral depression
thus broader.

DESCRIPTION. L: 1.99; W: 0.75; E/Pn L: 1.67;
E/Pn W: 1 .29; Pn W/L: 1 .42; E L/W: 0.91 ; Pr/Py:
1.13; Sterna: 0.50, 0.12, 0.62; Tibiae: 0.87, 0.93,
1.18. Body elongate, orange, mostly glabrous.
Frons about 1 .2 X as long as wide, sides weakly
arcuate, disk reticulopunctate, with a few short,
inconspicuous setae; labrum broad, apical margin
weakly bilobed, with a few short setae; antennal
scape angulate near middle, rounded at apex;

antennal club of female about two-thirds length
of scape, that of male about 1 .8 x length of scape.

Pronotum about 1 .5 x as wide as long, sides
margined, parallel in basal two-thirds, acutely
widened and somewhat elevated anteriorly;
lateral portions of anterior margin strongly
elevated, arcuate, separated from lower central
portion by stria which arcs behind base of lateral
elevation, then curved posteriorly, meeting lateral
pronotal margin at about the basal one-third;
pronotal disk slightly depressed in anterior
comers, with an acute, slightly transverse median
tubercle, reticulopunctate throughout.

Prostemum with anterior margin sinuate, not
grooved; prostemal keel transversely depressed
at middle behind anterior margin, narrowed
between procoxae, but widening slightly at apex,
acutely emarginate, reticulopunctate throughout.

Elytra widest near humeri, humeri strongly
though rather narrowly elevated, widened
anteriorly, bluntly projecting forward at sides, the
inner edges laminate, elevated above humeri
from anterior edge to central trichome opening,
lamina with a longitudinal setal fringe closely
appresed to its outer surface, projecting very
slightly above it, this fringe extending anteriorly
to a short groove (probable homologue of
‘superficial groove’ of other species); central
trichome opening mesal, small, circular, with a
concealing fringe of downwardly directed setae;
mediobasal elytral depression smooth, with low,
blunt transverse carinae; elytral disks otherwise
shallowly but uniformly reticulostrigose, with a
few conspicuous setae near apex of humeri,
otherwise glabrous; humeri of male, including
inner lamina, generally less prominent.

Mesostemum about 4 X as wide as long,
projecting at middle, reticulopunctate;
mesometastemal suture well impressed; median
metastemal suture visible as a dark line, but not
impressed, disk evenly, but very finely punctate;
stemite 1 somewhat more coarsely punctate
along basal margin and near metacoxae.

Legs more or less slender, slightly elongate, all
tibiae of female with outer margins arcuate, their
outer surfaces (and those of the femora less
uniformly) densely but finely punctate, nearly
alutaceous; protibia of male more nearly angulate
near the base.

Propygidium weakly depressed along basal
margin, otherwise evenly convex, shallowly
reticulopunctate; pygidium convex, reticulo-
punctate in basal half, smooth apically.

164

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 2. Lateral views of Chlamydopsis spp. A, C. striatipennis. B, C. leai. C, C. compressipes. D, C. pallida. E, C.
rana. F, C. antennata. Q C. trichonota. H, C. reticulata (9).

REMARKS. The male and female specimens
listed from Mt Glorious are slightly more
elongate in body shape than the type of C.
compressipes. However, with these three as all
the known material of the species, they are
considered to constitute a single variable species.

Chlamydopsis pallida Lea, 1918
" (Figs ID, 2D, 3)

Chlamydopsis pallida Lea, 1918: 86; Lectotype, hereby
designated: New South Wales, Sydney, SAM; examined,
2000; 2 paralectotypes: same data as type, BMNH.

REVIEW OF CHLAMYDOPSIS

165

DIAGNOSIS. L: 2.31; W: 0.75; E/Pn L: 2.08;
E/Pn W: 1 .34; Pn W/L: 1.58; E LAV: 0.98; Pr/Py:
1.58; Sterna: 0.40, 0.16, 0.59; Tibiae: 0.87, 0.93,
1.15. Chlamydopsis pallida has very distinctive
humeral trichomes. They are narrow and close to
the anterolateral comers of the elytra, enclosing
very little subhumeral space. This species also
possesses a small oblique lamina (oriented
posterolaterally) on the inner edge of the humeral
trichome, around which the lateral portion
appears to be curved. The pronotum is also
distinctive, having the sides strongly elevated
and the anterolateral groove deeply impressed.

REMARKS. Reported from the nest of a ‘small
reddish ant’, identified, apparently tentatively, as
Meranoplus hirsutus. An ant mounted with the
paratypes is a Meranoplus , but 1 cannot con-
fidently determine the species identity. Assuming
the reported species is accurate, this would refer
to what is now called M. minor Forel (based on its
range), formerly M. hirsutus minor. This species
is only known from the type series.

Chlamydopsis rana sp. nov.

(Figs IE, 2E, 3, 30B)

MATERIAL. HOLOTYPE (QMT 108574) 6 : Windsor
Tableland via Mt Carbine, N Qld, 12.xi-26xii. 1983. Storey
& Walford-Huggins/ MDP1 Intercept Trap, Site No. 14a, in
QMB. PARATYPES (12): 4 9 : same data as type; 3 9 9 ,
1 6 : same locality as type but 26.xii.l983-24.i. 1984; 1 9 :
Windsor Tableland, N Qld., 27.xii.88-9.i.l989, ES &
ANZSES Site 5, FIT; 1 9 : N Qld, Windsor Tableland,
38km from main road, 28.xi-20.xii. 1985, RS&JB/ MDPI
FIT site 14c; 1 6 : NEQ: 15°48’S 145°1TE, MtFinnigan,
1 080m, 4.xii. 1 990- 1 7.i. 1 99 1 , QMB & ANZSES, FIT. Site
5; 1 9 : NEQ. I5°52’S, 145°14’E, Mt Misery summit,
850m, 6.xii. 1 990- 1 7.i. 1991, QMB & ANZSES, FIT Site 3,
in QMB, DPIM, MSCC.

DIAGNOSIS. This species is easily recognised
by the combination of oblique, setose pronotal
trichomes, and the broad, arcuate, elongate
metatibiae. The only other Chlamydopsis which
possess pronotal trichomes, C. antennata sp. nov.
and C. trichonota sp. nov., have them restricted to
the lateral margin, not forming an oblique setose
depression. The metatibiae of these two species
are also elongate, but not as broad as those of C.
rana , at least in the known males.

DESCRIPTION. L: 2.06; W: 0.81; E/Pn L: 1.54;
E/Pn W: 1 .37; Pn W/L: 1 .46; E LAV: 0.77; Pr/Py:
1 .00; Sterna: 0.56, 0. 1 2, 0.65; Tibiae: 0.87, 1 .03,
1.34. Body dark, slightly rufescent brown,
dorsally glabrous (except for trichomes),
surfaces varied from smooth to coarsely strigose;

frons with sides weakly rounded, about 1 .2 X as
long as wide, reticulopunctate, with a pair of
prominent apical marginal setae; labrum wide,
faintly bilobed; antennal scape widest about
one-third from base, rounded at apex; antennal
club of male about 2x as long as scape,
somewhat compressed; antennal club of female
slightly shorter than scape.

Prothorax with lateral margins interrupted near
front by notch-like trichome, the fringe of which
extends more sparsely posteromesally along an
oblique depression; anterior pronotal margin
elevated, with central and lateral portions more or
less continuous, though a fine stria extends from
the anterior margin along the upper edge of lateral
portions; pronotal disk smooth at middle, strigose
at sides and along anterior margin.

Prosternum finely grooved along anterior
margin, this groove diverging slightly from the
margin at sides; anterior margin arcuate, more
prominent at sides than at middle; disk of pro-
sternal keel uniformly coarsely reticulopunctate,
narrowed posteriorly, emarginate at apex.

Elytra 1.5 x as wide as pronotal base, humeri
strongly elevated, rounded at apices; setae of
inner longitudinal fringe of trichome short,
decumbent posteriorly, fringe arched above
central opening of trichome, extending around
front into horizontal anterior groove; central
opening of trichome nearly obscured by
downward pointing fringe of setae: dorsal aspect
of trichome with elongate coarse setae, elytral
disk otherwise glabrous, strongly
reticulopunctate on upper and lateral surfaces of
humeri, with a few transverse reticulae between
trichomes, disk becoming smooth
posteromedially; elytral marginal stria nearly
complete, interrupted only slightly in mediobasal
depression, not abruptly divergent from margin
above metafemora.

Mesosternum about 5 x as wide as long,
projecting at middle, coarsely punctate; meso-
metasternal suture deeply impressed;
metastemum with median longitudinal suture
finely impressed, disk with only a tew punctures
in front of metacoxae; stemite 1 sparsely punctate
at sides.

Femora slender, the metafemur about 1 .3 x
length of pro- and mesofemur; protibia slender
and angulate near basal one-third; mesotibia
slightly broader, widest just beyond midpoint;
metatibia elongate, broad, with outer margin
more or less evenly rounded, dimorphic, with that
of female about 1 .3 X as broad as that of male.

166

MEMOIRS OF THE QUEENSLAND MUSEUM

Propygidium and pygidium faintly convex,
both coarsely punctate, the pygidium becoming
smooth and setose in apical one-third.

REMARKS: The name of this species refers the
frog-like metathoracic legs, as well as its general
appearance.

Chlamydopsis antennata sp. nov.

(Figs IF, 2F, 3)

MATERIAL. HOLOTYPE (QMT 108575) 6 : Windsor
Tableland, N Qld., 27Dec 88-8 Jan 1989, E.Schmidt &
ANZSES, Site 6, fit. intercept, in QMB. PARATYPES: 2
6 6 : same data as holotype, in QMB.

DIAGNOSIS. As above, only this species, C.
rana , and C. trichonota possess pronotal
trichomes. In this species and the following, these
are formed by short arcuate outgrowths of the
lateral margins which meet externally to enclose
a setose opening at the anterolateral pronotal
comer, whereas in C. rana an oblique setose
dorsal depression leads to a simple anterolateral
notch. Chlamydopsis antennata and C.
trichonota are very similar, and obviously closely
related. Chlamydopsis antennata appears (from
limited material) to be slightly smaller and, more
significantly, the setose inner edge of the humeral
trichome incises the elevated humerus more
deeply, forming a distinct mesal emargination
(Fig. IF vs 1G).

DESCRIPTION. L: 2.12; W: 0.75; E/Pn L: 1.83;
E/Pn W: 1 .42; Pn W/L: 1 .58; E LAV: 0.81; Pr/Py:

1.06; Sterna: 0.50, 0.12, 0.65;
Tibiae: 0.93, 1.06, 1.43. Body
dark, faintly rufescent, almost
entirely glabrous. Frons 1.3 x
as long as wide, sides rounded,
widest at middle, glabrous,
entirely reticulopunctate;
labrum broad, apical margin
only slightly arcuate, with a
few inconspicuous setae;
antennal scapes with outer
margins almost evenly arcuate,
widest near middle, reticulo-
punctate; antennal club (of
male) very large, about 3.5 x
as long as scape, apices project-
ing even when fully retracted,
strongly compressed.

Prothorax about 1.5 x as
wide as median length; lateral
margins with trichome, a small
setose circular opening
enclosed by anterior and
posterior outgrowths of lateral margin, inflated
base of posterior lobe with separate lateral setal
fringe; anterior pronotal margin elevated, with
central and lateral portions continuous, sinuate
around large antennal cavities; pronotal disk
glabrous, impunctate, shining.

Elytra with humeri strongly, rather narrowly,
elevated in basal half, inner edge of each
elevation emarginate, lined with continuous
fringe of conspicuous setae concealing central
opening of trichome, this fringe shorter anterior
to emargination, extending anteriorly along inner
edge around to anterolateral comer where it is
longer, opposing posterolateral fringe of
pronotum; mediobasal depression with strong
transverse carinae; each elytron impunctate
along suture, rather coarsely strigose at sides, the
strigae converging (mesally and laterally) to apex
of trichome; elytral marginal stria nearly
complete, interrupted only slightly in mediobasal
depression, not abruptly divergent from margin
above meta femora.

Prostemum with anterior marginal stria deeply
impressed, diverging from margin at sides,
ending beneath anterior edge of pronotal
trichome; prosternal disk shallowly reticulo-
punctate, narrowed posteriorly, keel depressed
between procoxae, apex deeply and acutely
emarginate.

Mesostemum short, about 6 x as wide as long,
projecting at middle, with a single row of

REVIEW OF CHLAMYDOPSIS

167

punctures; mesometasternal suture deeply
impressed; metastemal disk impunctate; longi-
tudinal metastemal suture faintly impressed;
visible stemite 1 impunctate.

Legs all relatively slender, slightly elongate,
the protibia and mesotibia about equal in length,
outer margins angulate one-third from base, the
metatibia 1 .3 x as long, with outer margin evenly
arcuate.

Propygidium arcuately depressed along basal
margin, otherwise convex, shallowly but evenly
reticulopunctate; pygidium evenly convex,
reticulopunctate in basal half, smooth and with a
few setae at apex.

REMARKS. Only males of this and the
following species are known. Given the sexual
dimorphism in metatibial shape, and possibly
color, in the preceding species, the females of
these species may differ slightly from the
descriptions. The name of this species refers to
the enormous antennal club of the males.

Chlamvdopsis trichonota sp. nov.

(Figs 1G, 2a 3)

MATERIAL. HOLOTYPE (QMT 108576) 8: Mt Lewis
Rd, NEQld, 16km from Highway, 18 Dec 1989-13 Jan
1 990, Monteith, Thompson, ANZSES, Site 2, 950m, Fit.
Intercept. PARATYPE: 1 8 : same data as type, in QMB.

DIAGNOSIS. L: 2.31; W: 0.81; E/Pn L: 1.85;
E/Pn W: 1 .45; Pn W/L: 1 .54; E LAV: 0.83; Pr/Py:
1.13; Sterna: 0.62, 0.12, 0.72; Tibiae: 1.00, 1.06,

1 .37. This species is very closely related to the
preceding, and is therefore not fully described
here. It differs most significantly in the form of
the trichome. Its inner edge in this species is
barely emarginate, the setal fringe arcuate
posteriorly, but much closer to the typical straight
line fringe of the striatipennis group. In addition,
the elytra of this species are very sparsely, but
evenly clothed with fine setae. Chlamydopsis
antennata has at most a few decumbent hairs on
the anterior surface of the trichome, but none
elsewhere. Otherwise the two species appear
virtually identical.

REMARKS: The name of this species refers to its
pronotal trichomes.

STRIGICOLLIS GROUP

The strigicollis group contains seven
apparently relatively generalised species. They
are characterised by an anterior superficial
humeral groove that extends more or less
obliquely and horizontally from the humeral

elytral comer inward to the mesal base of the
trichome. They are further restricted to those
species in which anterior prostemal marginal
stria does not depart from the margin to meet the
circumcoxal stria. This marginal stria does not
fonn a conspicuous groove in the first four
species included here. But it does in the last three,
which for that reason are included somewhat
tentatively. The fonn of the trichome suggests
relationships with the pygidialis group. However,
the latter is so readily characterised that both are
maintained as separate groups until a clearer
phylogenetic picture emerges. The first three
species of this group exhibit an oblique groove
running behind the lateral portion of the anterior
pronotal margin. This is identical in form to that
of most of the striatipennis group, although this
seems likely to be a symplesiomorphy.

Chlamvdopsis reticulata Lea, 1910
(Figs 1H-I, 2H-I, 6, 29D)

Chlamydopsis reticulata Lea, 1910: 199; Material. Holotype
(unique): reticulata Lea, Type, N.S.W./, on the underside
of mounting card: reticulata Lea TYPE, from King's coll./
Chlamydopsis reticulata Lea, Type, Australia: SAM,
examined, 2000; these specimen data conflict with Lea's
original citation of the type specimen: ‘Australia (a single
specimen, without locality label, from the late Rev. R. L.
King's collection)'; it is thus unclear why ‘N.S.W. ’
appears on the type label.

Chlamydopsis excavata Lea, 1910: 200; Type: Tasmania,
near Hobart; SAM, not examined; New Synonymy.
Chlamydopsis puncticollis Oke, 1923: 156; Lcctotype 8,
hereby designated: Femtree Gully, 26.5.1920, C. Oke,
Vic./ Chlamvdopsis puncticollis Oke Type/ Presented by
C.G. Oke/ '897 Type; Paralectotype 8: same locality,
20.6.20, 898 Paratype; MVM. examined, 2000; New
Synonymy.

RECORDS. AN1C: ACT: 35.16S. 149.06E, Black Mt,
600m, x. 1 987. NHM: NSW: Sydney [as C. excavata, dct.
by A. Lea], Lea (1919): NSW: Hunters Hill (nr Sydney),
Oct., nest of Ectatomma [as C. excavata]', Vic: Fern Tree
Gully, Dec. [as C. excavata]. Lea (1925): NSW: Como,
Ectatomma metaUicum. Oke ( 1 923): NSW: National Park;
Vic: Bcaconsfield; Vic: Belgrave; Vic: Femtree Gully [as
C. puncticollis]. MCZ: Vic: Beaconsfield.

DIAGNOSIS. L: 2.15; W: 0.72; E/Pn L: 2.00;
E/Pn W: 1 .39; Pn W/L: 1 .57; E L/W: 0.92; Pr/Py:

1 .00; Sterna: 0.62, 0. 1 2, 0.69; Tibiae: 0.8 1 , 0.87,
1.00. This and the following species are very
similar. Both exhibit sexual dimorphism in the
sculpturing of the elytra, with the males being
reticulostrigose everywhere outside the medio-
basal depression, and the females appearing
almost entirely impunctate. In both sexes the
pronotal texture differs slightly between the two
species, with the strigae of C. reticulata more
consistently impressed from edge to middle. In

168

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Dorsal views of Chlamydopsis spp. A, C. dimorpha (S). B, C. dimorpha (9). C, C. strigicollis. D, C.
mormolyce. E, C. monteithi ( 6 ). F, C. monteithi ( 9 ). G, C. setifera ( 9 ). H, C. lawrencei ( 6 ). I, C. pygidialis.

most specimens of C. dimorpha the often impunctate at the centre. While C.
strigae/punctures of the median portion of the reticulata is consistently slightly larger than C.
pronotal disk are less deeply impressed, and it is dimorpha , this is especially evident in the elytra

REVIEW OF CHLAMYDOPSIS

169

of the females, with those of C. reticulata much
broader relative to the pronotum. Possibly the
most consistent, though least substantial,
difference is that the mediobasal depression of C.
dimorpha always possesses a minute setigerous
pustule at the sides near the trichome
(occasionally more than one), whereas in all C.
reticulata examined so far, the mediobasal
depression is bare (except along the fine
transverse carina).

REMARKS. The new synonymies proposed here
are based partly on the sexual dimorphism
discovered in this species. In fact Lea (1910)
recognised the strong structural similarity
between his reticulata and excavata , but had no
reason to expect such a striking difference to be
sexual . Oke ( 1 923 ), however, seems to have been
simply unfamiliar with C. reticulata , as it is not
mentioned in his description of C. puncticollis.

Chlnmydopsis dimorpha sp. nov.

(Figs 4A-B, 5A, 6)

MATERIAL. HOLOTYPE (QMT108577) 6 (dissected
by the author): NEQ: 17°26’S, 145°42 ? E, Hughes Road
Topaz, 6 Dec 1 993-25 Feb 1 994, Monteith, Cook, Janetzki,
RF Intercept, 650m. PARATYPES (20): 1 1 <3, 1 9: same
data as holotype; I 9 : SEQ: 27°20’S, 1 52°48’ E, Stony Ck.,
via Samford 22.x.94-2.ii. 1 995, HJ&GM, RF FIT; 1 9:
same as preceding but 2.ii-8.iv. 1 995, open forest FIT; 1 6 :
NEQ: 17°24'S, 145°4rE, Westcott Rd, Topaz,
6.xii.93-25.ii.l994, GM,DC,HJ, RF FIT, 680m; 1 9:
NEQ: 17°24’S, 145°4rE. PEI Rd, Topaz, 6.xii.93-
25.ii.1994, GM.DC,HJ, RF FIT, 580m; I 3: NEQ:
Danbulla SF, 13km NE of Yungaburra, 20.xii.86-
13.1.1987, RS&SD, MDPI FIT site 27; 1 6 : Qld: 17.28S
145.29E, Longlands Gap BS1, 1150m 3.i-5.ii.l995, PZ,
FIT; 1 9: SEQ: 25°40’S 151 0 25'E, Nipping Gully, Site 2,
9.x- 18.xii. 1998, GM&Gough, RF FIT, 200m, 7399; 1 <J:
Qld: 28°08'S 152°40’E, Black Rock Scrub, 350m,
2.xii.2000- 1 3.V.200 1 , GM, RF FIT, 10162, in QMB,
DPIM, ANIC, MSCC.

DIAGNOSIS. See diagnosis under C. reticulata ,
above.

DESCRIPTION. L: 1.99; W: 0.69; E/Pn L: 1.91;
E/Pn W: 1 .29; Pn W/L: 1 .55; E LAV: 0.95; Pr/Py:
0.93; Sterna: 0.56, 0.12, 0.59; Tibiae: 0.75, 0.87,
0.87. Body dark reddish brown, subquadrate.
Frons slightly wider than long, sides
approximately parallel in apical 2/3, narrowed at
antennal bases and at apex; with strongly
developed reticulate sculpturing, the individual
cells varied but most elongate oval or elongate
polygonal; labrum semicircular, faintly rugose,
with a few elongate setae. Antennal scapes with
outer edges bluntly angulate, widest just apicad

of midpoint, reticulately sculptured as frons;
antennal club of female oval, approximately
equal in length to funicle; antennal club of male
more elongate, about 1 .5 x as long as funicle.

Pronotum almost 2 x as wide as long,
margined laterally, sides very slightly arcuate,
but more or less parallel, anterior margin
tripartite, in lateral thirds (above antennal
cavities) raised nearly perpendicular to disk,
oblique and slightly inwardly arcuate to side, in
middle third less strongly raised, the middle
section separated from lateral oblique sections by
a groove which continues posterolateral ly along
base of raised lateral margin, curving posteriorly
near side where it merges with lateral pronotal
margin; pronotal disk reticulately sculptured, the
reticulae elongated and oblique anterolateral ly,
less strongly sculptured posteromedially,
especially in female.

Prostemum with reticulately sculptured lobe
and keel separated from smooth depressions for
reception of legs by a strong carina extending
from lateral sternal margin to base of keel; keel
strongly narrowed posteriorly, acutely
emarginate at base; keel set off from lobe by a
weak transverse furrow at middle, reminiscent of
(though probably not homologous with) the
prestemal suture of many histerids; anterior
margin of prosternum simple (with weak
marginal stria but not grooved) weakly arcuate at
sides, transverse and faintly undulating at middle.

Each elytron about 2 x as long as wide, widest
just behind middle; transversely depressed across
middle at base, trichome with anterior and
posterior portions strongly elevated; anterior
elevation with anterior superficial groove
strongly oblique from anterior elytral comer to
near base of inner edge of trichome, bearing a
dense continuous fringe of golden setae from this
groove up inner edge of trichome to its apex, the
setae apparently increasing in length towards
apex, the apical-most setae extending oyer the
gap between anterior and posterior elevations of
trichome; gap between elevations incised nearly
to their bases internally and externally, the
external incision a narrow, inverted ‘Y’ shape,
lacking setae, the inner incision broadly open,
with anteriorly directed setae on posterior edge
from near base to apex; sculpturing of elytral disk
strongly sexually dimorphic: that of male with
elongated reticulae everywhere except in
depression between trichomes, that of female
absolutely smooth except on epipleuron and
anterior slope of trichome which are reticulate as
in male; elytral disk of both sexes with few

170

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Lateral views of Chlamydopsis spp. A, C. dimorpha (d). B, C. strigicollis. C, G mormolyce. D, C.
monteithi (d). E, G monteithi ( 9 ). F, C. setifera ( 9 ). G, C. lawrencei (d). H, G pygidialis.

conspicuous setae along basal half of elytral
suture, and a few at sides within the medial
depression; marginal stria of elytron continuous
along all edges except along base and medial
suture near scutellum.

Mesostemum about 3 x as wide as long,
acutely projecting at middle, with complete
anterior marginal stria, posterior margin sinuate,
raised relative to anterior portion of meta-
sternum, disk with scattered punctures and dense

REVIEW OF CHLAMYDOPSIS

171

intervening microsculpture. Metasternum
smooth, without hint of microsculpture, with
only a few weak punctures along anterior edge
and scattered inconspicuous setae. First visible
abdominal stemite similarly smooth and with
faint anterior marginal punctures.

Prothoracic legs densely reticulate on exposed
surfaces, tibia slender, with outer edge angulate
one-third from base; meso- and metathoracic legs
smoother, with only faint microsculpture along
outer tibial margins; posterior tibiae slightly
broader and more rounded than protibia, tibiae of
female markedly broader and rounder, and with
microsculpture confined to a discrete marginal
band.

Propygidium shallowly depressed at middle of
anterior margin, otherwise uniformly convex,
densely reticulate throughout; pygidium weakly
convex, with weakly developed reticulae in basal
one-third, otherwise smooth.

REMARKS. The name of this species refers to
the marked difference between the sexes.

Chlamvdopsis strigicollis Oke, 1923
(Pigs 4C, 5B, 6)

Chlamydopsis strigicollis Oke, 1923: 157; Lectotype, hereby
designated: Bclgrave, V., 17.10.21, C.Okc/ Chlamydopsis
strigicollis Oke, Type/ Presented by C.G. Oke/ 899.

MATERIAL. HOLOTYPE 6; MVM, examined, 2000.
Paralectotype: same data as lectotype; in MVM.

RECORDS. Oke (1923): Vic: Hurst’s Bridge; Vic:
Femtree Gully; Vic: Beaconsfield; Vic: Mooroolbark.
WAM: Vic: Femtree Gully. ANIC: ACT: 35. 1 9S, 1 48.5 1 E,
Wombat Ck, 6km NE Picadilly Circus, i. 1 984, 750m; Vic:
Withers, 2-42; NSW: 32.08S, I51.27E, Ailvn River,
Chichester SF, I0-Il.xi.l981. MCZ: Vic: Arthur’s Seat,
900ft., 29.iv.1951, ‘ w / Chalcoponera victoriae \

DIAGNOSIS. L: 1.93; W: 0.75; E/Pn L: 1.58;
E/Pn W: 1 .35; Pn W/L: 1 .42; E L/W: 0.83; Pr/Py:
1.21; Sterna: 0.56, 0. 1 2, 0.44; Tibiae: 0.69, 0.69,
0.69. Among members of this group, C.
strigicollis is unique in having the anterior
pronotal margin barely upturned relative to
dorsum. It also lacks the oblique lateral groove
behind the antennal cavities, present in C.
reticulata and in C. dimorpha. The pronotal
texture and setation are also unique; the disk is
reticulostrigose, with the strigac and the long,
decumbent setae convergent anteromedially.
Interestingly, C. strigicollis does not exhibit the
elytral dimorphism shown by C. reticulata and in
C. dimorpha ; both sexes are uniformly reticulo-
strigose.

REMARKS. Reported from nests of Chalcoponera
sp. (Oke, 1923).

Chlamvdopsis mormolyce Lea, 1925
(Figs 4D, 5C, 6)

Chlamydopsis mormolyce Lea, 1925: 255; Lectotype, hereby
designated: Mormolyce, Mundaring. Lea. Type/
Chlamydopsis mormolyce Lea Type, W. Australia,
mounted with host ant; SAM; examined, 2000.

RECORDS. WAM: WA: Culham, xi.1960, nest of
Chalcoponera inomata.

DIAGNOSIS. L: 2.18; W: 0.84; E/Pn L: 1.59;
E/Pn W: 1 .5 1 ; Pn W/L: 1 .30; E L/W: 0.8 1 ; Pr/Py:
1.11; Sterna: 0.65, 0. 1 6, 0.59; Tibiae: 1 .00, 1.12,
1.21. While C. mormolyce shares the orientation
of the anterior superficial humeral groove with
the other members of this group, the actual form
of the trichome differs somewhat from the others.
Firstly its upper mesal edge, densely fringed with
setae in the other species, is bare; the only
trichome setae are within its opening, barely
visible from above. The anterior surface ot
trichome is broad, slightly concave. There is also
no lateral gap between the anterior and posterior
humeral elevations. Although a conspicuous
shallow groove marks the junction, they are
solidly joined to their dorsal apices. A tew
additional distinguishing characters include:
pronotal reticulae all more or less polygonal,
none elongate; prosternum bearing setae;
metasternum and stemite 1 faintly reticulate
throughout, more conspicuously towards the
anterior margin of each: elytral disks with
elongate reticulation, and long, though sparse,
setae; legs of female all elongate, slender and
reticulate (posterior ones not smooth); inner apex
of meso- and metatibiae with an acute, fixed
denticle.

REMARKS. The numerous differences between
this and the other species ol the strigicollis group
leave some doubt that it belongs here phylo-
genetically. Its relationships will require
reassessment with more extensive samples.

Chlamvdopsis monteithi sp. nov.

(Figs 4E-F, 5D-E, 6, 28B)

MATERIAL. HOLOTYPE (QMT1 08578) 6: C.QId
26°57*S 148°02’E, Mt Moffatt NP. 1000m, Mahogany
Forest, 26 Sept-26 Nov 1995, G Monteith. Intercept.
PARATYPES: 7 9 9, same data as type, in QMB, DPIM,
MSCC.

DIAGNOSIS. This and the following two species
exhibit unusual anterior prostemal margins. Its
marginal stria does not diverge from the margin at

172

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 6. Collecting records for species of Chlamydopsis strigicollis group.

the sides to meet the
circumcoxal stria (as it does in
the preceding species), but it is
conspicuously grooved, as in
many Chlamydopsis outside
of this group. While the
prostemal strial configuration
is likely a symplesiomorphy,
the form of the superficial
humeral groove of the
trichome (strongly oblique,
entering the trichome near its
mesal base) is a more unusual
character supporting their
placement in the strigicollis
group. The females of this
species and C. setifera are
particularly similar (though
only the female of the latter is
known), sharing smooth
elytra bearing elongate sparse
setae, and essentially identical
trichome shape. The principal
difference between them is
that C. setifera bears elongate isolated setae on
the propygidia and all ventrites as well as
dorsally. The pygidia and venter of C. monteithi
are glabrous. The males of C. monteithi and C.
lawrencei are generally similar, but the latter is
smaller (2.37mm vs 2.31mm), less densely
strigose, and has the humeral trichomes smaller
and slightly more distant from the sides.

DESCRIPTION. L: 2.37; W: 0.84; E/Pn L: 1.81;
E/Pn W: 1.31; Pn W/L: 1 .56; E LAV: 0.89; Pr/Py:
0.95; Sterna: 0.75, 0. 1 6, 0.72; Tibiae: 0.87, 0.87,
0.97. Body rufescent, bearing numerous
elongate, single setae; frons slightly wider than
long, sides strongly rounded, apical margin
weakly rounded; frontal disk flat, densely
reticulopunctate, bearing 1-2 elongate setae;
labrum subacute at apex, faintly punctate, with
short sparse setae at apex; antennal scapes bluntly
angulate near middle, narrowed to base and more
gradually to apex, densely reticulostrigose.

Pronotum about 1 .4 x as wide as long, sides
unmargined (though appearing submargined in
posterior half where carina delimiting proleg
depression intrudes), widest near base, slightly
sinuately narrowed to apex; anterior margin
weakly elevated above antennal cavities (more
distinctly so in male), medial portion shallowly
inwardly arcuate; pronotal disk slightly
depressed in anterior comers, convex posteriorly,
densely reticulostrigose, strigae converging to

scutellum, bearing a few (exactly 4 in most of
type series) elongate setae, some of them
‘bundles’ from conspicuous punctures.

Prostemum with anterior margin finely but
distinctly grooved, the groove only slightly
diverging from margin at sides, not joining
circumcoxal stria, but meeting the
pronotal/prostemal marginal groove near the
anterolateral pronotal corner; anterior prostemal
margin slightly deflexed, sinuate, rounded at
middle, without lateral projections: prostemal
keel narrowed posteriorly, very shallowly
emarginate at apex, with fine marginal stria along
inner edge of leg depression, but not along apical
margin of keel; prostemal disk fairly evenly
punctate, more shallowly so along medial portion
of anterior margin, glabrous.

Elytra about 1.3 x as wide as base of
pronotum; widest close to base, sides parallel in
basal two-thirds, rounded to apices; humeri
strongly elevated (more so in females), anterior
and posterior elevations divided by transverse
furrow, this furrow deeper and broader mesally
than laterally, densely setose on posterior and
especially anterior mesal edges; anterior
superficial humeral groove fine, oblique, nearly
horizontal, entering mesal base of trichome
opening; mediobasal depression occupying less
than basal half of elytra, with fine transverse
carinae from just behind scutellum curving
posterolaterally to middle of trichome opening;
dorsum of elytral disk of female smooth,

REVIEW OF CHLAMYDOPSIS

173

impunctate except for widely scattered
setigerous punctures, each with elongate seta
(few w'ith multiple setae), these subserially
arranged behind trichome and along posterior
half of elytra! suture; elytral disk of male finely,
longitudinally reticulostrigose in apical
two-thirds, with similarly sparse setigerous
punctures; epipleuron strigose in both sexes,
strigae converging to trichome; posterior elytral
margin not carinate; elytral marginal stria
continuous around lateral and posterior margins,
but entirely absent along suture.

Mesostemum about 3 x as wide as median
length, weakly projecting at middle, disk densely
punctate; mesometasternal suture finely but
crenulately impressed, continuous at sides with
postmesocoxal stria; median longitudinal
metastemal suture not impressed, but visible;
metasternal disk and 1st abdominal sternite
impunctate, glabrous; legs short, slender, with
short fine setae; profemur and tibia faintly
punctate, meso- and metatibiae and femora
impunctate; outer margin of protibia acutely
angulate at basal one-third, meso- and metatibiae
more bluntly angulate nearer their midpoints.

Propygidium convex, densely but very
shallowly reticulopunctate in male, reticulae
only barely visible in female; pygidium of male
faintly reticulopunctate in basal half, impunctate
apical ly, that of female impunctate; pygidia in
both sexes glabrous.

REMARKS. This species is named in honor of
Geoff Monteith, collector of this species’ entire
type series, whose field efforts have contributed
enormously to chlamydopsine studies.

Chlamydopsis setifera sp. nov.

(Figs 4G, 5F, 6)

MATERIAL. HOLOTYPE (QMT108579) 9: SEQ:
26°53’S 152°09’E, Benarkin School, 14 Nov 1994-26 Jan
1995, G.B. Monteith, Intercept trap, open forest.
PARATYPE 9: Qld: 27°33’S 153°28’E, N.Stradbroke
I.,Enterprise Blackbutt #1, 90m, 8-22 Jan 2002,QM party
pitfall trap 10343.

DESCRIPTION. L: 2.3 1 ; W: 0.8 1 ; E/Pn L: 1 .85;
E/Pn W: 1 .53; Pn W/L: 1 .46; E L/W: 0.83; Pr/Py:
0.95; Sterna: 0.69, 0.12, 0.72; Tibiae: 0.87, 0.81,
0.90. This species exhibits only a few differences
from the females of the C. monteithi , described
fully above. They differ primarily in distribution
of setae. The elongate setae of this species are
more evenly distributed on the body. The
pronotum bears setae along the anterior and
lateral margins, rather than just at middle; setae

of the elytra extend up the entire length of the
suture (rather than being confined to its apical
half); the pygidia, legs and ventrites all bear
elongate setae completely lacking in the
preceding species; pronotal reticulae faint at
middle, obsolete in apical half. There is an
additional possible difference in the form of the
trichome. On the left side of the type (but not the
right) the furrow between anterior and posterior
elevations is more deeply and finely incised than
in the preceding species. However, this may
result from ant damage, as this is where ants are
known to grasp the beetles in their mandibles.

REMARKS. Although males of this species are
not yet known, it is likely that the species is
dimorphic, and that the males will have the elytra
densely reticulostrigose, as in the preceding
species.

Chlamydopsis lawrencei sp. nov.

(Figs 4H, 5G, 6)

MATERIAL. HOLOTYPE 6 : 35. 16S 149.06S [sic] ACT,
Black Mtn. Canberra, 25 Oct. 1990, J.F.Lawrcnce,
sweeping; in ANIC. PARATYPE: 1 6 : ACT: 36.16S
149.05 ACT Black Mtn.W. slope,5.xii.l989, T. Weir,
Berlesate ANIC 1 125, open forest liner; in ANIC.

DIAGNOSIS. As above, this species is unusual
in having the anterior prostemal margin grooved
but with the groove not diverging at the sides to
meet the circumcoxal stria. This species, known
only from the two males, is likely sexually
dimorphic like the preceding two. This male
differs from the male of C. monteithi in its smaller
body size, shorter, more convex pronotum, and
smaller, more narrowly separated humeral
trichomes.

DESCRIPTION. L: 2.3 1 ; W: 0.8 1 ; E/Pn L: 1 .85;
E/Pn W: 1 .43; Pn W/L: 1 .42; E L/W: 0.9 1 ; Pr/Py:
0.80; Sterna: 0.69, 0.12, 0.72; Tibiae: 0.84, 0.87,
1.00. Body dark rufescent; frons about 1.2 X as
wide as long, sides strongly rounded, margins
elevated, mostly Hat, faintly depressed between
antennal bases; frontal disk reticulopunctate,
with a few fine setae; labrum rounded, shallowly
punctate; outer margin of antennal scapes bluntly
angulate near midpoint, baso- and apicolateral
margins straight, apex rounded; antennal club
about 1 .4 x as long as antennal scape.

Prothorax about 1.3 x as wide as median
length, approximately equal in width basally and
apically, slightly narrowed at middle,
unmargined in anterior half, outer edge of
prosternal leg depression projecting beyond

174

MEMOIRS OF THE QUEENSLAND MUSEUM

margin in posterior half (such that it appears
margined from above); lateral thirds of anterior
margin strongly arcuate, elevated; median
portion only very weakly elevated; pronotal disk
shallowly depressed in anterior comers, other-
wise convex, with subacute median tubercle.

Prosternum with anterior margin linely
grooved, the groove only slightly diverging from
margin at sides, not joining circumcoxal stria, but
meeting the pronotal/prostemal marginal grove
near the anterolateral pronotal comer; anterior
margin slightly deflexed, uneven, not projecting
at sides; prostemal keel narrowed posteriorly,
shallowly emarginate at apex; prostemal disk
reticulopunctate, glabrous.

Elytra about 1.3 x as wide as base of pro-
notum, sides nearly straight, slightly convergent
to front; humeri moderately elevated, not incised
laterally, trichome a mesally open semicircle,
with an uneven inner setal fringe, the posteriorly
directed setae of the anterior edge overlaying
those of the posterior edge; anterior superficial
humeral groove fine, oblique, extending from the
humeral comer medially along the anterior
elytral margin for a short distance before curv ing
back to enter the anteromesal base of the
trichome opening; mediobasal depression short,
confined to basal one-third, with low, obliquely
transverse basal carinae; apical elytral margin
weakly carinate; elytral disk reticulostrigose;
with scattered elongate setae throughout.

Mesostemum about 4x as wide as median
length, weakly projecting anteriorly, disk densely
punctate; mesometastemal suture finely im-
pressed, continuous at sides with postmesocoxal
stria; median longitudinal metastemal suture
faintly impressed; metastemal disk with few fine
punctures, glabrous, with faint alutaceous
microsculpture; 1st abdominal stemite with row
of fine punctures along anterior margin, with a
few scattered punctures behind; legs short,
slender, minutely setose; outer surfaces of
profemur and tibia faintly punctate, meso- and
metatibiae and femora impunctate; outer margin
of protibia acutely angulate at basal one-third,
meso- and metatibiae more bluntly angulate.

Prppygidium shallowly reticulopunctate, with
sparse elongate setae; pygidium very shallowly
reticulopunctate in basal one-third, nearly
smooth apical ly.

REMARKS. The female of this species is not
known. Discovering that it shared the dimorph-
ism in elytral texture with C. monteithi , above,
would support its position in this group. The

species is named in honor its type’s collector,
coleopterist par excellence John Lawrence.

PYGIDIALIS GROUP

While a few of the species in this group are
extremely distinctive and closely related, it is in
fact a rather varied group, with only a few species
showing the radical serrate and tuberculate
surfaces seen in the nominate species. All of the
species possess some form of frontal tubercles,
usually in two longitudinal rows of three each,
although 2 and 4 tubercles/row have also been
seen. The humeral trichome is very consistent in
shape (with the exception of C setipennis ; see
below), forming a horizontal C-shaped arc
originating nearly at the humeral comer. Its setal
fringe (again with the exception of G setipennis )
extends along this arc, and is either continuous
through the lateral notch (the posteriormost point
of this arc) and along the anterior edge of the
posterior trichome elevation, or is interrupted at
the lateral notch. The anterior superficial humeral
groove in these species is also continuous with
this arc, extending from anteromesal base of the
trichome to the humeral comer. Most species also
exhibit the following characters: transverse
propygidial carina, median pronotal tubercle
present, often forming short transverse ridge
(often also with longitudinal carina leading to
anterior pronotal margin), lateral pronotal
margins elevated, metatibiae often expanded.
There is little question that the species included
here are closely related. However, some
characters suggest derivation of the longipes
group from among them.

Chlamvdopsis pvgidialis Blackburn, 1900
(Figs 41, 5H, 9)

Chlamvdopsis pvgidialis Blackburn, 1900: 206; Type:

Australia; NHM, examined, 2000.

Chlamydopsis pvgidialis var. minor Oke, 1923: 153; Mazur,
1984; Lcctotype, hereby designated: Femtrce Gully,
16.5.1920, C.Oke, Vic./ Chlamydopsis pygidialis var.
minor Type/ Presented by C. Oke/ 891 Type; MVM,
examined, 2000.

RECORDS. NHM: Vic: nr. Melbourne, vi.1920; ACT:
Canberra, Black Mt, vii.1974, under stones. WAM: Vic:
Femtrec Gully. AMS: Vic: Millgrave. Lea (1925): Vic: nr
Melbourne, May, with Iridomyrmex. Oke (1923):
Beaconsfield; Bclgrave; Healesville; Warburton; Yarra
Junction; Emerald; Pakenham [all Vic].

DIAGNOSIS. Chlamydopsis pygidialis shares
many characteristics with the following 3
species. All have the lateral and anterior pronotal
margins strongly elevated, and the posterior
margin of each elytron strongly carinate, with the

REVIEW OF CHLAMYDOPSIS

175

carina extending forward along the lateral
margins (extending nearly to the elytral base in C.
pygidialis , C. serricollis, and C. carinicollis).
These last three also have a strong median longi-
tudinal carina from the median pronotal tubercle
extending forward to the margin. Of the four
species, all but C. carinicollis have this carina at
least slightly to strongly serrate. Chlamydopsis
setipennis is easily separated from the remaining
three by the humeral trichomes in which the
anterior elevation is nearly or fully vertical, and
separated from the posterior elevation by a broad
notch lined by a long, unkempt setal fringe.
Chlamydopsis pygidialis and C. serricollis are
the most similar of the group, having pronotum,
elytra and propygidium strongly serrately margined.
The most distinctive character separating the two
is the oblique apical margin of the metatibia,
which in pygidialis departs about 75° from the
long axis of the tibia. This angle is closer to
45-50° in C. serricollis. The metatibia is less
strongly narrowed towards the base in C.
pygidialis. Lea (1914b) cited the medial longi-
tudinal carina of the pronotum as being more
conspicuously serrate in C. serricollis than in C.
pygidialis.

REMARKS. Blackburn (1891) reported that the
type was ‘obtained by beating dead branches and
probably connected with some species of
Hymenoptera inhabiting the dead wood’.

Chlamydopsis carinicollis Lea, 1919
(Figs 7A, 8A, 9)

Chlamydopsis carinicollis Lea, 1919: 178; Type locality:

Vic: Beaconsfield; SAM, not examined.

RECORDS. NHM: Vic, (Del. A. Lea); Oke (1923): Vic:
Femtree Gully; Vic: Upwey, Chalcoponera sp.

DIAGNOSIS. As above, this species shares the
carinate apical elytral margin and the median
longitudinal pronotal carina with C. pygidialis
and C. serricollis. However, in C. carinicollis
these carinae arc not serrate as they are in the
other two. The humeral trichome of this species is
also relatively small, and does not incise the
lateral aspect of the humeri.

REMARKS. The type was reported from a nest
of the ant Aphaenogaster longiceps in July (Lea,
1919). Of currently recognised subspecies, only
A. /. longiceps (Smith) occurs near the type
locality of C. carinicollis.

Chlamydopsis serricollis Lea, 1914
(Figs 7B, 8B, 9)

Chlamydopsis serricollis Lea, 1914b: 217; Type: serricollis
Type Lea, Pt. Hacking/ Chlamydopsis serricollis Lea type,
N. S. Wales; mounted with 1 ant, reportedly Ectatomma
metallician (Lea, 1914b); in SAM; examined.
Chlamydopsis serricomis; Mazur, 1997: 4; misspelling.

RECORDS. NHM: NSW: Sydney. Lea (1925): NSW: nr
Sydney, Ectatomma metallicum.

DIAGNOSIS. See the diagnosis under C. pygidialis
above.

Chlamydopsis setipennis Oke, 1923
(Figs 7C, 8C, 9)

Chlamydopsis setipennis Oke, 1923: 154; Lectotypc, hereby
designated: Evelyn. V., 5.6.22, C.Oke/ Chlamydopsis
setipennis Oke Type/ Presented by C.Oke/ 893 Type,
MVM, seen, 2000; Paralectotypes; Belgrave, Vic.,
16.10.21, C.Oke/ Chlamydopsis setipennis Oke Co-Type/
2149 Paratype/ F.E. Wilson Collection, MV; Belgrave,
Vic., 3.7.21, C. Oke/ Chlamydopsis setipennis Oke
Co-Type/ Presented by C. Oke/ 894 Paratype, MVM;
Belgrave, Vic., C. Oke, AMS; Femtree Gully, Vic.. C.
Oke, NHM; Victoria: Emerald, Sep 26, 1920, Oke, NHM.

DIAGNOSIS. Chlamydopsis setipennis is quite
distinctive within the pygidialis group. The form
of its humeral trichome is unique, having the
anterior and posterior elevations quite broadly
separated, with a large lateral notch between
them. The setal fringe of each is also unique, with
the setae of the anterior elevation being elongate,
rather sparse, and nearly erect, while those of the
posterior elevation, also unusually elongate, are
clustered into two distinct, adjacent ‘clumps’.

REMARKS. Described as inhabiting the nests of
Notorious foreli Andre var. dentatus Forel (Oke,
1923). This species is known only from the type
series.

Chlamydopsis convergens sp. nov.

(Figs 7D, 8D, 9)

MATERIAL. HOLOTYPE (QMT 108580) 8: NEQ:
1 7^26’S, 145°42’E, Hughes Rd, Topaz. 6 Dec 1 993-25 Feb
1994, Monteith, Cook, Janetzki, RF Intercept, 650m.
PARATYPES (11): 8 ex.: same data as holotype; 1 ex.:
NEQ: 17°14 ? S 145 D 25‘E, 3km W of Bones Knob,
10 xii.I995-9.ii.l996, GM,DC,GT, RF PT, 1140m; 1 ex.:
Qld: 1 7.37S, 145.34E 1000m, BS3 Massey Ck.,
1 xii.l994-3.i.l995, PZ, FIT JCU (West); 1 ex.: Qld:
17.28S 145.29E, BS1 Longlands Gap, 2.x-l.xi.l995, LU,
1 150m, FIT JCU, in QMB, DPIM, ANIC, MSCC.

DIAGNOSIS. The surface texture is unique among
members of the pygidialis group. Both sexes
have shiny, only very shallowly punctatostrigose
elytra. In the females, apart from the humeri and
epipleurae, the elytra may be entirely impunctate.

176

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 7. Dorsal views of Chlamydopsis spp. A, C. carinicollis. B, C. serricollis. C, C. setipennis. D, C. convergens.
E, C. erupta. F, C. transversa. G, C. longipes. H, C. inaequalis. I, C. agilis.

The males typically have sparse shallow although it is more variable in this regard than the
punctures on the dorsal elytral apices as well. The elytra. An additional, almost unique feature is a
pronotum may also be largely impunctate, secondary erect fringe of setae above the

REVIEW OF CHLAMYDOPSIS

177

decumbent fringe along the anterior inner edge of
the trichome. This is seen in one of two
individuals of C. coronis as well.

DESCRIPTION. L: 1.87; W: 0.69; E/Pn L: 1.73;
E/Pn W: 1 . 1 0; Pn W/L: 1 .77; E LAV: 0.88; Pr/Py:
1.14; Sterna: 0.59, 0. 1 6, 0.53; Tibiae: 0.69, 0.75,
0.90. Body nearly black, faintly rufescent
(especially extremities) parallel-sided. Frons as
wide as long, sides rounded, disk with 6
prominent tubercles arranged in two vertical
rows, the middle pair most prominent, the upper
pair least, the anterior pair on the apical margin;
frontal disk with slightly irregularly spaced
rounded punctures; labrum rounded, with a few
small punctures, about 6 inconspicuous setae on
apical margin; antennal scapes arcuate, with
outer margin somwhat undulating, obliquely
truncate apically, anterior surface convex,
subcarinate along the longitudinal axis.

Pronotum margined laterally, convex, with a
small acute tubercle medially, concave behind
upturned lateral and anterior margins; lateral
margins outwardly arcuate and widening slightly
towards the front, continuous with inwardly
arcuate, oblique anterior margins; middle third of
anterior margin somewhat separated from lateral
oblique portions, its edge slightly interrupted at
middle; median pronotal elevation extended
anteriorly as a faint carina; pronotal disk densely
puncate in anterior and lateral depressions, less
so on medial convex area; posterior edge of
pronotal disk with a fine, distinct marginal stria.

Prosternum with anterior margin faintly
sinuate, not grooved, transversely depressed
behind anterior margin, rising slightly to
narrowed posterior apex of prosternal keel,
which is faintly tuberculate at middle, acutely
emarginate apically, with circumcoxal stria
carinate; prosternal disk uniformly punctate,
with strong microsculpture between punctures.

Mesostemum about 2 x as wide as long,
acutely projecting at middle, with complete
anterior marginal stria, punctate as prostemum,
raised slightly above anterior margin of
metasternum; metasternal disk with coarse
punctures along anterior margin, more finely but
uniformly punctate elsewhere; abdominal
stemite 1 similarly punctate anteriorly, but more
finely behind.

Elytra with humeral trichomes well developed,
strongly depressed between, coarsely rugose in
anterior (except along elytral suture), smoother,
with shallow punctures posteriorly (and
extending faintly anteriorly along suture) in

male, more or less impunctate in most females;
trichome basically a transverse incision between
anterior and posterior elevations, with dense setal
fringe extending inwardly and ventral ly from
apex, curving anteriorly along inner basal edge
where it meets the inconspicuous, horizontal
anterior superficial groove; each elytron with a
poorly developed transverse carina in outer half,
about one-third from apex; posterolateral
marginal carina from elytral apex (well
developed in, e.g., C. transversa sp. nov.) faint,
confined to apical corner; epipleurae with
shallow striae converging to apex of trichome.

Outer surfaces of protibia and femur uniformly
covered with shallow punctures, those of meso-
and metathoracic legs with only very fine
punctures: protibiae slender, angulate at middle
of outer edge, with margins arcuate to angle;
meso- and metatibiae much broader, though still
more or less angulate, their inner edges with
discrete band of microsculpture where they
overlap the inner edge of femora; the tarsal
grooves of meso- and metatibiae parallel to the
apical half of lateral margin (rather than parallel
to the longitudinal axis of the tibia),

Propygidium depressed along basal margin,
but strongly transversely carinate along apical
margin, this carina weakly interrupted at middle,
with an additional median tubercle just anterior to
the carina, uniformly punctate with punctures
separated by slightly less than their widths;
pygidium weakly convex at middle, flat to weakly
depressed along apical margins, punctures
smaller and more widely separated than those of
propygidium, almost uniformly distributed, only
slightly denser in anterior comers.

REMARKS. A single male specimen from Qld:
2km SE Mt Spurgeon (QMB) has a particularly
well developed median pronotal tubercle, and
conspicuous posterolateral pronotal tubercles,
which are only faintly detectable in the type
series of C. convergent Its pronotal punctation is
also distinctly coarser. This specimen is excluded
from the type series. However, a subspecies
designation might be appropriate if additional
material from this area shares these differences.
The name of this species refers to the con-
vergently strigose sculpturing of its epipleurae.

Chlamydopsis coronis sp. nov.

(Fig. 9)

MATERIAL. HOLOTYPE (QMT108581) 9: 3km N.
Lansdowne via Taree, N.S.W., 10 Feb. 1988, G Williams,
intercept trap; ex rainforest-wet scler. forest margin.

178

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. Lateral views of Chlamydopsis spp. A, C.
carinicoUis. B, C. serricollis. C, C. setipennis. D, C.
converge ns. E, C. <?rw/?/a. F, C. transversa. G, C.
longipes. H, C. inaequalis. I, C. ag/7/.s.

REVIEW OF CHLAMYDOPSIS

179

DESCRIPTION. L: 2. 1 8; W: 0.78; E/Pn L: 1 .80;
E/Pn W: 1 .30; Pn W/L: 1 .60; E LAV: 0.87; Pr/Py:
1.13; Sterna: 0.65, 0.22, 0.62; Tibiae: 0.69, 0.84,
1.00. This species is extremely similar, and
closely related, to the preceding species. Only the
differences are described here. Pronotum with
transverse row of 4 median tubercles, with a
carina running from the base of the middle two to
the anterior marginal ridge, with which it merges;
with a pair of posterolateral tubercles on each
side, and with a low but distinct pair of
posteromedial tubercles immediately in front of
scutellum; lateral pronotal margin slightly
undulating, widening slightly towards front,
entire pronotal disk densely and uniformly
punctate. Elytra with trichome similar in shape,
posterior transverse carina of elytron developed
as a row of three tubercles, the middle the most
prominent; row of tubercles present from elytral
apex anteriorly along elytral suture, diminishing
in prominence towards the front, detectable to
about the elytral midpoint; apical margin of
elytron carinate and prominent, particularly in
about the middle third. Punctures of posterior
portion of metastemum slightly larger than in C.
convergent. Propygidium with apical transverse
carina interrupted to form row of 5 tubercles,
with an additional medial tubercle anterior to this
row; pygidium with faintly developed medial
tubercle at basal one-third.

REMARKS. A second specimen, probably of
this species (ACT: Blundells Ck, 3km E of
Piccadilly Circus; AN1C), is not included in the
type series due to several differences. Most
obviously the pronotal tubercles are much less
prominent and the secondary, erect row r of
trichome setae is not present. Additional material
from these and intervening (they are separated by
about 700km) localities needs to be studied to
determine the significance of these differences.
The name of this species refers to the crown-like
appearance of the anterior pronotal margin.

Chlamydopsis erupta sp. nov.

(Figs 7E, 8E, 9)

MATERIAL. HOLOTYPE (QMT1 08582): NEQ:
1 7°26'S, 145°42’E, Hughes Road, Topaz, 6 Dec 1993-25
Feb 1994, Monteith, Cook, Janetzki, RF Intercept, 650m.
PARATYPES (21): 4 ex.: same data as holotype; 1 ex:
NEQ, 19km NE of Mareeba, 20.xii. 1984-73.1985, RS &
Titmarsh/MDPI Site 26; 2 ex.: NEQ, Windsor Tableland.
28.ii-6.iii. 1992, JH, FIT: 1 ex.: Qld, 17.28S 145.29E, BS1
Longlands Gap, 30.xi.l 995-33.1 996, LU, 1150m, FIT
JCU; 1 ex.: NEQ, Hugh Nelson Range, 21kmS Atherton,
l.xii. 1983-93. 1984, RS & JB/MDPI Site 17; 1 ex.: NEQ,

Millaa Millaa Falls, 1 3 .iii- 1 03 v. 1 990,RS & KH, MDPI FIT
Site 34; 2 ex.: same but 73i-13.iii.1990; 1 ex.: same but
4.v-6.vi. 1 990; 2 ex.: NEQ, Wongabel SF, 6km S Atherton,
26.vii-3.ix.1984, RS & JB/MDPI Site 18; 1 ex.: same but
3.ix-l.xi.l984; 2 ex.: NEQ, Charmillin Ck Xing, 950m,
Tully Falls Rd. 8.xii. 1989-53. 1990. GM,GT,HJ, PT&FIT;

1 ex.: Qld, 17.06S 145.37E, Mt Edith GS2,1050m,
17.iii-6.iv.l995,PZ, FIT; 1 ex.: NEQ, 6km S Kuranda,
15.i-203i.1985, RS & KH, MDPI Site 22; 1 ex.: Qld,
17.33S 145.32E, BS2, Mt Fisher, 1150m, 3.x-2.xi.l995,
LU, MT, in QMB, DPIM, ANIC, MSCC.

DIAGNOSIS. While this species shows a couple
of obvious similarities to the preceding, in par-
ticular the transverse row of 4 median pronotal
tubercles and the row of tubercles along the
posterior half of the elytral suture, it is otherwise
one of the most distinctive of all Chlamydopsis ,
with the entire dorsal surface tuberculate. Most
tubercles on the elytra, especially those on the
anterior surface of the humeral trichome, are
without obvious homologues in any other species.

DESCRIPTION. L: 1.81; W: 0.69; E/Pn L: 1.64;
E/Pn W: 1.35; Pn W/L: 1.41; EL/W: 0.86; Pr/Py:
1.08; Sterna: 0.50, 0.12, 0.44; Tibiae: 0.56, 0.65,
0.75. Body dark rufescent in color, with
conspicuous tubercles on most dorsal surfaces,
more or less reticulately punctate throughout.
Frons slightly longer than wide, sides (below
antennal insertions) nearly straight, slightly
convergent to apex, surface with two longitudinal
rows of 3 tubercles each (in most individuals;
specimens with only 2 or as many as 4 tubercles
in each row' have been seen), the anteriormost
pair at the apical margin, the uppermost row
between the antennal insertions; frontal surface
otherwise reticulately punctate; clypeolabral
suture slightly incurved; labrum rounded,
shallowly punctate; antennal scape arcuate,
widest at middle, outer edge undulating, surface
texture as on frons with from 2-3 tubercles in a
longitudinal row; antennal club of male slightly
longer than scape, that of female about 2/3 as long
as scape.

Pronotum about twice as wide as long, sides
margined, more or less inwardly arcuate,
interrupted by subbasal and, usually, subapical
tubercles; anterior margin strongly upturned,
with lateral supra-antennal portions arcuate and
separated from medial portion, the medial
portion sinuate, often forming a continuous ‘Y’
with a longitudinal carina in the anterior third of
disk, some individuals with anterior marginal
carina isolated and only a medial anterior tubercle;
medial portion of disk with numerous tubercles,
somewhat varied in specific arrangement,

180

MEMOIRS OF THE QUEENSLAND MUSEUM

O C. pygidialis
□ C. carinicollis
A C. serricollis
* C. setipennis

usually with a transverse
arcuate row of 4 at middle, a
pair on each side
posterolateral to this row, and
a prominent pair in front of the
scutellum, in addition to those
along the lateral margins;
often also tubercles between
scutellar pair and transverse
median row; ground texture
uniformly densely punctate;
posterior margin of pronotum
with fine marginal stria, this
interrupted at a small
prescutellar emargination.

Anterior margin of
prosternum simple, middle
third weakly outwardly
arcuate, prosternal disk
transversely depressed behind
anterior margin, an isolated
transverse stria visible in
some individuals in this
depression, prosternal keel
with tubercle at middle, narrowed posteriorly,
acutely emarginate at apex, with circumcoxal
stria carinate, separating smooth leg cavities
from reticulate prosternal disk.

Mesostemum slightly more than 2 x as wide as
long, acutely projecting anteriorly, without an
obvious marginal stria, subcarinate along midline
and raised in posterolateral comers, elsewhere
depressed, coarsely punctured and micro-
sculptured throughout; mesometastemal suture
finely but distinctly impressed; metastemal disk
coarsely punctured and microsculptured
throughout, but punctures smaller and sparser at
middle; 1st visible abdominal stemite with a
continuous row of punctures along anterior
margin, elsewhere more sparsely punctate, with
conspicuous microsculpture throughout.

Elytra parallel sided, rather abruptly narrowed
at base and apex, tuberculate throughout, most
consistently along elytral suture (the anteriormost
of these less prominent), at sides immediately
behind trichomes, and on the dorsum of the
anterior elevation of the trichome; humeral
trichome well developed, though not as strongly
elevated as in most species of this group, forming
a sinuate transverse groove, widening at middle,
curved anteriorly to base of elytron, continued
anterolaterally by a fine, oblique anterior
superficial groove; trichome densely lined on
inner edges by inclinate golden setae which

C. convergens
C. coronis
C. erupta
C. transversa

FIG 9. Collecting records for species of Chlamydopsis pygidialis group.

completely obscure its opening; surface of elytra
mostly reticulately punctate, with reticulae more
elongate posteromedial to trichomes; posterior
marginal carina well developed, and undulating,
particularly at the centre, extending forward at
side (where it diverges from the elytral margin)
approximately one-fifth of the distance from
apex.

Legs as in C. convergens , except outer surfaces
of meso- and metathoracic legs slightly more
densely punctate, though still less so than
prothoracic legs.

Propygidium depressed along basal margin,
elevated as a transverse row of tubercles along
apical margin, with prominent medial, and less
prominent lateral, tubercles in front of this row;
pygidium slightly convex, with three prominent
tubercles in an arcuate row near base, and with
slightly elongate tubercles at middle of each
lateral margin; propygidium and pygidium
densely punctate throughout.

REMARKS: A single specimen from near
Lansdowne (NSW: Lorien, 4. i. 1987; CMN) is
excluded from the type series. This locality is
over 1 000km from the next nearest site for the
species. Although its dorsal tubercles clearly
associate it with this species, differences in
metastemal punctation and trichome shape leave
its specific identity uncertain. The name of this
species refers to the numerous tubercles on its
body.

REVIEW OF CHLAMYDOPSIS

181

Chlamydopsis transversa sp. nov.

(Figs 7F, 8F, 9)

MATERIAL. HOLOTYPE (QMT108582) 8 :
AUSTRALIA: N Qld, Hann Tableland, 13km WNW of
Marecba, 9.xi-7.xii.l988, Storey & Dickinson/ MDPI
Intercept Trap Site No. 3 1 . PARATYPES (22): 3 ex.: same
data as holotype; 3 ex.: same but 17.ii-20.iii. 1989; 1 ex.:
same but 13.x-9.xi.l988;2ex.: same but 1 .iii-12.iv.1994; 2
ex.: same but 7.xii. 1 988- 17.i. 1989; 2 ex.: same but
13.i-l.tii. 1994; 1 ex.: same but 13.vii-4.viii.1988; 1 ex.;
same but 4.viii-9.ix. 1 988; 1 ex.: same but
20.vi-13.vii.1988; 1 ex.: SEQ: 26°16'S 151°25’E, Roy
Property at Brigooda (Top site), 26.i-20.iv. 1995, GM, FIT,
vine scrub; 1 ex.: SEQ: 25°3TS I52°18'E, Fairlies Knob,
0.5km S, 2 1 .vii-20.x.2000, 300m, DC, Wright Vanderduys,
vine scrub PT 9464; 1 ex.: SEQ: 25°08'S I51°59’E,
Nangur SF, 2nd site, 24.xi.1995-3.ii. 1996, GM, FIT, RF.
320m, 5853; 1 ex.: SEQ: 27°14'S 152°I5’E, Mt
Dcongwar, 3km S, site 2, 30.xii.98-26.iii.1999, GM, RF
PT, 460m, 7652; 1 ex.: SEQ: 25°40’S 151°26’E, Nipping
Gully, site 2, 2 1 . viii-9.x. 1 998, GM. RF FIT, 200m, 7258; 1
ex.: Qld, 23°37’S 150°2S’E, Mt Gavial, 3km SSW,
18.xii.98-14.iii.1999, EXT, vine forest FIT, 320m, 7492, in
QMB, DPIM. ANIC, MSCC.

DIAGNOSIS. This species is easily recognised
by the combination of well developed transverse
medial and posterolateral pronotal carinae, well
developed subapical elytral carinae (in lateral
half of each elytron), absent or at most weak
median longitudinal pronotal carina, slender
metatibia acutely angulate at its midpoint, and
elytra densely reticulopunctate (except in
mediobasal depression) in both sexes.

DESCRIPTION. L: 1.90; W: 0.69; E/Pn L: 1.77;
E/Pn W: 1 .26; Pn W/L: 1 .59; E L/W: 0.89; Pr/Py:
1.14; Sterna: 0.56, 0.16, 0.47; Tibiae: 0.75, 0.78,
0.97. Dark rufescent brown, elongate, parallel-
sided, near entire dorsum densely reticulately
punctured. Frons nearly as wide as long, sides
rounded, disk deeply reticulately punctate, with 6
prominent, more or less acute, tubercles arranged
in two vertical rows of 3, the middle pair the most
prominent; frontal punctures as elsewhere (e.g.,
antennal scapes, pronotum); labrum arcuate,
semicircular, with two long and several short
setae; antennal scapes with outer edges
irregularly arcuate, widest at middle, anterior
surface convex and reticulately punctured;
antennal club of male very elongate,
approximately 1.25 x as long as scape.

Pronotum about 1.3 x as wide as long,
margined laterally, sides more or less inwardly
arcuate, but interrupted at basal one-third by a
low acute tubercle; anterior pronotal margin strongly
upturned, obliquely arcuate above antennal

cavities, these edges discontinuous from medial
portion of margin, which is bilobate, the two
halves meeting in a very shallow 4 V’; pronotal
disk with a strong transverse carina in medial
one-third, rising about equally high as the
upturned anterior pronotal margin, the dorsal
edge of this carina weakly sinuate, and with
additional lateral tubercles immediately
posterolaterally, these adjacent to, but not con-
tinuous with those along the lateral pronotal
margins; pronotal disk also with relatively small
tubercles along posterior margin, immediately in
front of and on each side of the scutellum.

Prostemum with anterior margin faintly sinuate,
not grooved, transversely depressed behind
anterior margin, rising slightly to narrowed
posterior apex of prostemal keel; prostemal keel
acutely emarginate apically, with circumcoxal
stria carinate; prostemal disk densely punctate.

Elytra parallel-sided, with prominent humeral
trichomes, the anterior elevations of which are
strongly convex, with complete marginal fringe
of conspicuous golden setae from inner base to
apex, apparently continuous, through a shallow
lateral groove, with the anteriorly directed setae
of the apex of the posterior elevation; anterior
superficial groove inconspicuous but entering the
trichome horizontally at the inner base; each
elytron with an isolated transverse carina in lateral
half, one-fourth from the apex, and a lateral
longitudinal carina, continuous with the posterior
elytral margin, delimiting the epipleuron in the
apical half elytral disks reticulately strigose
basally and laterally (especially on epipleuron),
becoming more finely reticulate at middle and
towards apex.

Mesostemum acutely projecting at middle,
slightly raised at middle and along posterior
margin, with complete anterior marginal stria,
densely punctate and microsculptured;
metasternal disk more finely and sparsely
punctate than mesostemum, especially laterally;
visible abdominal stemite 1 densely punctate
along basal margin and at sides, less densely
posteriorly.

Prothoracic legs slender, reticulately punctate,
the protibia acutely angulate one-third from base;
meso- and metathoracic legs slightly more
elongate, and less conspicuously punctured,
although with microsculpture, both posterior
tibiae similarly angulate to protibia.

Propygidium depressed along basal margin,
but strongly transversely carinate along apical
margin, this carina weakly interrupted at middle,

182

MEMOIRS OF THE QUEENSLAND MUSEUM

with an additional median tubercle anterad;
pygidium weakly convex at middle, weakly
depressed along apical margins, and with a
median tubercle one-third from basal margin;
both propygidium and pygidium uniformly
reticulately punctate.

REMARKS. The name of this species refers to
the transverse pronotal carina.

LONGIPES GROUP

The longipes group is clearly monophyletic,
with three known species. In addition to the
extremely elongate metatibiae, the species all
share: prominent pyramidal frontal protuberance,
elongate arcuate labrum, curved humeral trichome
with the anterior superficial stria entering at the
mesal base, and clavate metafemora. Of these, all
are probable synapomorphies except for the
trichome character, which resembles the possibly
related pygidialis and striatipennis groups.

Chlamvdopsis longipes Lea, 1910
(Figs 7G, 8G, 12)

Chlamydopsis longipes Lea, 1910: 201; Type locality: Vic:
Bannockburn; repository: SAM; not examined.

RECORDS. SAM: SA: Katarapko Game Res. 34°24’S,
140°34’E, pitfalls, 9-26 May, 1991. NHM: Vic: Melton,
x.1920. WAM: Vic: Melton. Lea (1912): Vic: Ararat Oke
(1923): Vic: Melton; Vic: Bacchus Marsh.

DIAGNOSIS. Despite sharing the basic trichome
plan with the other two species of this group, its
actual shape in C. longipes is highly distinctive.
The inner edge is deeply and broadly emarginate,
the humerus consequently appearing as a thin,
inwardly open arch. This species also has the
most elongate meso- and metatibae of the three,
as well as a generally more elongate body,
particularly noticeable in the form of the
pronotum.

Chlamvdopsis inaequalis Blackburn, 1891
(Figs 7H, 8H, 12, 29C)

Chlamydopsis inaequalis Blackburn, 1891: 94; Type:
Australia, NHM; examined, 2000.

RECORDS. ANIC: ACT: 35.17S 149.07E, Black Mt,
4.i. 1 989, light trap (1); 4 others, same locality, all from light
traps, Jan. (3), Feb. (1).

DIAGNOSIS. L: 2.93; W: 1.06; E/Pn L: 1.76;
E/Pn W: 1 .3 1 ; Pn W/L: 1 .53; E L/W: 0.88; Pr/Pv:
1.10; Sterna: 1.00,0.19, 1.12; Tibiae: 1.25, 1.62,
2.49. This species and C. agilis are very similar in
body form, both being much more quadrate than
C. longipes, and their trichome shape is

essentially identical. The two may be separated
by the form of the metatibia, which in C.
inaequalis is much more slender, with the
subapical angulation about one-third from the
apex. In C. agilis the angulation is less
prominent, and nearer one-sixth from the apex.

Chlamvdopsis agilis Lea, 1914
(Figs 71, 81, 12)

Chlamydopsis agilis Lea, 1914b: 216; Type locality: NSW:

Sydney; repository: SAM; not examined.

RECORDS. NHM: NSW: Sydney (Det. by Lea). Lea
(1919): SA: Naime, nest of Ectatomma metallicum. Lea
(1925): NSW: Como; NSW: Lane Cove.

DIAGNOSIS. See above under C. inaequalis.

REMARKS. The type was collected from a nest
oi Ectatomma metallicum (Lea, 1914b).

ECTATOMMAE GROUP

The C. ectatommae group is primarily defined
by a short anterolateral pronotal groove that
originates along the upper edge of the antennal
cavity, extends posterolaterally, incising the
lateral portion of the anterior pronotal margin,
curves around the lateral pronotal margin (which
in most of these species is poorly defined), and
then merges with the lateralmost extent of the
circumcoxal stria. In all but a couple of species
this groove is very distinct. Where it appears only
weakly impressed (C. pecki sp. nov., C. malice
sp. nov.), additional similarities to other species
leave little doubt as to their placement. The
bifovaecoUis subgroup is placed here with some
uncertainty. These arc unique in having a
pronotal groove, originating at the same point
behind the antennal cavities, that extends straight
backward and ends in a conspicuous dorsolateral
pit (Fig. 29A) In addition to this putatively
homologous groove, the trichome shape is quite
similar to that of several other unquestioned
members of the ectatommae group. That said,
trichome shape is rather varied across the entire
ectatommae group. In most species the anterior
and posterior elevations converge to a single
apex, are uninterrupted laterally, and have a
mesal emargination fringed with a simple arc of
setae, concealing the trichome opening beneath.
The anterior superficial groove of the trichome
usually extends somewhat inwardly and
obliquely from the humeral corner to the
anteromesal origin of the setal fringe. The major
variables are the height of the trichome and the
size of the mesal emargination (and consequently
the extent of the setal fringe.) One extreme is

REVIEW OF CHLAMYDOPSIS

183

found in G loculosa , in which the trichome
constitutes no more than a small humeral pit at
the apex of the anterior superficial groove.
Unlike species in the preceding groups, but like
most of those that follow, the ectatommae group
species have the anterior prosternal margin
deeply grooved, with the groove diverging from
the margin laterally to meet the circumcoxal stria.

This is a large group, containing more than 20
known species, ranging across temperate and
tropical Australia, and with two members occurring
in New Guinea (C. papuae Lewis and C.
jayawijaya sp. nov.).

Chla mvdopsis ectatommae Lea, 1912
(Figs 10 A, 11 A, 12)

Chlamydopsis ectatommae Lea, 1912: 66; Lectotype. hereby
designated: ectatommae Lea Type, Gladesville / cotype
[on underside of same label]/ 14671, Chlamydopsis
ectatommae Lea N. S. Wales, Type; SAM, examined
2002. It is uncertain whether syntypes of this species exist
in other collections, as the ‘cotype’ label would imply.
The ‘cotype’ written on the bottom of the main label is
partially cut off, and may not have been intended to
pertain to this specimen.

Chlamydopsis ectatommae var. rufomaculatus Oke, 1923:
153; Mazur, 1984; Type: Victoria: Melton; Bacchus
Marsh, in nests of Chalcoponera metallica ; SAM, not
examined.

RECORDS. NHM: NSW: Como; NSW: Sydney (Det. A.
Lea). WAM: NSW: Cooma, v.1935. ANIC: ACT: 35.16S
149.06E, Black Mt 600m, Mar. (sweeping), Oct. (FIT);
SA: 34.2 IS, 139.3 IE, Brookfield Con.Pk. x-xi.1992, FIT
(2). Lea (1918): NSW: Hunters Hill, 30.V.1917. Lea
(1925): Vic: Lakes Entrance, October, SA: Mount Lofty
Range, May. Oke (1923): Vic: Bacchus Marsh,
Chalcoponera metallica.

DIAGNOSIS. L: 1.87; W: 0.75; E/Pn L: 1.50;
E/PnW: 1.31; Pn W/L: 1.33; E L/W: 0.86; Pr/Py:
1 .00; Sterna: 0.56, 0.12, 0.56; Tibiae: 0.69, 0.69,
0.75. Chlamydopsis ectatommae , C. kununurra
sp. nov., and C. acutricha sp. nov. are all very
similar in the shape of their humeral trichome and
are diagnosed together here. In all of these the
apex of the trichome is subacute, projecting
slightly above the middle of the inner setal fringe,
and bears a few dorsally or mesally directed
setae, which are separate from the inner arc. This
projection and a small area around its base are
distinctly smoother than the surrounding
integument. (The only other known species that
has a similar form of trichome is C.
myrmecophila sp. nov., in which the apex is only
barely or not at all projecting, and the lateral
surface of the trichome is deeply vertically
furrowed). Chlamydopsis acutricha is the
smallest of these three ( 1 ,62mm), has a median

pronotal tubercle which is blunt rather than
distinctly acute, has the elytra nearly smooth
behind the trichomes rather than coarsely reticulo-
punctate throughout, and has the metastemal disk
entirely impunctate. Chlamydopsis ectatommae
and C. kununurra are more similar, differing
primarily in pronotal shape. In C kununurra the
median pronotal tubercle is more strongly
elevated and acute, the medial portion of the
anterior pronotal margin is more distinctly
elevated, shallow oblique furrows extend
anterolaterally from the scutellum, and the
pronotal reticulae, particularly along the anterior
margin, are more distinctly elongate.

REMARKS. The type was reportedly collected
in a nest of Ectatomma metallicum (Lea, 1912).

Chlamydopsis kununurra sp. nov.

(Figs 10B, 1 1 B, 12, 28D)

MATERIAL. HOLOTYPE (QMT 108584) <J: N WA,
Kununurra, 22.XI1. 199 1-5.1. 1992, R.I. Storey.

DIAGNOSIS. See diagnosis above under C.
ectatommae.

DESCRIPTION. L: 1 .93; W: 0.72; E/PnL: 1.70;
E/Pn W: 1 .3 1 ; Pn W/L: 1 .52; E L/W: 0.85; Pr/Py:
0.94; Sterna: 0.56, 0.16, 0.56; Tibiae: 0.81, 0.75,
0.84. Body subquadrate, rufescent (the type is
likely teneral), dorsal surfaces mostly reticulo-
strigose; Frons with sides rounded, disk
reticulopunctate, with sparse elongate setae;
labrum rounded; antennal scape widest at basal
one-third, abruptly narrowed to base, distal
margin straight, narrowed to rounded apex,
reticulate and setose as frons; antennal club of
male 1 .25 x length of scape.

Prothorax 1 .5 X as wide as long, sides uneven,
wide near base, bluntly projecting, narrowed in
anterior half, w r ider, subacute in front of
anterolateral groove; medial portion of anterior
margin weakly elevated, notched at middle,
separated from lateral portions; lateral portions
more strongly elevated, arcuate, ending laterally
where intersected by well developed antero-
lateral groove; pronotal disk depressed behind
anterior margin, more broadly so at sides,
elevated and acute at middle, posterolateral
comers convex, separated from medial elevation
by shallow, oblique antescutellar furrows; disk
entirely reticulopunctate, reticulae varied in size,
generally elongate, diverging around median
tubercle.

Anterior prosternal margin grooved, sinuate,
projecting on either side of middle; prosternal

184

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 10. Dorsal views of Chlamydopsis spp. A, C. ectatommae. B, C. kununurra. C, C. acutricha. D, C.
myrmecophila. E, C. variolosa (d). F, C. variolosa ($). G, C. mallee (d). H, C. mallee ( 9).

REVIEW OF CHLAMYDOPSIS

185

disk slightly depressed behind anterior margin,
entirely reticulopunctate, narrowed posteriorly,
acutely emarginate at apex.

Elytra about one-fifth wider than pronotum,
parallel-sided, narrowed to base and apex;
humeral trichomes small, closed laterally, open
mesally with an arc of mesoventrally directed
setae, with an acute peak above, the inner surface
of this peak with a separate cluster of mesally
directed setae, its outer surface smooth; anterior
superficial groove oblique, extending from humeral
comer to inner base of opening of trichome;
mediobasal depression with transverse, arcuate
carinae; elytral disk reticulostrigose; apical
elytral margin subcarinate, with elongate setae
(some setae in "bundles'); elytral disk otherwise
very sparsely setose.

Mesostemum about 4x as wide as median
length, projecting at middle, densely punctate,
crenulately so along posterior margin; meso-
metastemal suture impressed, metastemal disk
with large punctures behind suture and at sides,
with only few small punctures at middle; 1st
visible abdominal stemite with continuous row
of large punctures separated by about their widths
along anterior margin and inner postcoxal stria,
more sparsely punctate behind.

Legs generally slender, pro femur and tibia
punctate on outer surfaces, meso- and metafemora
and tibiae less so; protibia acutely angulate
one-third from base, meso- and metatibiae bluntly
angulate.

REMARKS. The name of this species refers to
the type locality.

Chlamydopsis acutricha sp. nov.

(Figs IOC, 11C, 12)

MATERIAL. HOLOTYPE 9: 12.40S 142.39EQld, 3km
W Batavia Downs, 23 Aug - 16 Sep 1992, Flight Intercept
Trap, P.Zborowski & L.Miller, in ANIC.

DIAGNOSIS. See above under C. ectatommae.

DESCRIPTION. L: 1.62; W: 0.69; E/Pn L: 1.36;
E/Pn W: 1.19; Pn W/L: 1.23; E LAV: 0.94; Pr/Py:
1.00; Sterna: 0.44, 0.09, 0.47; Tibiae: 0.56, 0.50,
0.62. Body rufescent, elongate, nearly parallel
sided; frons about as wide as long, sides rounded,
disk reticulate and with a few elongate setae;
labrum short, rounded, reticulate; antennal
scapes widest just beyond basal one-third,
slightly narrowed to apex, more abruptly to base,
surface reticulate and bearing 2-3 elongate setae;
antennal club of female about three-fifths length
of scape.

Prothorax about one-fourth wider than long;
sides weakly margined near base; anterior margin
distinctly but not strongly raised above antennal
cavities, interrupted by a groove extending
posterolateral ly from edge of antennal fossa to
circumcoxal stria; medial portion of anterior
margin more or less flat; anterior and lateral
margins with a few elongate setae; pronotal disk
shallowly depressed in anterior corners,
otherwise" strongly and uniformly convex; disk
rather shallowly reticulately punctured, with
sparse mostly deeply bifid setae.

Prosternum with anterior margin sinuate,
strongly grooved, this groove continuous with
circumcoxal stria; prosternal keel narrowed
posteriorly, emarginate at apex, prosternal disk
mostly densely punctate, with small impunctate
spaces medially.

Elytra about one-fifth wider than pronotum,
parallel sided, narrowed to base and apex; convex
in posterior two-thirds, depressed basally;
humeral trichome small, shallowly furrowed
laterally, open mesally with a fringe of ventral ly
directed setae, with an acute peak above, the
inner surface of this peak with a separate cluster
of mesally directed setae; superficial groove of
anterior elevation curved, mostly horizontal,
curving from anterior elytral comer to inner base
of trichome opening; elytral disk shallowly
reticulostrigose, mostly smooth in a longitudinal
band behind trichome; apical elytral margin with
elongate setae (some setae in ‘bundles ), a few of
these extending up the apical hall of the elytral
suture, elytral disk otherwise almost entirely
glabrous.

Mesostemum acutely projecting anteriorly,
punctate, elevated along mesometastemal suture;
metastemal disk punctate along anterior margin,
elsewhere entirely impunctate, very smooth; 1st
visible abdominal stemite similarly punctate
anteriorly (and with a few punctures laterally),
impunctate behind.

Propygidium and pygidium of approximately
equal length medially; propygidium flat along
base, becoming strongly convex apically;
pygidium weakly convex; both propygidium and
pygidium with reticulate sculpture, that of
pygidium fading in apical one-third.

REMARKS. The name of this species refers to
the acute dorsal apex of the humeral trichome.

186

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1 1 . Lateral views of Chlamvdopsis spp. A, C. ectatommae. B. C. kununurra. C, C. acutricha. D, C
myrmecophila. E, C. variolosa (<J ). F, C. variolosa ( 9 ). G, C. mallee (<S). H, C. mallee ( 9 ).

Chlamvdopsis myrmecophila sp. nov.
(Figs 10D, 1 1 D, 12, 29B)

MATERIAL. HOLOTYPE (QMT1 08585) 6: Australia:
N Qld, Hann Tableland, 13km WNW of Mareeba,
8.XII. 1993-13.1. 1994, R.Storey & S.DeFaveri, MDPI

F.I.T.-site 3 1 . PARATYPES (14): 4 ex.: same as holotype;
2 ex.: same but 13.i-l .iii. 1994; 1 ex.: same but
5.xi-8.xii.l993; 1 ex.: same but 17.ii-20.iii.1989; 1 ex.:
same but 9.xi-7.xii. 1 988; 2 ex.: same but 1 7.i- 1 7,ii. 1 989; 2
ex.: NEQ: Evelyn, 21.xii.1965; 1 ex.: same locality
28.77.1967; in AMB, DPIM, ANIC, MSCC.

REVIEW OF CHLAMYDOPSIS

187

DIAGNOSIS. While this species is similar to the
preceding three in that a separate setal bundle is
present at the apex of the trichome above the
mesal setal fringe, this setal bundle is not borne
on a prominent elevated tubercle, as in the others.
The lateral surface of its trichome is also unique
in the ectatommae group, bearing a deep vertical
furrow from the apex of the trichome nearly
halfway down the epipleuron, expanding slightly
ventrad. This is only an indentation, and not a
lateral opening to the trichome. This species is
uniquely setose, bearing rather sparse, but quite
long setae on essentially all surfaces.

DESCRIPTION. L: 2. 1 2; W: 0.8 1 ; E/Pn L: 1 .62;
E/Pn W: 1 .29; Pn W/L: 1 .46; E LAV: 0.86; Pr/Py:

1 .06; Sterna: 0.62, 0. 12, 0.59; Tibiae: 0.90, 0.87,
1.00. Body subquadrate, rufescent brown, the
pronotum appearing very slightly darker than
elytra, most surfaces reticulately sculptured,
bearing sparse but conspicuous long setae.

Frons about as broad as long, sides and anterior
margin slightly rounded, disk reticulate; labrum
short, arcuate, reticulate, with 6-10 long setae;
antennal scapes arcuate, about twice as long as
median width, lateral margin abruptly expanded
one-third from base, evenly arcuate distally, with
a few long discal setae and a lateral fringe of
shorter setae, surface reticulate; antennal club of
female about 2/3 as long as scape, antennal club
of male about 1 .3 x as long as scape.

Pronotum 1.6 x as wide as long, sides
margined, not raised, nearly straight and slightly
convergent; anterior margin slightly inwardly
arcuate at middle, oblique and inwardly arcuate
laterally above antennal cavities, middle portion
slightly reflexed, lateral portions more so;
anterior pronotal margin with a continuous
groove running laterally halfway along dorsal
margin of antennal cavity, departing from margin
posterolaterally, joining circumcoxal stria;
pronotal disk uniformly convex, reticulately
sculptured, bearing sparse conspicuous setae.

Prosternum about twice as wide as long;
anterior margin slightly deflexed, sinuate,
projecting on each side, deeply grooved, this
groove departing from margin laterally, curving
posteriorly and continuous carinate circumcoxal
stria; prostemal keel narrowed posteriorly, acutely
emarginate at apex; prostemal disk reticulately
punctate and with elongate setae.

Elytra with sides nearly parallel, slightly
sinuate beneath trichomes; trichomes strongly
produced dorsally, appearing transversely
‘pinched’, anterior and posterior elevations

meeting narrowly at apex, deeply impressed
laterally, mesally excavate and lined with
ventromesally directed setae; anterior elevation of
trichome with oblique superficial groove from
anterolateral comer of elytron to inner apex of
trichome; elytra broadly transversely depressed
between trichomes; elytral disks mostly
uniformly reticulate, the reticulae between the
trichomes transverse; marginal stria of elytra
complete except along basal half of elytral suture
and on basal margin.

Mesostemum slightly more than twice as broad
as long, acutely projecting anteromedially,
reticulate, marginal stria not evident; meso-
metastemal suture impressed; metasternal disk
shallowly reticulopunctate in basal 1/5, laterally,
and on lateral half of posterior margin between
metacoxae, elsewhere smooth, with sparse
elongate setae throughout; 1st abdominal stemite
with" row of shallow punctures along anterior
margin, and sparsely scattered elsewhere.

All tibiae elongate, slender, angulate one-third
from base; profemur and protibia shallowly
reticulate, others smooth.

Propygidium 2/3 as long as wide, depressed
along basal margin, convex elsewhere, uniformly
reticulopunctate throughout; pygidium weakly
convex, sparsely punctate in basal 1/3, otherwise
smooth.

REMARKS. One of the paratypes offers a unique
insight into the trichome functionality. It died in
the death grip of an ant, presumably its host, with
its mandibles eternally locked on the inner and
outer furrows of the beetle’s humeri. This ant
(missing its caster and one petiolar segment)
appears to be an Aphaenogaster. Based on
“Australian Ants Online (Shattuck & Barnett,
2001) two species of this genus are known to
occur near the type locality of C. myrmecophila ,
4 longiceps Smith and A. pvthia Forel. I he
former of these has previously been reported as
host of Chlamydopsis carinicollis (Lea, 1919).
The ant’s position on the specimen tends to
support Oke’s ( 1 923) contention that the humeral
modifications were perhaps little more than
handles for the hosts to carry the beetles around.
The name of this species refers to its habits, this
being one of the few new species where a host
record confirms myrmecophily.

Chlamydopsis variolosa Lea, 1910
(Figs 10E-F, 1 1E-F, 15)

Chlamydopsis variolosa Lea, 1910: 206; Type: [illegible if
Dalby, as published]/ 14169, Chlamydopsis variolosa Lea,

188

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 12. Collecting records for species of Chlamydopsis longipes and
ectatommae groups.

Queensland; SAM. examined. 2000; type locality: Dalby,
Queensland [27°10 , S,151°15 , E].

Chlamydopsis atra Lea, 1914a: 250; Lectotype <$, hereby
designated: atra Type Lea Mt Tambourine/ T.1300
Chlamydopsis atra Lea Queensland, Type [in red]; SAM,
examined. Paralectotype 6 : same data as type; these two
type specimens are mounted side by side on a single card.
TY’ had previously been written on the card behind the
left-most specimen; this is the specimen designated here as
the Lectotype. New Synonymy.

MATERIAL. QMB: SEQ: 26°53’S 152°09’E, Benarkin
School, 14.xi.94-26.i.l995, GM, FIT, open forest [1 6. 1
9 ]; SEQ:25°1 3 ’S 1 49°0 1 ’E, Expedition Ra. Nat. Pk, 5729,
‘Amphitheatre’ yards, 440m, 19.xii.974.iii. 1998, DC &
GM, open for. FIT. NHM: SEQ: Brisbane, Samfoid,
27.xii.1982. HAHC: SEQ: 27°19'54 t, S, 152°45 , 29 ,, E, Mt
Glorious, 26.xi.1997, MT. CMN: AUST: Qld, Mt
Glorious, 630m, 28.ii-9.iii. 1984, L.Masner,MT. Although
also reported from northern Queensland (Lea, 1914a), it
seems likely that this record should refer to one of the other
species from that area described here. The specimen so
referred to has not been seen.

DIAGNOSIS. L: 2.09; W: 0.75; E/Pn L: 1.79;
E/Pn W: 1.38; Pn W/L: 1.42; E LAV: 0.91; Pr/Py:
1.19; Sterna: 0.56, 0.12, 0.62; Tibiae: 0.81, 0.84,
1.00. Chlamydopsis variolosa , C. pecki sp. nov.,
and C. mallee sp. nov. are a closely related and
morphologically similar trio of species. The most
striking character uniting them is the strong
dimorphism in the sculpturing of the elytra,
pronotum, and in the degree of development of
the humeral trichomes. The putative males of all
three have most of the body reticulostrigose,

while the females are almost
entirely smooth. The elytra
and pygidia are the most
distinct points of contrast
between the sexes (and the
frons and prosternum the
least), but the difference is
marked throughout. The
trichomes of the females are
more strongly elevated
dorsal ly, and in C. mallee ,
have the inner edge of the
trichome more broadly (and
acutely) emarginate. The
females also have the
pronotum more distinctly
margined laterally, resulting,
in C. pecki and especially C.
mallee , in the near total
obliteration of the
characteristic anterolateral
pronotal groove. Features
shared by both sexes include
relatively small body size
(<2.1mm), lack of median
pronotal tubercle, deeply incised anterior
superficial humeral groove, transverse series of
setal bundles originating at the apex of the
humeral trichome continued for a varied distance
down the lateral surface, and generally small
trichome opening, the inner setal fringe of which
originates from a short arc, but consists of
elongate, ventrally directed setae, which largely
conceal the inner opening of the trichome (except
in female of C. mallee). Most also have
prominent setae on the frons, and along the
pronotal and elytral margins (except in male of C.
mallee). Chlamydopsis variolosa may be
separated from the other two species by the very
restricted origin of the inner trichome fringe. The
arc spans a longitudinal distance equivalent to
about half the width of an antennal club. The
setae themselves project mesoventrally for a very
short distance, then bend ventrally, extending
nearly to the floor of the mediobasal elytral
depression. In the other two species the originat-
ing arc of this fringe is more extensive,
equivalent in length to at least an antennal width
or greater. In the female of C. variolosa the
anterior elevation of the trichome is more
strongly elevated, projecting well above the
posterior elevation, and the pronotum is
margined, although the anterolateral groove from
the antennal cavity remains distinct.

REVIEW OF CHLAMYDOPS1S

189

REMARKS. Describing C. atra, Lea appears to
have forgotten C. variolosa entirely. When
diagnosing atra from similar species, C.
variolosa is not even mentioned. The two type
localities are separated by less than 200km in
southeastern Queensland, and their synonymy
appears clear. Lea (1914a) reported collecting the
types of C. atra "in January by means of the
sweep net, used on low herbage, ferns, etc., late in
the afternoon’.

The sexual dimorphism in this and the
following two species requires additional study.
Originally the putative females of C. variolosa
and C. pecki , especially, had not even been
considered members of this species group, due to
the vague or absent anterolateral pronotal groove.
At present, only single females are known for
either of these two. Nonetheless, in all cases
putative males and females have been collected
at identical localities and trap dates, and no other
candidates for conspecifics (morphospecies
consisting of only a single sex) are known. It is
also relatively clear (in 2 of 3 cases) that the
males and females of each putative species share
similarities, primarily in trichome morphology,
that would place them closer to each other than to
any other species. Finally it is worth pointing out
that this same mode of dimoiphism, with females
showing smoother elytra, is the same as that
found in the strigicollis and, to a lesser extent,
pygidialis groups. This represents at least two,
and possibly three separate origins.

Chlamydopsis niallee sp. nov.

(Figs 10G-H, 1 1G-H, 15)

MATERIAL. HOLOTYPE 8: 34.2 IS 139.3 IE SA.
Brookfield Con. Pk., 31 Mar.-29Apr. 1 992, E.S.Nielsen
F.I.T. m F.I.T. ANIC 1233 mallee #2; in ANIC.
PARATYPES ( 1 5): 8 males: same data as holotype; 1 8 , 2
9 9 : same but 28.v-5.viii. 1 992, FIT. ANIC 1 260 mallee U
2; 1 9 : same but 7.x-7.xi. 1 992, FIT, ANIC 1 266 mallee #2:
1 d: same but 20.V-1.V. 1993, FIT, ANIC 1297, mallee #2;
1 8: SA: 34.19S 139.30E, Brookfield Con. Pk.,

31.111- 29.iv.1992, EN, FIT, ANIC 1232 mallee with
Triodia #1; 1 8 SA: 34.22S 139.27, Brookfield Con. Pk.

30. 1 11- 3. i v. 1992, Site 3, Berlaseate ANIC 1231,
Euc.largijlorvm, clay pan, litter; in ANIC, QMB, SAM.
MSCC.

DIAGNOSIS. Both males and females of this
species are very similar to those of C. variolosa
and C. pecki , and are only diagnosed from these
two here. See above to separate these three from
other Chlamydopsis. First, the differences
between the sexes of this species are more
pronounced than in the above species, and they
are diagnosed separately. Males are nearly

glabrous, lacking frontal and anterior marginal
pronotal setae, and the originating arc of inner
trichome setae is more extensive, forming a
complete semicircle, rather than a short arc as in
the preceding, while the humeral region as a
whole is not strongly developed. The humeral
trichomes of the females, on the other hand, are
very strongly elevated, projecting well above the
surrounding elytral surface, with the apex higher
than the mesal opening. The inner trichome
fringe is unique, forming a slightly rounded ‘V’,
with only short setae projecting horizontally over
the broadly exposed opening, not directed
ventrally. As above, the female has conspicuous
elongate setae on the frons and pronotal and
elytral margins, but unlike the preceding species,
has setae on the disk of the pronotum, as well as
the margin.

DESCRIPTION. L: 1.93; W: 0.65; E/Pn L: 1.95;
E/Pn W: 1.38; Pn W/L: 1.52; E L/W: 0.93; Pr/Py:
1.29; Sterna: 0.56, 0.12, 0.56; Tibiae: 0.72, 0.75,
0.87. Body subquadrate, rufescent brown; frons
about as wide as long, sides rounded, disk
reticulate, setose in female; clypeolabral suture
present, labrum rounded, reticulate; antennal
scapes widest near middle base, gradually
narrowed to rounded apex; antennal club ol male
about 1 .3 x length of scape, that of female about
one-half as long as scape.

Prothorax about 1.3 X as wide as long; sides
parallel, margined; anterior margin elevated,
more so above antennal cavities than medially;
anterior pronotal margin above antennae
intersected by a groove extending postero-
lateral ly from upper edge of antennal cavity to
supracoxal carina in male, in female this groove
is entirely obsolete; lateral and anterior pronotal
margins bearing a few elongate setae; pronotal
disk shallowly depressed in anterior corners,
convex posteromedially, not tuberculate at
middle; disk of male reticulostrigose, with
strigosity longitudinally parallel anteriorly but
convenient posteromedially, disk ol female
largely smooth, very shallowly reticulate along
anterior margin.

Prostemum strongly grooved along anterior
margin, this groove curving posteriorly at sides,
meeting carinate circumcoxal stria; anterior
margin sinuate; prosternal keel narrowed
posteriorly, emarginate at apex; prosternal disk
reticulate throughout, shallowly so in female.

Elytra parallel sided, convex in posterior
two-thirds, depressed between humeral
trichomes; humeri elevated, more strongly in

190

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 13. Dorsal views of Chlamydopsis spp A, C. pecki ( <J ). B, C. pecki ( 2 ). C, C. loculosa. D, C. degallieri. E, C.
papuae. F, C.jayawijaya. G, C. lucifer. H, C. bataviae. I, C. burnetta.

REVIEW OF CHLAMYDOPSIS

191

female than in male; trichome of male semi-
circular, with mesal fringe of ventromedially
directed setae, that of female forming a mesally
open ‘V’, with short horizontal setal fringe;
anterior elevation of trichome with deep
superficial groove extending from anterior
elytral comer to inner apex of elevation, with a
few additional erect setae at apex, these arranged
in a subtransverse row of bundles; anterior half of
basal elytral depression with low transverse,
slightly oblique elevation; elytral disk and
epipleurae reticulostrigose throughout in male,
smooth in female; apical half of elytra of female
with numerous elongate setae, particularly along
margin; elytra of male v/ith setae only along
apical margin and posterior one-fifth of elytral
suture.

Mesostemum acutely projecting at middle,
disk strongly strigose, depressed in anterior
comers, carinate along mesometastemal suture;
metastemal disk with punctures along edges, but
broadly impunctate at middle; 1st visible
abdominal stemite with punctures along basal
margin, laterally, and, more faintly, along apical
margin, but impunctate at middle; legs slender,
moderately elongate, tibiae angulate at basal
third, the protibia most acutely, the others less so.

Propygidium and pygidium approximately
equal in length; propygidium depressed along
extreme anterior margin, otherwise uniformly
convex; pygidium slightly convex; propygidium
and pygidium reticulately punctured and
glabrous in male, smooth and bearing elongate
setae in female.

pronotal margin are also unusually elongate
(especially in the female), while the pronotal disk
is glabrous. The originating arc of the inner
trichome fringe is short and only slightly curved,
but about twice as long as in C. variolosa ,
approximately equivalent in length to the
antennal club width in the male, and slightly
longer in the female. In both sexes the setae of
this inner fringe project almost directly ventrad,
curving anteromesally just at their tips.

DESCRIPTION. L; 1.81; W: 0.62; E/Pn L: 1.90;
E/Pn W: 1 .33; Pn W/L: 1 .50; E L/W: 0.95; Pr/Py:

1 .07; Sterna: 0.50, 0. 12, 0.50; Tibiae: 0.69, 0.75,
0.84. Points of difference from C. mallee sp. nov.,
described in full above, are noted in the
diagnosis.

REMARKS. This species is named in honor of
Stewart and Jarmila Peck, collectors of the types,
whose worldwide work has furnished valuable
material for countless beetle studies.

Chlamydopsis loculosa Lea, 1925
(Figs 13C, 14C, 15)

Chlamydopsis loculosa Lea, 1925: 254: Lcctotype 9, hereby
designated: Lea, Type, loculosa. Beverley / 15775
Chlamydopsis loculosa Lea, W. Australia Type; SAM,
examined, 2002; paralectotype 9: same data as type
(mounted on the same card); these two type specimens are
mounted side bv side on a single card. ‘TY‘ had
previously been written on the card behind the left-most
specimen; this is the specimen designated here as the
Lectotype. An additional paralectotype has been
designated in the AMS from the same locality. The type
series originally included specimens from Swan River,
WA; their current repository is unknown.

REMARKS. The name of this species refers to
the vegetation at the type locality, derived from
the common growth form of the local Eucalyptus.

Chlamydopsis pecki sp. nov.

(Figs 13 A, 14A, 15)

MATERIAL. HOLOTYPE 6: Walpole NP, Peaceful Bay
WA, 17-26 Jun. 1980, S.& J. Peck, SBP58/ pan traps
coastal sand heath; in ANIC. PARATYPES: 1 9: same
data as type; in ANIC.

DIAGNOSIS. Both males and females of this
species are very similar to those of C. variolosa
and C. mallee sp. nov., above, and are only
diagnosed from these two here. See above to
separate these three species from other
Chlamydopsis. The transverse series of outer
humeral setae of this species is longer than in
either of the other two, extending from the apex
of the trichome down approximately one-third of
the epipleural surface. The setae of the anterior

RECORDS. NHM: WA: Beverley: WAM: all WA:
Beverley; Culham; Bejoording; New' Norcia, Spencer s
Br<x)k, Midland, Murchison River. SAM: SA: Gammon
Ra NP Weetootla Gorge, under stones, 16.ix. 1990; ANIC:
WA: Midland, 7.vi.48; NSW: 15km N of Coombah,
31 x 1975, on worker in Rhytidoponera maniac nest
(#1438), P.S. Ward (1); NSW: Kinchcga NP, 32.30S.
14 ? ?0E iii 1986, rod sand dune. McMillan (1950): all
WA~ Btinbury; Perth (King’s Park); Guildford: West
Midland; Spencer’s Brook; Northam; Bolgart; Albany.

DIAGNOSIS. L: 2.52; W: 0.93; E/Pn L; 1.70;
E/Pn W. 1 .42; Pn W/L: 1 .27; E L/W; 0.94; Pr/Py:
1.17; Sterna: 0.69. 0.3 1 , 0.72; Tibiae: 1 .25, 1 .37.
L71. C. loculosa can be easily distinguished
from all other Chlamydopsis by its small
nonsetose trichomes. The humeri are not
elevated: a deep humeral groove extends from the
anterolateral elytral comers into a simple deep
pit, the diameter of which is only about 2 x the
width of the humeral groove. The legs of C.
loculosa are also unusally elongate, with nearly

192

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 14. Lateral views of Chlamydopsis spp. A, C. pecki ( 6 ) B,C. pecki ( 9 ). C, C. loculosa. D, C. degallieri. E,
C. papuae. F, C.jayawijaya. G C. lucifer. H, C. bataviae. I, C. burnetta.

REVIEW OF CHLAMYDOPSIS

193

one-fourth of the meso- and metafemora
extending above the dorsal surface of the elytra
when the legs are retracted; the outer marginal
angulations of the meso- and metatibiae are also
poorly developed.

REMARKS. McMillan (1950) published some
interesting observations on the biology of C.
loculosa (erroneously as C. duboulayi
Westwood; McMillan, 1954). The very elongate
legs are apparently not locomotory specialis-
ations, as is commonly assumed, but facilitate
phoresy, serving to grasp the thorax of their
ponerine hosts (see Table 1).

Chlamvdopsis degallieri sp. nov.

(Figs 13D, 14D, 15)

MATERIAL. HOLOTYPE 8: 20km E Norseman WA,
12-13 Jan. 1986, Reid, Gullan&Lewis, light trap in tall
mal lee/Melaleuca scrub; in ANIC. PARATYPES (6): 5
males: same data as holotype; I 8.: SA: -15km SSE of
Streaky Bay, 20-2 l.xii. 1985, Reid, Gullan&Lewis, ex.
light trap in mallee scrub; in ANIC, SAM, MSCC.

DIAGNOSIS. Chlamydopsis degallieri sp. nov.
is a distinctive member of this group. While it
bears a distinct anterolateral pronotal groove, this
groove lies outside prominently elevated lateral
pronotal margins, which are continuous with the
more weakly elevated anterior pronotal margin.
This is a unique configuration in Chlamydopsis.
In general body form, and in its essentially
unelevated humeral trichomes, it resembles C.
loculosa , and the two may be related. However
the trichome of C. degallieri bears conspicuous
mesal setae, and its legs are not unusually
elongate.

DESCRIPTION. L: 2.43; W: 0.78; E/Pn L: 2.12;
E/PnW: 1.37; Pn W/L: 1.52; E LAV: 1.02; Pr/Py:
1.13; Sterna: 0.62, 0. 1 6, 0.8 1 ; Tibiae: 0.8 1 , 0.93,

1 .03. Body elongate, light rufescent brown, most
surfaces nearly uniformly reticulately punctate,
with conspicuous granulate microsculpture
within punctures. Frons about as wide as long,
sides rounded; clypeolabral suture straight to
slightly outwardly arcuate; labrum short,
rounded; antennal scape widest near middle,
outer margin subacutely angulate, narrowed to
rounded apex; antennal club of male about 1 .2 x
as long as scape.

Prothorax about 1.5 x as wide as long; sides
margined, elevated, continuous with
anterolateral portion of anterior margin, slightly
narrowed to front; anterior pronotal margin
somewhat interrupted between medial and

anterolateral portions; pronotal disk depressed
along lateral and anterior margins, but convex
medially.

Anterior margin of prosternum deeply
grooved, sinuate, acutely projecting on either
side; prosternal disk slightly transversely
depressed behind anterior margin; prosternal keel
narrowed posteriorly, elevated between coxae,
emarginate at apex; prosternal disk uniformly
punctate, punctures with granulate
microsculpture within.

Elytra 1.5 X as wide as pronotum, parallel
sided, narrowed in basal one-fifth and apical
one-fourth; humeral trichomes weakly elevated
trichome opening small, mesal, with fringe of
setae, some of which are erect and extend
dorsally above subacute peak of trichome,
elevation of trichome slightly furrowed laterally,
humeral groove present, slightly arcuate,
extending from anterolateral elytral comer to
opening of trichome, meeting an oblique, blunt
ridge extending from anterior margin next to
scutellum; blunt, transverse tubercles present on
each side behind scutellum; scutellum small.

Mesosternum about 2 x as wide as long,
nterior margin projecting at middle; disk punctate
s in prostemum; mesometastemal suture visible
ot strongly impressed; metastcmal disk elevated,
trongly convex in posterior three-fourths,
moderately punctate in anterior one-fourth and at
ides, punctures smaller and sparser along
lidline and posterior margin; 1st visible
bdominal sternite uniformly moderate ' y
unctate; legs not elongate, all tibiae blunt y
ngulate about one-third from base.

Propygidium and pygidium about equally long
lediailv; propygidium shallowly de P ress£ j d
long anterior margin, otherwise moderately
onvex; pygidium convex medially, fain y
cpressed 'along apical edges; p r °pyg|dium
niformly reticulate/granulate, the pygidium
. . Vtolf

VlARKS. This species is named in honor of
valued chlamydopsine collaborator, Nicolas

Chlamvdopsis papuae Lewis, 1913
(Figs 13E, 14E, 15)

mvdopsis papuae Lewis, '913: 86; MATERIAL.

^ . Laloki, Papua, F. Muir, 1910, NHM,

DIAGNOSIS. Chlamydopsis papuae and the
following species are closely related, and represent
the only Chlamydopsis known from New Guinea.

194

MEMOIRS OF THE QUEENSLAND MUSEUM

Both have the anterolateral pronotal groove
distinct and interrupting the lateral portion of the
anterior pronotal margin. Both have a small but
acute median pronotal tubercle. Although both
species have the dorsal surfaces entirely reticulo-
strigose, the scuplturing of C . papuae is generally
less dense. The elytra of C. papuae exhibit some
flat surface among the posterior strigosity,
whereas the dense sculpture of the elytra of C.
jayawijaya is uninterrupted. The pronotal
punctation differs, with that of C. papuae more
generally consisting of nearly polygonal punctures
and that of C. jayawijaya entirely strigose. The
epipleural sculpturing is also somewhat different
with the strigosity of C papuae arched, but
largely continuous under the trichome, whereas
that of C. jayawijaya converges to the trichome,
with nearly all strigae discontinuous beneath it.

REMARKS. This species is known only from the
type.

Chlamydopsis jayawijaya

sp. nov.

(Figs 13F, 14F, 15)

MATERIAL. HOLOTYPE 6 :
Diuremna (nr. Nalca) [~4°22’S,
1 39°5 1 *E], 1 900-2 1 00m,

9-11. XI. 1992/ IRIAN JAYA,
Jayawijaya Prov., leg. A. Riedel; in
BMNH.

DIAGNOSIS. See above
under C. papuae.

DESCRIPTION. L: 2.87; W:
1.00; E/Pn L: 1.88; E/Pn W:
1.50; Pn W/L: 1.63; E L/W:
0.77; Pr/Py: 1.00; Sterna:
0.87, 0.25, 0.97; Tibiae: 1.21,

1 .34, 1 .56. Body large, elytra
broad, quadrate; pronotum
narrower; nearly black, very
slightly rufescent, strongly
reticulostrigose on all dorsal
surfaces. Frons and
antenennal scapes strongly
reticulated, sides of frons
slightly rounded; labrum
rounded, reticulate; antennal
scape widest one-third from
base, slightly tapered to
rounded apex; antennal club
of male about 1 .25 x length of
scape.

Prothorax about 2 x as wide
as long, sides unmargined, inwardly arcuate,
slightly narrowed toward front but abruptly
expanded around antennal cavities; anterior
pronotal margin strongly elevated but interrupted
in several places; an oblique groove extending
posterolateral ly from each antennal cavity to the
supracoxal carina; transverse anteromedial
portion of pronotal margin separated from nearly
semicircular portion delimiting each antennal
cavity; transverse medial portion also notched at
middle; pronotal disk with acute median tubercle,
and less conspicuous tubercles on each side near
base; disk depressed behind anterior margin,
elevated at middle, with small but deep medial
depression in front of scutellum.

Prostemal anterior marginal groove continuous
with circumcoxal stria; anterior prosternal
margin sinuate, projecting on each side; pro-
sternal keel narrowed between coxae, widening
out slightly behind, acutely emarginate at apex,
disk densely reticulopunctate.

REVIEW OF CHLAMYDOPSIS

195

Elytra parallel-sided, narrowed basally and
apically, transversely elevated just behind
middle, depressed basally between trichomes;
anterior and posterior elevations of trichomes
enclosing circular dorsal opening, nearly, but not
quite meeting mesally, all inner and upper
margins of elevations apparently lined with short
setae; anterior elevation of trichome higher than
posterior, nearly acute at apex, with a few
dorsal ly and posteriorly directed setae near apex,
with conspicuous, curved anterior groove from
anterior elytral comer to inner apex; anterior half
of basal depression differentiated from posterior,
slightly elevated, punctate, and setose, elsewhere
reticulostrigose and more or less glabrous; elytral
epipleurae reticulostrigose, strigosity convergent
to trichome; marginal stria of epipleuron
delimiting a conpicuous smooth marginal bead.

Mesostemum projecting anteriorly, about 4 x
as wide as median length, densely
reticulopunctate; mesometastemal suture deeply
and crenulately impressed; longitudinal meta-
stemal suture shallowly impressed but visible,
disk impunctate except near metepistemum; first
visible abdominal stemite with punctures along
anterior margin and at sides but impunctate at
middle; legs slender, elongate; tarsal groove of
tibiae straight, parallel to tibial axis; outer margin
of tibiae angulate at apex of tarsal groove (near
basal one-third).

Propygidium about two-thirds length of
pygidium at middle, depressed along anterior
margin and at sides, but convex medially;
pygidium very slightly convex; propygidium and
pygidium uniformly reticulopunctate, with
sparse but conspicuous setae.

REMARKS. This species is named for the New
Guinean province encompassing the type locality.

Chlamvdopsis lucifer sp. nov.

(Figs 13G, 14G, 15)

MATERIAL. HOLOTYPE (QMT 108586) 6 : N.T., 6km
E of Humpty Doo H2.6°S,131.25°E], 6-19.X.1990, R.I.
Storey at UV light. PARATYPE: 1 9 : NEQ: Mann R. N of
Laura [~15.6°S,144.4°E], 20.iii-24.iv.1994, PZ, FIT.

DIAGNOSIS. This species and the following are
similar and diagnosed together here. The two
may be separated from other species of the
ectatommae group by the combination of a very
prominent, acute median pronotal tubercle, and a
well delimited, outwardly arcuate trichome
which is somewhat removed from the lateral
elytral margin, with a distinct, horizontal
‘shoulder’ laterad. This mesal displacement also

results in the anterior superficial humeral groove
extending more obliquely inward from the
humeral comer, curving upward to the inner
corner of the trichome’s arcuate setal fringe. The
trichomes of C. lucifer define a much deeper arc
than do those of C. bataviae (compare Figs 13G
and 13H). In both, the origins of the inner setal
fringe are horizontally toward the centre of their
arc. In C. lucifer the apices of these setae are
curved ventrad, whereas in C. bataviae they
curve dorsad.

DESCRIPTION. L: 1.93; W: 0.75; E/Pn L: 1.58;
E/Pn W: 1 .29; Pn W/L: 1 .42; E LAV: 0.86; Pr/Py:
1.00; Sterna: 0.53, 0.12, 0.59; Tibiae: 0.62, 0.65,
0.75. Body rufescent, elytra light orange,
pronotum darker, body reticulately punctate
throughout (though with some variation in
density); most surfaces with sparse short setae.
Frons slightly wider than long, sides rounded,
disk reticulopunctate, glabrous; labrum rounded;
antennal scapes arcuate, widest at middle,
narrowed slightly to rounded apex, surface
punctate, microsculptured between punctures.

Pronotum one-fifth wider than long, sides
margined, narrowed to front; anterior margin
slightly elevated above antennal fossae, medial
portion above head not elevated; anterolateral ly
with groove extending from edge of antennal
fossa posterolaterally to supracoxal carina.
pronotum raised at middle, with a single acute
dorsal tubercle near the middle; entire pronotal
disk, including tubercle, strongly reticulo-
strigose, lateral strigosity tends to diverge around
tubercle and converge both posteriorly and

anteriorly.

Prosternum with anterior margin sinuate,
grooved, this groove continuous with
circumcoxal stria; prosternal keel narrowed
posteriorly, emarginate at apex, prosternal disk
mostly densely punctate, with interstices
conspicuously microsculptured.

Elytra about one-third wider than pronotum,
markedly lighter in color, parallel-sided,
narrowed in basal and apical fifths; humera
trichomes, more or less shelf-like laterally, but
with discrete, elevated bead at base of semi-
circular setal fringe, with dense, inwardly
directed fringe of convergent setae, curving
ventrad at their apices, open beneath this fringe;
anterior surface of trichome with oblique groove
extending from anterolateral comer of elytron to
leading edge of setal fringe; most of elytral disk
reticulostrigose, except upper one-third of lateral
surface of trichome smooth; sparse but

196

MEMOIRS OF THE QUEENSLAND MUSEUM

conspicuous setae along apical margin of elytron
and along apical half ot elytral suture; lew short
setae on anterior elevation of trichome and in
basal elytral depression.

Mesostemum projecting, densely punctate;
mesometastemal suture impressed; metasternal
disk with row of punctures along all edges but
only sparsely punctate in middle ot disk, 1st
visible abdominal sternite sparsely punctate
throughout, with punctures slightly denser along
anterior margin; legs not elongate, moderately
slender, the posterior (lower) surface ol pro lemur
densely punctate, the protibia and the meso- and
metafemora and tibiae not conspicuously
punctate but with distinct microsculpture and
sparse setae.

Propygidium slightly longer than pygidium
along midline; propygidium depressed just along
anterior margin, otherwise strongly convex,
densely reticulopunctate throughout, interstices
microsculptured; pygidium flat to slightly
convex, punctate in basal halt, microsculptured
throughout.

REMARKS. The single female from Queensland
differs slightly as follows: opening of trichome
slightly larger, particularly extended slightly
forward along anterior edge; also base ot
trichome’s upturned edge slightly constricted,
such that there is a distinct ‘shelf’ surrounding it;
elytra less strongly strigose, especially postero-
lateral ly. Additional material is needed to determine
whether these differences are individual, sexual,
or geographic. The name of this species refers
primarily to the fact that the type was collected at
light, which is very unusual in histerids.
Secondarily, the setae of the trichome converge
and recurve in a manner suggesting horns.

Chlamydopsis bataviae sp. nov.

(Figs 13H, 14H, 18)

MATERIAL. HOLOTYPE (QMT108587) 6: 12.43S
142.42E Qld, 7km S. of Batavia Downs, 19 Jun - 22 Jul
1992, Right Intercept Trap, P.Zborowski & J.Cardale.
PARATYPE: 1 <5: same locality as type, 22.vi-23.vii. 1992,
PZ&EN, FIT; in DPIM.

DIAGNOSIS. See above where this species and
the previous are diagnosed together.

DESCRIPTION. L: 1.96; W: 0.78: E/Pn L: 1.52;
E/Pn W: 1 .27; Pn W/L: 1 .48; E L/W: 0.81; Pr/Py:
1.13; Sterna: 0.56, 0.12, 0.59; Tibiae: 0.78, 0.72,
0.84. This species is very similar to, and closely
related to C. lucifer above, and a complete
description is not given here. The form of the

humeral trichome is the primary distinguishing
character. In C. bataviae the humerus is not as
strongly elevated, and the mesal fringe is not
borne on a distinctly elevated ring as it is in the
preceding species. The mesal edge forms a blunt,
obtuse, 4 V’, with the setal fringe curving upward
and slightly outward, converging above opening,
with just a few short inwardly and ventrally
directed setae beneath the upper fringe. Lateral to
the trichome there is a broad, nearly flat humeral
‘shelf. Elytral surface thoroughly reticulostrigose,
but shallowly so, particularly posterolaterally.

REMARKS. This species takes its name from the
type locality.

Chlamydopsis burnetta sp. nov.

(Figs 131, 141, 18, 28C, 29E)

MATERIAL. HOLOTYPE (QMT 108588) 9, dissected
by the author: SEQ: 25°40’S 1 5 1°26'E, Nipping Gully, Site
4. 90ct- 1 8Dec 1 998, Monieith&Gough,200m, Open forest
Intercept. 7502, in QMB. PARATYPE: 1 9 : SEQ: 25°40’S
151°26’E, Nipping Gully, Site 4. 18.xii.98-26.i.l999, GM
& Gough, 280m, O/For., FIT, 7570, in QMB.

DIAGNOSIS. This and the four species that
follow form a very distinctive, and clearly
monophyletic, subgroup within the ectatommae
group. The most conspicuous characteristic of
the group is the dense bundle of setae on the
transverse carina of the mediobasal elytral
depression. In C. pilosipes sp. nov. this bundle
consists of only a few thin setae, but in all the
others it forms a large patch of at least eight or
more setae. The trichome structure of the group is
somewhat varied, but is in all cases distinct from
any others of the ectatommae group, never
forming a simple mesal arc of setae. These five
also share: relatively elongate narrow body;
elongate legs; numerous pronotal, elytral, and
pygidial discal setae (many of them small setal
bundles); convex pronotal disk with weakly
produced, short transverse median tubercle
(except in C. contorta). Most are known from
both sexes, and show no obvious dimorphisms.
Chlamydopsis burnetta , C. zborowskii and C.
pluriseta are particularly similar, with a humeral
trichome that has a somewhat obliquely
transverse, quadrate dorsal opening, filled with a
very short but dense setal mat. In neither C.
burnetta nor C. pluriseta does the setose region
of the trichome reach the lateral edge of the
elytra. A short smooth transverse furrow,
approximately half the width of the setose area
itself, incises the lateral aspect of the humeri. In
C. zborowskii the setose patch is larger, its lateral

REVIEW OF CHLAMYDOPSIS

197

comer extending most of the way to the lateral
humeral edge. The principal difference between
C. bumetta and C. pluriseta is size, (2.8 v. 2.4mm
PE length, respectively) along with the probably
correlated longer legs of C. bumetta. Also in C.
bumetta , the metastemal disk is densely and
contiguously punctate throughout, while in C.
pluriseta all specimens show some impunctate
space, and are often entirely smooth at sides.

DESCRIPTION. L: 2.74; W: 1.03; E/Pn L: 1.67;
E/Pn W: 1 .43; Pn W/L; 1 .39; E L/W; 0.83; Pr/Py:
1.11; Sterna: 0.75, 0.22, 0.90; Tibiae: 1 .37, 1 .50,
1.74. Body elongate, dark rufescent, mostly
reticulate above, and with sparse elongate setae
(a few bundles of setae) nearly throughout. Frons
about as wide as long, sides rounded, anterior
margin very slightly outwardly arcuate; disk
deeply reticulate, with a few conspicuous setae;
labrum rounded, reticulate; antennal scapes
widest at middle, slightly narrowed to rounded
apex, reticulate, setose; antennal club of female
approximately as long as scape.

Prothorax about one-third wider than long,
widest near base, sides margined, sinuate to front;
anterior margin elevated, the anterolateral
portions more so than medial portion; antero-
lateral portion of margin delimited laterally by a
sinuate groove extending from above antennal
fossa posterolateral ly where it joins supracoxal
groove; pronotal margins with approximately
evenly spaced, prominent bundles of setae;
pronotal disk depressed behind anterior margin
and just in front of scutcllum, otherwise strongly
convex, acutely produced at middle, bearing
scattered setal bundles.

Prosternum with deep anterior marginal
groove, margin sinuate and acutely projecting on
each side; prostemal keel narrowed posteriorly,
emarginate at apex, convex; prosternal disk
reticulate and with scattered setae.

Elytra about 1.5 X as wide as pronotum,
parallel sided, narrowed in basal one-sixth and
apical one-fourth; elytra convex in most of apical
two-thirds (depressed in posterolateral comers)
transversely depressed in basal one-third, this
basal depression with low transverse carina
bearing large bundle of setae close to elytral
suture; humeral trichomes prominent, anterior
aspect nearly vertical (in lateral view); anterior
and posterior elevations of trichome separated by
shallow lateral furrow which is continuous with
mesal opening of trichome; opening of trichome
nearly rectangular, widest transversely but offset
about 30° from horizontal (towards anterior

comers); mesal fringe of trichome dense and
erect but very short; anterior elevation with
nearly straight superficial groove, extending
from anterior comers of elytra to anterior comers
of rectangular trichome opening, and with
several prominent setal bundles near apex; elytral
disk reticulostrigose and with scattered setae
except within basal depression.

Mesostemum 2.2 x as wide as long; anterior
margin acutely projecting at middle; disk deeply
reticulate; mesometasternal suture deeply
impressed; metastemal disk and 1st abdominal
stemite uniformly densely punctate (though
slightly less densely than mesostemum). Legs
elongate, slender, about one- fifth of metafemoral
length projecting above elytra when held
vertically; femora slightly clavate; outer margins
of tibiae angulate at about their midpoint.

Propygidium about as long as pygidium along
midline, slightly depressed along basal margin,
but otherwise strongly convex; pygidium
convex; both propygidium and pygidium
strongly reticulate, with sparse elongate setae.

REMARKS. This species takes it name from the
watershed (Burnett River) encompassing the
type locality.

Chlamvdopsis zborowskii sp. nov.

(Figs 16A, 17A, 18)

MATERIAL. HOLOTYPE (QMT108589) d: 12.40S
142.39E Qld, 3km W Batavia Downs, 24 Oct-23 Nov
1992, Flight Intercept Trap, P.Zborowski & A.Calder.
PARATYPES (5): 1 d . 1 9 : same locality as holotype but
16.ix-24.x.l992; 1 d, 1 9: same but 23.viii-16.ix.1992; 1
9: Old: 12.40S, 143.00E, 13km E by S Weipa,
15.viii-12.ix.1993, FIT,PZ&SS, in QMB, DPIM, MSCC.

DESCRIPTION. L: 2.40; W: 0.90; E/Pn L: 1.66;
E/Pn W: 1.41; Pn W/L: 1.41; E L/W: 0.83; Pr/Py:
1 .06; Sterna: 0.72, 0. 1 9, 0.8 1 ; Tibiae: 1 .09, 1 • 1 X
1.34. This species and the following are very
similar to the preceding species and are only
described to the extent that they differ from it.
Setal patch of humeral trichome larger, extending
laterally to the side of the humeral dorsum; inner
apex of posterior elevation of trichome more
protuberant dorsally and anteriorly, nearly
closing the mesal opening above; strigosity of
pronotum and elytra finer (narrower and more
elongate), and on elytra less deeply impressed,
some individuals (especially the females) almost
impunctate behind trichomes; legs less elongate,
metafemur barely projecting above elytra when
held vertically; punctures of metasternum
confined to middle portion of disk, sides

198

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 16. Dorsal views o f Chlamydopsis spp. A, C. zborowskii. B, C. pluriseta. C, C. contorta. D, C. pilosipes. E,
C. bifovaecollis. F, C. nielseni. G, C. australis. H, C. lepida. I, C. epipleuralis.

REVIEW OF CHLAMYDOPSIS

199

impunctate; lateral portion of 1st visible
abdominal stemite also less punctate, with no
punctures within postmetacoxal depression.

REMARKS. It is unusual to have two such
similar and obviously related species as this and
the following collected at the same locality, over
several of the same trapping periods. Yet both
forms are known from both sexes, and the
differences between them are absolutely con-
sistent among all specimens. It would be very
interesting to know if they share hosts. This
species is named in honor of Paul Zborowski,
collector of this type, as well as many other
interesting chlamydopsines.

Chlamvdopsis pluriseta sp. nov.

(Figs 16B, 17B, 18)

MATERIAL. HOLOTYPE (QMT 108590) 6 : 12.39S
142.42E Qld, 4km NE Batavia Downs, 24 Oct-23 Nov,
1 992, P. Zborowski &A.Calder. PARATYPES (11): 1 6 ,1
9 : same data as holotype; 1 <5 , 1 9 : same but 23.xi-l 1 .xii.
1992; 4 dd, 3 9 9: same but 1 6.ix-24.x. 1 992, in QMB,
DPIM, AN1C, MSCC.

DESCRIPTION. L: 2.31; W: 0.87; E/Pn L: 1.64;
E/Pn W: 1 .32; Pn W/L: 1 .46; E LAV: 0.85; Pr/Py:
1 .00; Sterna: 0.69, 0.22, 0.8 1 ; Tibiae: 1 .09, 1.12,
1.34. See C. burnetta above to separate this
species and the preceding two from all other
Chlamydopsis. This species differs from C.
burnetta in only a few characters: size smaller,
(2.4mm v. 2.8mm PE length); legs less elongate,
tips of metafemora not or barely projecting above
elytral dorsum when held vertically; inner edge
of posterior elevation of trichome projecting
further forward, nearly closing trichome opening
mesally; metastemal disk with fewer punctures at
sides, often entirely impunctate.

REMARKS. The name of this species refers to
the conspicuous setae on most body surfaces
(which it shares with most of this subgroup).

Chlamvdopsis contorta sp. nov.

(Figs 16C, 17C, 18)

MATERIAL. HOLOTYPE (QMT 108591) <J: C. Qld:
22°02’S 148°03’E, Moranbah, 5km S. 240m. 20 Dec 97-26
Apr 1998, GMonteith, FlLlnterccpt,Bendee Scrub. 5799.
PARATYPES: 3 males: same locality as holotype but
25.vi-20.xii. 1997, GM&E. Kruck, FIT, Bendee Scrub,
5642, in QMB, MSCC.

DESCRIPTION (compared with C. burnetta ,
described in full above). L: 2.62; W: 0.97; E/Pn
L: 1.71; E/Pn W: 1.39; Pn W/L: 1.48; E LAV:
0.83; Pr/Py: 1.17; Sterna: 0 . 75 , 0.22, 0.87; Tibiae:

1.12, 1.21, 1.46. Body broader, reticulation
generally less deeply impressed; body setae
fewer and single (no bundles of setae, with one
exception noted below); lateral margins of
pronotum mostly straight, convergent to near
apex, then expanded around antennal lossae,
without lateral marginal setae; oblique groove
between antennal fossa and circumcoxal stria
absent (though a faint vestigial furrow can be
seen); pronotal disk not as strongly convex, with
slight indication of transverse medial carina in
two of four specimens, with a lew single setae
near middle, but lacking prescutellar setae (seen
in both C. burnetta and C. pilosipes ); humeral
trichomes not at all rectangular almost
W’-shaped in dorsal view (the bottom ot the v ^
directed anteriorly) , the outer arm of the \
continuous with a rather deep lateral iurrow;
anterior elevation of trichome divided by anterior
superficial groove, the outer halt more strongly
produced, more or less acute in lateral view;
bundle of setae along mediobasal transverse
carina more extensive, occupying the entire apex
of the carina; metasternum and 1st visible
abdominal stemite almost entirely impunctate,
each with only a few shallow punctures along
anterior margin; legs not as elongate, ang e o
outer margin of tibiae nearer basal third
midpoint.

REMARKS. The name of this species refers to
the sinuate contours of the humeral trichome.

Chlamydopsis pilosipes sp. nov.

(Figs 16D. 17D, 18)

MATERIAL. HOLOTYPE (QMT108592)?: SEQ:
?S°1VS 149°0rE, Expedition Ra. Nat. rK, >
Arnphhhe^re yards. 44(kn. 1 9 Dec 974 Mar 1 998. Cook
& Monteith. open tbr.intercept.

DIAGNOSIS. Chlamydopsis pilosipes is the
most distinctive species of this subgroup. The
opening of the humeral trichome is entirely
different from the others, consisting mainly ot a
small, but quite deep dorsal pit. continuous
anteriorly with the superficial humeral groove,
and almost concealed above by a n acutc - ^ a '^'
posteriorly directed projection from the nne,
edge of the anterior humeral elevation
Additionally the setal bundles of the mediobasal
elytral carinae consist of only a tew setae, and the
legs are covered with fairly dense decumbent
setae, whereas those of the other species have
sparser, mostly erect setae.

DESCRIPTION. L: 2.43; W: 0.93; E/Pn L: 1.60;
E/Pn W: 1 .53; Pn W/L: 1 .27; E L/W: 0.83; Pr/Py:

200

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 17. Lateral views of Chlamydopsis spp. A, C.
zborowskii. B, C. pluriseta. C, C. contorta. D, C.
pilosipes. E, C. bifovaecollis. F, C. nielseni. G, C.
australis. H, C. lepida. I, C. epipleuralis.

REVIEW OF CHLAMYDOPSIS

201

1 .38; Sterna: 0.62, 0.22, 0.69; Tibiae: 1 .06, 1 .03,
1.15. Body elongate, subquadrate, rufescent,
reticulostrigose throughout, most surfaces
(except for posterior two-thirds of elytra) with
long, prominent setae or bundles of setae; frons
about as wide as long, sides rounded, anterior
margin straight, with about 6 prominent setae;
labrum rounded; mandibles bearing a couple
setae on outer surfaces; antennal scape widest
near middle, faintly tapered to rounded apex,
with a few setae.

Prothorax about one-third wider than median
length, sides unmargined, sinuate, widest near
base, narrowing anteriorly, then abruptly
widened around antennal fossae; anterior mar-
gins of pronotum elevated, anterolateral portions
strongly raised, rounded, interrupted by groove
extending from edge of antennal fossa postero-
laterally to supracoxal hypomeral groove; medial
portion of anterior pronotal margin not as
strongly elevated as anterolateral portions but
more or less continuous with them; anterior and
lateral pronotal margins with prominent bundles
of setae; pronotal disk depressed behind anterior
margin, otherwise strongly convex, subacutely
produced at middle, prescutellar region slightly
depressed: pronotal disk with numerous
conspicuous setae, their arrangement not quite
symmetrical; one especially prominent pair of
setae just in front of scutellum.

Prosternum with anterior margin strongly
grooved, sinuate, acutely projecting on either
side; prostemal keel narrowed posteriorly, not
elevated, reticulate and sparsely setose.

Elytra about 1 .5 X width of pronotum, sides
more or less parallel, narrowed in apical
one-third and basal one-fourth; transversely
depressed in mediobasal one-third, with low,
setose, transverse carinae within depression;
humeral trichomes prominent, with conspicuous
bundles of setae on anterior and lateral surfaces;
anterior elevation of trichome divided by deep
superficial groove, prominent and acutely
produced on either side of this groove, the inner
portion tapered, leaflike, extending posteriorly to
cover much of dorsal opening of trichome, the
small, depressed opening apparently lacking a
marginal fringe of setae; posterior elevation of
trichome as high as anterior ones, but merely
convex above; elytral disk entirely reticulo-
strigose, but this texture less strongly developed
at middle; apical elytral margin with fringe of
conspicuous setae.

Mesostemum about 2.2 x as wide as median

length, acutely projecting at middle, densely
reticulopunctate and sparsely setose; mesometa-
stemal suture finely impressed; metastemum
densely punctate anteriorly and laterally but less
so posteromedial ly, with punctures rounder,
shallower and separated by about one-half their
widths; 1st visible abdominal stemite almost
uniformly punctate, the punctures separated by
slightly less than their widths, those of the
anterior and posterior margins more closely
spaced; Legs elongate slender, the meso- and
metafemora slightly clavate, outer margins of all
tibiae angulate near middle; outer surfaces of
meso- and metatibiae smooth near bases but
increasingly strigose towards apices; all legs
densely setose, most or all setae single (not in
bundles).

Propygidium about one-fourth longer than
pygidium, slightly depressed just along basal
margin, but otherwise strongly convex; pygidium
convex; both propygidium and pygidium
strongly reticulate, with sparse elongate setae.

REMARKS. The name of this species refers to its
conspicuously setose legs.

Chlamvdopsis bifovaecollis (Oke, 1923)
(Figs 16E, 17E, 18, 29A)

Orectoscelis bifovaecollis Oke, 1923: 159, New
Combination; MATERIAL. HOLOTYPE prob. 9: Natya,
Vic 29 9 1922, C. Oke/ Orectoscelis bifovaecollis , Uke.
Type./ Presented by C.G. Oke/ 904 Type/ MATERIAL.
HOLOTYPE. T. 904 Orectoscelis bifovaecollis Oke,
1 Q? V MVM. examined. 2000.

DIAGNOSIS. This species and the following
three represent a distinctive clade, which may or
may not belong in the ectatommae group. They
are highly distinctive, having an anterolateral
pronotal groove extending from the posterior
edge of the antennal cavity posteriorly to
conspicuous pits on the pronotal dorsum (Fig.
29A). Placement of these species in the present
group is based primarily on the (questionable)
assumption of homology between these grooves
and the oblique grooves of the other members o
the ectatommae group. The shape of the humeral
trichome is also similar to that of some other
ectatommae group species (particularly C.
lucifer). The anterior prostemal groove, on the
other hand, would be somewhat unusual for this
group. This marginal groove is well impressed,
and divergent from the margin at the sides, but it
does not curve directly back to meet the circum-
coxal stria; it meets the pronotal/prostemal suture
somewhat anteriorly to it. In appearance this
condition is intermediate between the state

202

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 1 8. Collecting records for species of Chlamydopsis ectatommae group.
The species C. nielseni , C. australis , and C. lepida all share a type locality
and are collectively represented by a single point.

observed in C. setifera and C.
bumetta , and that of the rest of
the ectatommae group.

Chlamydopsis bifovaecollis
(known only from the female
holotype) itself is unique
among members of the group
in having the elytral dorsum
smooth rather than
reticulopunctate. In body
shape, and especially
trichome shape, this species
and C. lepida sp. nov. are
otherwise quite similar.

REMARKS. This species was
placed originally in
Orectoscelis ‘with some slight
doubt’ (Oke, 1923). However,
despite some general
similarity in body shape, this
species lacks any of the
characteristics of Orectoscelis
or related genera. Most
notably, the scutellum in
bifovaecollis is fully exposed.

The species was reported from near nests of
Euponera lutea and a small black Iridomyrmex ,
under a log.

Chlamydopsis nielseni sp. nov.

(Figs 16F, 17F, 18)

MATERIAL. HOLOTYPE <J: 34.2 IS 139.3 IE SA,
Brookfield Con. Pk., 31 Mar.-29 Apr. 1992, E.S.Nielscn
F.I.T. #2; in ANIC.

DIAGNOSIS. This species is the most distinctive
of the four placed in this subgroup. It is about
1 .5 x the size of any of the others, with the body
darker and more coarsely sculptured. Its pronotal
pits are deeper and located immediately behind
the upturned pronotal margin. The humeral tri-
chomes also differ somewhat, with a very limited
mesal arc of setae, which themselves are quite
elongate, extending downward and then
posteriorly for a short distance along the elytral
dorsum.

DESCRIPTION. L: 2.46; W: 0.87; E/Pn L: 1 .82;
E/Pn W: 1 .36; Pn W/L: 1 .57; E LAV: 0.85; Pr/Py:
1.05; Sterna: 0.75, 0.12, 0.81; Tibiae: 0.97, 1.03,
1.09. Body subquadrate, nearly black, very
slightly rufescent, reticulostrigose throughout.
Frons about as wide as long, sides rounded,
surface reticulate, glabrous, apical margin
slightly rounded; labrum arcuate; antennal

scapes widest near middle, slightly narrowed to
rounded apex, reticulate, glabrous; antennal club
retracted (full length not visible).

Prothorax 1.7 x as wide as median length;
sides margined, sinuate, shorter, ending
anteriorly where anterolateral margin is strongly
elevated above antennal fossae; anterolateral
margin oblique, arcuate, interrupted near sides by
groove extending from edge of antennal fossa to a
very conspicuous anterolateral pronotal pit, this
pit about half the diameter of the exposed
antennal fossa (in dorsal view), and immediately
behind the raised anterolateral margin; medial
portion of pronotal margin elevated less strongly
than lateral portions, but not separated from
them; pronotal disk depressed anteromedially
and posterorly from pits, otherwise broadly
convex, subacute at middle.

Prosternum with anterior margin strongly
grooved, sinuate, acutely projecting on either
side, the anterior groove curving posteriorly at
sides joining circumcoxal stria; prostemal keel
narrowed and elevated posteriorly, emarginate at
apex; entire disk finely reticulate.

Elytra about 1.5 x pronotal width, parallel
sided, narrowed to base and apex; convex in
apical two-thirds, flat to slightly depressed in
basal third between humeral trichomes;
trichomes widely separated, only slightly
elevated, subacute, with an apical bundle of

REVIEW OF CHLAMYDOPSIS

203

elongate setae which descend the posteromedial
edge of elevation, then curving posteriorly where
they meet the elytral surface; anterior aspect of
trichome with fine superficial groove arising
inward about one-eighth from lateral comer,
extending to the apex of the trichome at the base
of the setal bundle; elytral surface entirely
reticulostrigose, mostly glabrous with a few short
setae near apex.

Mesostemum short (<l/8 median length of
metastemum) with anterior margin projecting at
middle, disk reticulate; mesometastemal suture
strongly impressed at sides but not at middle;
metastemal disk with single row of punctures
along anterior and posterior margins, but disk
otherwise impunctate, strongly convex ('raised’
relative to metastemum and abdominal
stemites); 1st visible abdominal stemite punctate
along anterior margin, more sparsely so behind.
Legs slender and moderately elongate; profemur
shallowly punctate on lower surface, tibiae, and
meso- and metafemora with distinct
microsculpture but impunctate; outer margin of
protibia angulate near basal one-third; meso- and
metatibiae bluntly angulate.

Propygidium about one fourth longer than
pygidium, slightly depressed along basal margin,
but mostly strongly convex; pygidium flat at sides,
weakly convex at middle; propygidium and
pygidium uniformly reticulate, with sparse, short
setae.

REMARKS. It must be noted that this and the
following two closely related species were all
collected at virtually the same locality, and some
share sampling dates as well. This species and the
following arc both known from males only, while
C. lepida sp. nov. is known only from females. It
is conceivable that some of these should be paired
up (C. australis sp. nov. and C. lepida are the
most similar), but this cannot be determined with
available material. The species is named in
honour of the late Ebbe Nielsen, collector of the
type, and noted lepidopterist and biodiversity
advocate.

Chlamvdopsis australis sp. nov.

(Figs 16G, 17G, 18)

MATERIAL. HOLOTYPE 6 : 34.19S 139.30E SA,
Brookfield Con. Pk., 20 Feb.-31 Mar. 1992, A.Calder,
W.Dressler, F.I.T. #1; in ANIC.

DIAGNOSIS. This species’ elytral strigosity is
sufficient to distinguish it from C. bifovaecollis .
The shape and size of the humeral trichome will
separate it from C. lepida sp. nov., below. Those

of the present species are much smaller, in fact
rather inconspicuous from above, while those of
C. lepida are prominent, occupying much of the
outer half of the anterior one-third of each elytron.

DESCRIPTION (compared to the preceding,
fully described, species). L: 1.78; W: 0.62; E/Pn
L: 1.85; E/Pn W: 1.39; Pn W/L: 1.55; E L/W:
0.86; Pr/Py: 1 .23; Sterna: 0.50, 0.09, 0.59; Tibiae:
0.65, 0.65, 0.72. Body lighter, rufescent, more
elongate, body more rounded, most surfaces with
very" fine sparse setae; antennal club visible,
about 1.5 x length of scape; pronotum very
different, groove from antennal fossa to pronotal
pit welTdefined, delimiting rather than
interupting the anterolateral portion of pronotal
margin; pronotal pit oblong, situated at middle of
pronotal side, near margin (posterolaterally
relative to that of C. nielseni ), with lateral
pronotal margin elevated and slightly curved
around its outer edge: medial portion of anterior
pronotal margin not elevated; pronotal disk
depressed near posterior margin, but posterior
margin itself raised, carinate, especially towards
sides; middle of pronotal disk simply convex, not
subacutely projecting. Prosternal keel not
elevated relative to mesosternum, slightly
depressed between procoxae; metasternum and
1st abdominal stemite distinctly rnicrosculptured
throughout. Elytra about one-third wider than
pronotum, with sides slightly arcuate, humeral
trichomes situated more medially, very weakly
elevated, with semicircular dorsal opening, the
mesal fringe of setae directed downward,
partially obscuring cavity beneath; depression
between trichomes with blunt postscutellar
protuberance; elytral surface entirely reticulo-
strigose, with uniform microsculpturc. Legs
shorter, broader; outer margins of tibiae angulate
about one-fourth from base. Propygidium
uniformly reticulate and with uniform
microsculpture; pygidium reticulate only in basal
one-fourth, rnicrosculptured throughout.

REMARKS. The name of this species refers to
the southerly position of the type locality.

Chlamydopsis lepida sp. nov.

(Figs 16H, 17H, 18)

\TERIAL. HOLOTYPE 9: 34.19S 139.30E SA,
ookfield Con. Pk., 3 Jun.-28 Aug. 1993, S. Williams,
T #1; in ANIC. PARATYPE 9: SA: 34.2 IS 139.3 IE,

, ni. 11 ::: 'in 1 0OO CM PTT inAMIP

DIAGNOSIS. See diagnoses for C. bifovaecollis
and the preceding species.

204

MEMOIRS OF THE QUEENSLAND MUSEUM

DESCRIPTION. L: 1.68; W: 0.62; E/Pn L: 1.70;
E/Pn W: 1 .43; Pn W/L: 1 .50; E L/W: 0.79; Pr/Py:
1.14; Sterna: 0.44, 0.09, 0.53; Tibiae: 0.53, 0.56,
0.62. As mentioned above, this species possibly
represents the female of the preceding species.
The two differ substantially in trichome shape,
with those of C. lepida much larger (Figs 1 6G, H).
The elytral discs are also less strongly strigose in
the present species (a frequent sexual difference
in Chlamydopsis ), the setae of pronotum and
elytra, although fine, are more conspicuously
scale-like, and the overall body size is slightly
smaller. Otherwise , the two are very similar. An
additional feature worth noting is the antennal
club. Its outer surfaces are largely sclerotised,
with only the oblique apex and a smaller
subapical patch tomentose.

REMARKS. The name of this species refers to its
scale-like dorsal setae.

EPIPLE URALIS GROUP

The epipleuralis group is a rather
heterogeneous group of mainly temperate
species. The group is primarily characterised by
trichome characters. The setal fringes of both
anterior and posterior humeral elevations tend to
have separate inner and outer origins. The inner
and outer origins of the anterior elevation are
divided at the entry point of the anterior humeral
groove. In most species the inner apices ot
anterior and posterior humeral elevations nearly
meet, with their respective setal fringes opposing
and separating the mediobasal elytral depression
from a dorsally open, rounded lateral cavity. The
setal fringe of this lateral cavity may be
continuous, interrupted at an epipleural incision,
or, rarely, absent. All members of this group have
a well developed anterior marginal prostemal
groove that departs from the margin laterally to
meet the circumcoxal stria. Most also have the
anterior and lateral margins of the pronotum
strongly elevated. This group contains several of
the largest known Chlamydopsis species.

Chlamydopsis epipleuralis Lea, 1912
(Figs 161 171, 21, 30 A)

Chlamydopsis epipleuralis Lea, 1912: 68; MAIERIAL.
HOLOTYPE: epipleuralis Lea Type, Hornsby/ 14673
Chlamydopsis epipleuralis N. S. Wales, Type; mounted
with two host ants; SAM, examined.

Chlamydopsis epipleuralis var. mastersi Lea, 1912: 68;
Mazur, 1984: 1 10.

RECORDS. NHM: NSW: Sydney; Vic: Ballarat; SA: Ml
Lofty Rep. (mounted w. id’d Iridomyrmex anceps ), SA:
Adelaide. WAM: NSW: Sydney. QMB: NSW: Sydney.
AMS: NSW: Sydney; NSW: Como, June. MCZ: WA:

Swan River [has two labels on pin (which has two
specimens)] other says NSW: Sydney, more likely. Lea
(1925): Vic: Sea Lake; WA: Perth ‘a small species of
Iridomyrmex \ Oke (1923): Vic: Whittlesea, with
Iridomyrmex gracilis.

DIAGNOSIS. L: 2.43; W: 0.90; E/Pn L: 1.69;
E/Pn W: 1 .30; Pn W/L: 1 .48; E L/W: 0.88; Pr/Py:
1.15; Sterna: 0.59, 0.22, 0.69; Tibiae: 1 .03, 1.18,
1.31. Chlamydopsis epipleuralis is one of the
most common and widespread species of the
group. It is very closely related to C. sculplus
Oke, and the two are difficult to separate.
Together they are most easily recognised by the
form of the humeral trichome. The anterior
surface is nearly flat, and incised by a moderately
deep, straight superficial groove. The inner
anterior and posterior elevations of the trichomes
nearly meet, and are pointed apically so as to
meet at opposing, setose triangular faces. Lateral
to these inner elevations is a nearly circular
opening, setose on at least its anterior (and often
the entire) edge. Chlamydopsis convexa is
somewhat similar to these in its general trichome
shape. But its anterior superficial humeral groove
is displaced laterally, with the anterior trichome
elevation strongly convex mesal to the groove,
and the outer circular opening of the trichome
lacks setae. Chlamydopsis epipleuralis and C.
sculplus differ from each other mainly in pronotal
sculpture, with the punctures of epipleuralis
extremely shallowly impressed, those of sculptus
being much more conspicuous, and even with
intermingled finer punctures. The trichomes also
differ slightly, most noticably in the anterior
superficial groove, which has its outer edge
carinate and distinct in epipleuralis , but rounded,
indistinct in C. sculptus.

REMARKS. Not all of the above specimens have
been studied, and some may refer to C. sculptus
(below), as several such misidentifications have
been found. Oke’s (1923) host record of
Iridomyrmex gracilis refers to a valid species, but
several subspecies are now recognised. The
nominate subspecies, occurring in coastal
Victoria, is likely the proper one.

Chlamydopsis sculptus Oke, 1923
(Fig. 21)

Chlamydopsis sculptus Oke, 1923: 158; Lectotype, hereby
designated: Maccdon, Victoria]., 23.4.21, C.Oke/
Chlamydopsis sculptus , Oke. Type/ Presented by C. G.
Oke/ 902 Type; MVM. examined, 2000. Paralectotype,
hereby designated: Bendigo, V, 1.10.21, C.Oke/
Chlamydopsis sculptus , Oke. Co-Type/ 903 Paratype;
AMS. An additional specimen of this species was found
bearing a type label Bendigo, 4.10.24 [Vic]. However, the

REVIEW OF CHLAMYDOPSIS

205

collection apparently postdates the description, and the
type label should be disregarded.

RECORDS. SAM: SA: 13km E Marion Bay, Yorke Pen.
6.xi. 1981 , pitfall trap behind beach. WAM: Vic: Bendigo,
Sept and Oct. QMB: SA. ANIC: SA. AMS: Vic: Ballarat,
Vic: Maldon; SA: Adelaide. USNM: SA: Lucindale. Oke
(1923): all Vic: Sea Lake; Daylesford; Ballarat; SA: Mount
Lofty.

DIAGNOSIS. L: 2.55; W: 1.00; E/Pn L: 1.56;
E/Pn W; 1.30; Pn W/L; 1.38; E L/W: 0.88; Pr/Py:
1.26; Sterna: 0.69, 0.19, 0.78; Tibiae: 1.0, 1.0,
1.12. See diagnosis under C. epipleuralis , above.

REMARKS. Reported from nests of Iridomyrmex
sp. (Oke, 1923).

Chlamydopsis convexa sp. nov.

(Figs 19 A, 20A, 2 1 )

MATERIAL. HOLOTYPE 8 : 34.2 1 S 1 39.3 1 E SA [South
Australia] Brookfield Con. Pk. 7 Oct.-7 Nov. 1992, J.
Stelman, S. Williams, FIT #2/ F.l.T, ANIC 1266, mallec
#2; in ANIC. PARATYPES (4): 1 8 : same data as
holotype; 1 9 : same but 2.xii.91-2.i. 1992, FIT, ANIC 1224
mallee #2; 1 9 : same but 7.xi-24.xi. 1992, FIT, ANIC 1268
mallee #2; 1 8 : SA: 34.19S 139.30E, 20.x-3.xi.1991, E.
Edwards, SS, FIT, ANIC 1208 mallee with Triodia #1: in
ANIC, MSCC.

DIAGNOSIS. This species is similar to the
previous two in general body and trichome shape.
The most pronounced difference is that the inner
anterior elevation of this species’ trichome is
uniquely convex (rather than flat) and displaces
the anterior superficial groove (and the reduced
outer portion of the anterior elevation) laterally.
Its body is also somewhat narrower overall. Other
differences include: glabrous dorsum, weak to
indistinct elytral strigosity, and lack of setae
lining the outer opening of the trichome.

DESCRIPTION. L: 2. 1 2; W: 0.78; E/Pn L: 1 .72;
E/Pn W: 1 .23; Pn W/L: 1 .60; E L/W: 0.88; Pr/Py:
1.29; Sterna: 0.56, 0.12, 0.65; Tibiae: 0.72, 0.75,
0.78. Body orangeish brown, appearing largely
smooth and glabrous; punctures on many
surfaces extremely shallow, appearing granulate
within; frons about as wide as long, sides parallel
at middle, narrowed to base and apex, anterior
margin straight , disk nearly flat, glabrous, very
sparsely and shallowly punctate; labrum
rounded, impunctate with numerous short apical
setae; antennal scape with outer margin forming a
blunt 90° angle near basal one-third, apex
rounded, disk microsculptured and finely setose;
antennal club of male 1 .2 x as long as scape; that
of female 0.6 x as long a scape.

Pronotum about 1.7 x as wide as long,
margined laterally, sides weakly outwardly
arcuate, widest near middle, distinctly but not
strongly elevated; anterior margin elevated, with
medial and lateral portions continuous with each
other and with elevated pronotal sides; disk
convex at middle, with sparse, inconspicuous
punctures, separated by slightly more than their
widths.

Prostemum with anterior margin deflexed,
deeply grooved, sinuate and acutely projecting
on either side, marginal groove curving post-
eriorly at sides and merging with circumcoxal
stria; prostemal disk depressed behind anterior
margin, narrowed posteriorly; apex of keel
shallowly emarginate; prostemal disk faintly
punctate, without grooves along the proleg
depressions.

Elytra about one-fifth wider than base of
pronotum; humeral trichomes prominent; inner
edges of anterior and posterior elevations broad,
strongly convex, their opposing edges with dense
fringes of very short setae, nearly meeting; outer
edges of humeral elevations less prominent,
displaced laterally, a thin outer wall between
them closing the trichome laterally; trichome
opening round, its cavity deeper than floor of
mediobasal elytral depression; outer edge of
rounded opening finely carinate, without setae;
anterior humeral groove large, deep, extending
from humeral corners to anterior edge of lateral
trichome opening; mediobasal depression with
setose, blunt, transverse tubercles; elytral disk
flat to weakly convex posteriorly, with very fine
setae throughout, a few shallow, elongate
punctures posteriorly; posterior elytral margin
finely carinate, this carina continuous with
blunter ridge extending forward to outer edge of
trichome; epipleuron strigose, strigae converging
to trichome.

Mesosternum short, about 5 x as wide as
median length, bluntly projecting at middle, more
acutely protecting at sides, in front of mesocoxae;
disk sparsely punctate and finely setose;
mesometastcrnal suture impressed; median
metasternal suture very finely elevated;
metastemal disk sparsely and finely punctate; 1st
abdominal stemite similar in texture. Legs short,
rather broad, impunctate, with minute setae;
outer surface of profemur with well developed
oblique carina; meso- and especially metafemora
with margins arcuate; tibiae angulate at basal
one-third; tarsi laterally compressed.

206

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 19. Dorsal views of Chlamydopsis spp. A, C. convexa. B, C. striatella. C, C.formicicola. D, C. dispersa. E,
C. weiri. F, C. crowcrofti. G, C. latipes. H, C. macmillani. I, C. nullarbor.

REVIEW OF CHLAMYDOPSIS

207

Propygidium about 1 .2 x as long as pygidium,
shallowly depressed along anterior margin,
otherwise evenly convex, with sparse, very
shallow punctures, finely setose; pygidium
nearly flat, weakly depressed posterolateral ly,
faintly punctate.

REMARKS. The name of this species refers to
the anterior convexity of the humeri.

Chlamydopsis striatella Westwood, 1869
(Figs 19B, 20B, 21)

Chlamydopsis striatella Westwood, 1869: 318; Type locality

‘Swan River [WA]; type specimen not located, sought at

Oxford in 2001.

REMARKS. It is extremely unfortunate that the
identity of this species, the type of the genus,
seems terminally uncertain. Westwood's type
specimen seems to be lost. Although the species
was illustrated by Westwood, not all diagnostic
characters are clearly visible. The only clues we
have to the its identity come from a debate in the
literature (Lea, 1912, responding to Lewis, 1910).
Lewis removed C. striatella and C. formicicola
King from synonymy, and offers characteristics
for separating them from each other and from C.
inquilina Lewis. Lea agreed with removing the
two species from synonymy, but goes further to
suggest that Lewis is incorrect concerning the
identity ol C. striatella . Lea cited specimens of C.
striatella from only Western Australian localities
(Swan River [1912], and Fremantle [1910]), and
apparently bases his identification on the fact that
Swan River is the published type locality of C.
striatella . Lewis's C. striatella, on the other hand,
has only been reported from the southeastern part
of the continent. Geography would seem to favor
Lea s concept. However, I have studied specimens
from both authors' collections, and Lewis's
determination (as a species close to C. formici-
cola) is a much better match to Westwood's
figure than is Lea's detenu ination; the Fremantle
specimen cited above, housed at SAM, though
lacking prothorax and head is identifiable as a
member of the ectatommae species group, and
shares no obvious similarities with Westwood's
figure. Ultimately, as neither author makes
reference to having studied a type of any kind,
this identity may be unknowable. But for the
present, Lewis’s conception is considered more
likely correct. That being said, I have been unable
to discern any differences between Lewis's
specimens of C. striatella and C. formicicola
King (and cite his specimens under the latter
species).

RECORDS (assuming the identity asserted above holds).
WAM: NSW; Tahmoor.

Chlamydopsis formicicola (King, 1869)
(Figs 19C, 20C, 21)

Chlamydopsis formicicola King, 1 869: 74; Lectotype, hereby
designated: Byzenia formicicola RLK, ants nests,
Liverpool, NSW./ K26343; AMS, examined, 2001; 2
paralectotypes, same data as type; AMS.

Chlanmlopsis formicicola var. darwinensis Lea, 1918: 85;
Mazur, 1984: 110.

RECORDS. NHM: NSW: Glen Jones, 1 4.vii. 1921; NSW:
Sydney. AMS: NSW: Sydney; NSW: Liverpool. QMB:
23°32'S 147°18’E, Bogantungan, 13.5km N,
26.x-17.xii.2000, 880m, DC & GM, Ironbark woodland,
FIT, 9825. Lea ( 1 925): nests of Camponotus aeneopi/osus ,
Brooklyn, Hawkesbury River [both NSW]. Oke (1923):
Vic: Sea Lake; Vic: Bendigo.

DIAGNOSIS. L: 2.74; W: 1.00; E/Pn L: 1.75;
E/Pn W: 1 .13; Pn W/L: 1 .66; E L/W: 0.93; Pr/Py:
1.10; Sterna: 0.69, 0. 1 6 , 0.75; Tibiae: 1 .06, 1.21,
1 .40. This species is similar and closely related to
C. dispersa sp. nov., described in full below. The
most diagnostic difference is in the shape of the
humeral trichome. In C. formicicola , its dorsal
opening is larger, occupying most of the apex of
each elevation (rather than just the inner half).
The anterior superficial groove is deeper, and the
anterior surface of the elevation is fiat on each
side of this groove, and shallowly inclined
toward it. A small tubercle present in this groove
in C. dispersa is not evident in C. formicicola. In
addition the pronotal disk of formicicola is less
densely punctate, although the punctation is still
rather dense, and the apices of the elytra are
fringed with setae. (In the Queensland specimen,
much of the body is sparsely setose; other
specimens are otherwise bare).

REMARKS. King (1869) originally described
this species in the Byrrhidae. The reported host,
Camponotus aeneopilosus remains a valid
species, but with two subspecies, both apparently
within the range of C. formicicola. There is some
possibility that the following species represents
the male of C. formicicola, see Remarks below.

Chlamydopsis dispersa sp. nov.

(Figs 19D, 20D, 21, 29G)

MATERIAL. HOLOTYPE 6 : 12.48S, 132.42E,
Nourlangie Ck, 8km N. of Mt Cahill. N.T., 19.xi.1972,
M.S.Upton; in ANIC. PARATYPES (13): 1 ex.: same data
as holotype; 1 : same but 26.X.72, at light E.B. Britton; 2
ex.: NT: I2.47S, 132.5 IE, 19km NE. by E. of Mt Cahill,
16.xi.1972, MU; 1: NEQ: 23km E. of Mareeba (Kanervo
Rd), 29.i. 1 989, A. Howden, at light; 1 ex.: NEQ: Mt Spec,
i.69; 2 ex.: NT: Tindal, 14.31 S 132.22E, l-20.xii.1967,

208

MEMOIRS OF THE QUEENSLAND MUSEUM

HvS. •

FIG. 20. Lateral views of Chlamydopsis spp. A, C.
convexa. B, C. striatella. C, C. formicicola. D, C.
dispersa. E, C. weir/. F, C. crowcrofti. G, C. latipes.
H, C. macmillani. I, C. nullarbor.

REVIEW OF CHLAMYDOPSIS

209

light trap, WJ.M.Vestjens; 1 ex.: NT: 16.28S 136.09E,
46km SSW of Borroloola, 28.x. 1975, MU; 1 ex.: NT:
Muriella Park, Kakadu, 1 8.v. 1 987, Fay & KH, at UV light;

1 ex.: NT: Kakadu NP, S. Alligator R., Gungaree RF,
17.xii.1993, uv light, S&J Peck; 1 ex.: NT: Kakadu NP,
North Point, RF FIT, 24.xii-7.i. 1 993, S&J Peck; 1 ex.: N.
Qld: Edungalba, 14.xii.1988, H&A Howden; in ANIC,
DPIM, QMB, CMN, MSCC.

DIAGNOSIS. See under C. formicicola, above.

DESCRIPTION. L: 2.68; W: 1.03; E/Pn L: 1.61;
E/Pn W: 1.19; Pn W/L: 1.58; EL/W: 0.85; Pr/Py:

1 .26; Sterna: 0.75, 0. 1 6, 0.8 1 ; Tibiae: 1 .03, 1 . 1 5,
1.31. Body dark rufescent, subquadrate; most
surfaces densely punctate, with dense
microsculpture between punctures; firms about
1.2 X as long as wide, sides nearly straight,
narrowed abruptly at antennal insertions and
gradually to apex; clypeolabral suture straight;
disk densely punctate, with both large and
intermingled smaller punctures, ground texture
between punctures somewhat granulate; labrum
rounded; antennal scapes widest about one-third
from base, abruptly narrowed to base, gradually
narrowed to rounded apex; antennal club of male
1 .5 x as long as scape.

Prothorax about 1.5 x as wide as long, sides
margined, widening slightly towards front;
anterior pronotal margin strongly elevated,
tripartite, with the lateral portions angled back
obliquely from transverse medial portion, lateral
portions slightly arcuate; pronotal disk strongly
depressed behind anterior margin, convex in
posterior half, strongly elevated, subacute at
middle; disk densely and uniformly punctate, the
punctures somewhat convergent toward
scutellum.

Prosternum with anterior margin strongly
grooved, this groove curving posteriorly at sides
and continuous with circumcoxal stria; anterior
margin slightly deflexed, sinuate, acutely
projecting on each side of middle; prostemal keel
narrowed posteriorly, acutely emarginate at apex;
prostemal disk densely punctate throughout.

Elytra together about as wide as long, sides
rounded, widest behind middle; humeral
trichomes prominent, occupying outer two-thirds
of each elytron; anterior elevation broadly raised,
anterior surface transversely concave, with
oblique superficial groove from anterolateral
corner: superficial groove with very small
tubercle anterior to where the groove widens
around mesal opening of trichome; mesal
opening nearly circular, with setose fringe which
continues straight down inner edges of anterior

and posterior elevations; trichome with con-
spicuous lateral notch as well, this (in lateral
view) bent obliquely posterad; outer edge of
posterior elevation continuous with low marginal
carina which continuous around posterior elytral
margin; elytral disk strongly depressed between
trichomes, impunctate within depression, with
low arcuate carina from elytral suture to base of
trichome opening; disk elsewhere densely punctate
(strigose near middle) and microsculptured in the
intervening space, except smooth on trichome
elevations, particular the anterior.

Mesostemum about 3 x as wide as median
length, anterior margin sinuate, projecting
medially; anterior and lateral margins sub-
carinate; disk raised along longitudinal midline,
finely but densely punctate; mesometastemal
suture impressed; metastemum convex, with
small punctures separated by slightly more than
their widths throughout, but denser along midline
and posteriorly; 1st visible abdominal stemite
densely punctate anteriorly, more sparsely so
posteriorly, the more posterior punctures each
bearing minute seta; legs slender, elongate; pro-
and mesotibiae bluntly angulate about one-third
from base; mctatibial margin simply rounded;
outer surfaces of all legs more or less uniformly
covered with setigerous punctures, the tibiae more
finely microsculptured toward outer margins.

Propygidium about one-third longer than
pygidium along midline, weakly depressed along
anterior margin, otherwise strongly convex;
pygidium flat to slightly convex; propygidium
and basal half of pygidium densely punctate and
microsculptured, the apical half of pygidium very
sparsely punctate.

REMARKS. The two Queensland specimens of
this species are markedly broader than those from
Northern Territories, but differ in no other obvious
characters. The entire type series consists of
males, while, where checked, the few C. formici-
cola specimens examined have all been females.
Their known distributions nearly abut in central
Queensland, prompting the possibility that the
two species are sexes of the same species.
However, the differences between them would
not correspond to any other known Chlamydopsis
dimorphisms. The name of this species alludes to
its relatively broad distribution.

210

MEMOIRS OF THE QUEENSLAND MUSEUM

Chlamvdopsis weiri sp. nov.

(Figs 19E, 20E, 21)

MATERIAL HOLOTYPE <J: 12.39S142.42E Qld,4km
NE Batavia Dns. 1 6Sept.-24()ct. 1 992, FIT, P.Zborowski &
T.Weir; in AN1C.

DIAGNOSIS. This species is similar to the
preceding two, sharing an upturned anterior (but
not lateral) pronotal margin and fairly simple,
rounded, mesally open trichomes, the anterior
elevations of which are broad and more or less
flat. The outer border of the trichomc opening is
subacutely elevated in the present species, and
smooth on its outer surface, whereas in the
preceding two, while the trichome is closed
laterally, a shallow groove extends laterad from
the trichome opening to a deep epipleural
invagination.

DESCRIPTION. L: 2.62; W: 0.97; E/Pn L: 1.71;
E/Pn W: 1 .28; Pn W/L: 1 .6 1 ; E L/W: 0.83; Pr/Py:
1.17; Sterna: 0.75, 0. 1 6, 0.87; Tibiae: 0.97, 1 .06,
1.18. Body rufescent brown; frons slightly wider
than long, with sides broadly arcuate, slightly
elevated relative to disk, acutely projecting near
antennal insertions, disk coarsely reticulopunctate,
glabrous; apical frontal margin shallowly
emarginate; labrum rounded, shallowly punctate,
with numerous very short setae along apical mar-
gin; antennal scapes strongly angulate one-third
from base, basal portion of outer margin inwardly
arcuate, apical portion mostly straight, narrowed

to rounded apex; antennal club
of male about 1 .5 x as long as
scape.

Pronotum 1 .75 x as wide as
long, narrowest at base,
widening slightly to
intersection with lateral
portions of anterior pronotal
margin; sides margined, disk
slightly depressed along
margin; anterior margin
strongly elevated, the medial
portion interrupted by a notch
at middle and at sides; lateral
portions projecting forward
slightly beyond medial
portion, arcuate to sides; disk
convex, with very short, faint
transverse median carina
densely punctatostrigose, with
minute setae interspersed, the
strigae converging toward
scutellum.

Prostemum with anterior margin deflexed,
deeply grooved, sinuate and bluntly projecting on
either side of middle, the marginal groove
curving posteriorly at sides and merging with
circumcoxal stria; prostemal disk transversely
depressed behind anterior margin, and somewhat
longitudinally depressed between procoxae;
edge of proleg depression strongly carinate, and
with deep groove along inner margin of this
carina from apex of keel around to sides; keel
narrowest between procoxae, slightly widened to
apex, emarginate apically; prostemal disk densely
punctate throughout.

Elytra about 1.4 x as wide as base of
pronotum, sides more or less parallel, evenly
narrowed to base and apex; humeri strongly
elevated, particularly at sides, with lateral ‘wall’
slightly overhanging trichome opening; trichome
open mesally, anterior and posterior edges
parallel, outer edge more rounded, lined with
nearly continuous fringe of setae (though
anterior, lateral, and posterior fringes are
interrupted underneath overhanging side), those
of the inner anterior comer and outer edge
longest; anterior humeral grooves, nearly straight,
extending from humeral comer to anterolateral
comer of trichome opening; mediobasal elytral
depression occupying slightly less than basal
half, smooth, with prominent, arcuate, transverse
carinae; elytral disk outside of depression
densely punctatostrigose, slightly elevated near
posterolateral comers; apical elytral margin

REVIEW OF CHLAMYDOPS1S

211

carinate, carina diverging from margin at sides,
and extending forward about one-fourth of
elytral length.

Mesostemum about 4 x as wide as long, raised
along midline, depressed at sides, acutely
projecting at middle, disk punctate; meso-
metastemal suture strongly impressed, crenulate,
continuous at sides with postmesocoxal line;
longitudinal metasternal suture shallowly
impressed, complete; metasternal disk uniformly
but sparsely covered with small punctures,
interstices microsculptured, faintly alutaceous;
1st abdominal sternite similarly sculptured,
though punctures smaller. All legs short, slender;
tibiae angulate near basal one-third; outer
surfaces of pro femur and tibia sparsely punctate,
meso- and metafemora and tibiae with smaller
punctures, most with minute setae.

Propygidium depressed along anterior margin,
strongly convex along midline, depressed at
sides, disk coarsely reticulopunctate; pygidium
slightly convex, densely punctate in basal
one-half, more sparsely apically.

REMARKS. This species is named in honour of
Tom Weir, who collected the type and has offered
considerable assistance during this study.

Chlamvdopsis crowcrofti sp. nov.

(Figs 19F, 20F, 24)

MATERIAL. HOLOTYPE 6: 34.2 IS 139.3 IE SA,
Brookfield Con. Pk.. 12 Sept.-20 Oct. 1991, J.Stelman,
S.Williams, F.I.T.#2/ F.I.T. AN1C 1206 mallee #2; in
AN1C. PARATYPE: 1 6 : same data as holotype but 7.x-
7.xi.l992; in AN1C.

DIAGNOSIS. This and the following five
species form a relatively distinctive grouping
(referred to here informally as the ‘ latipes
subgroup’ ), united by humeral trichome shape. In
all the inner edges of the anterior and posterior
elevations of each trichome are well separated
from each other, but have opposing bundles of
setae which nearly or actually touch, while
laterally the trichome is flat, broadly rounded,
bearing an arc of marginal setae projecting
mesally above an internal ‘shelf’, which
variously conceals the opening of the trichome.
In addition, all have the pronotum at least slightly
acutely tuberculate, the anterior pronotal margin
(but not the lateral) elevated, and metatibiae
moderately to markedly elongate. The present
species is distinguished from the others
principally by the form of the anterior superficial
groove of the trichome, which is slightly
‘undercut’ toward the outside, such that the outer

edge of the groove is strongly carinate, while its
inner edge is nearly flat and noncarinate. The
sparse dorsal setae are more conspicuous,
particularly on the posterior portions of the
elytra, than in any other species in this group.

DESCRIPTION. L: 2.71; W: 1.00; E/Pn L: 1.72;
E/Pn W: 1 .26; Pn W/L: 1 .78; E LAV: 0.76; Pr/Py:
1.24; Sterna: 0.78, 0.28, 0.93; Tibiae: 1.21, 1.43,

1 .74. Body rufescent, somewhat rounded, almost
entirely densely reticulately punctate, with
sparse, elongate setae throughout; frons about as
long as wide; sides rounded, gradually narrowed
to apex, abruptly narrowed at antennal bases;
apical margin very slightly sinuate; frontal disk
reticulate, bearing -5 irregularly placed long
setae; labrum rounded, nearly semicircular,
reticulate, with anterior fringe of shorter setae;
outer basal surfaces of mandibles reticulate;
antennal scape with inner margin arcuate, outer
margin angulate, widest about one-third from
base, reticulate; antennal club of male 1.3 X as
long as scape.

Prothorax 1.5 x as wide (at base) as median
length; sides margined, parallel in basal
two-thirds, abruptly widened in apical third, with
sparse setal fringe; anterior pronotal margin
strongly elevated, the lateral thirds arcuate,
oblique" to medial third and slightly projecting
forward of it where they meet; median portion not
quite as high as lateral, distinctly notched at
middle; pronotal disk shallowly depressed
behind anterior margin, convex posteriorly,
weakly acutely produced at middle; disk entirely
reticulopunctate, the posteromedial reticulae
slightly more elongate, converging toward
scutellum; disk with sparse elongate, though
mainly decumbent, setae.

Prosternum with anterior margin deflexed,
deeply grooved, sinuate and bluntly projecting on
either side of middle, the marginal groove
curving posteriorly at sides and merging with
circumcoxal stria; prostemal disk transversely
depressed behind anterior margin, convex
posteriorly between procoxae; prostemal keel
narrowed to between coxae, then slightly
widened to apex, apex deeply emarginate; entire
prostemal disk reticulate.

Elytra approximately 1 .5 X wider than base of
prothorax; sides of elytra slightly sinuate, slightly
wider anteriorly, rounded to apex; humeral
trichomes prominent, their openings nearly
circular, broadly open mesally (and beneath),
with a lateral arc of dense, erect setae, and
separate opposing dense bunches of setae on the

212

MEMOIRS OF THE QUEENSLAND MUSEUM

inner edges of the posterior and anterior
elevations; shelf above opening where lateral
trichomal setae are inserted open in lateral view;
anterior elevation subacutely produced, with
deep, almost straight superficial groove
extending from anterolateral elytral comer to
outer edge of trichome opening, this groove
slightly undercut toward the outside such that its
outer edge is overhanging; basal elytral
depression with two arcuate carinae from elytral
suture behind scutellum to anterior edge of
opening of trichome; posterior half of elytra
unevenly convex, surface undulated in
posterolateral comers; posterior elytral margins
carinate, these carinae diverging from margin at
posterolateral corners and extending about
one-fifth around the side before diminishing.

Mesostemum about 3 x as wide as median
length; anterior margin sinuate, projecting at
middle; mesostemal disk depressed in anterior
comers, elevated at middle, uniformly reticulate;
mesometastemal suture crenulately impressed;
metastemum only very sparsely and finely
punctate, with faint alutaceous microsculpture;
postmesofemoral line ending short of metepi-
stemum; 1st visible abdominal stemite with
conspicuous row of small punctures along
anterior margin, but more sparsely punctate
behind; postmetafemoral line not well
developed, ending behind coxa; femora widened
to apex, exposed surfaces densely punctate, with
setae along anterior and posterior margin;
protibia slender, angulate about one-third from
base, densely punctate on outer surface, approx-
imately equal in length to profemur; meso- and
especially metatibiae conspicuously (about
one-sixth in the latter case) longer than their
respective femora, widened, with dense
punctures only along edges; mesotibia angulate
one-third from base; metatibia rounded, only
bluntly angulate near middle.

Propygidium 1.3 X longer than pygidium,
slightly depressed along anterior margin,
otherwise strongly convex, reticulopunctate,
sparsely setose; pygidium convex, densely
punctate in basal half, punctures slightly more
widely spaced in most of apical half.

REMARKS. This species is named for renowned
mammologist Peter Crowcroft, in recognition of
his role in establishing Brookfield Conservation
Park, type locality of this and several other
Chlamydopsis species. Although his primary
concern at the time was the southern hairy nosed

wombat ( Lasiorhinus latifrons), numerous
additional species benefit from his efforts.

Chlamydopsis latipes Lea, 1919
(Figs 19G, 20G, 24)

Chlamydopsis latipes Lea, 1919: 176; HOLOTYPE $: latipes
Lea Type, Mt Henry/ Chlamydopsis latipes Lea, W.
Australia Type. 10675; mounted with 2 hosts, reportedly
Dolichoderus (Hypoclinea) scabridus (Lea, 1919); SAM
examined, 2000.

MATERIAL. WAM: WA: Chidlow’s Well [31°86’S,
1 16°26'E], 22.V.1953, in association with R/hytidiponera]
convexa , under stone very dull day, quiescent in gallery,
East aspect. Lea (1925): WA: Armadale [32°09’E,
1 16°00'S], Rhytidoponera comexa violacea.

DIAGNOSIS. L: 3.61; W: 1.40; E/Pn L: 1.58;
E/Pn W: 1 .59; Pn W/L: 1 .29; E LAV: 0.77; Pr/Py:
119; Sterna: 0.93, 0.37, 1.2 1 ; Tibiae: 1 .74, 2. 1 2,
2.55. See above to diagnose the latipes subgroup
Within this group C latipes itself is unique in
having the elytra behind the trichomes virtually
impunctate. While faint strigosity is detectable, it
is inconspicuous at lower magnifications.
However, apart from this character and its larger
body size, C. latipes is very similar to the
following, and there is a chance that they are
sexes of the same species. There is precedent for
this sort of dimorphism within Chlamydopsis
(see strigicollis group above). But the two are
known from no identical localities, and from too
few specimens to be able to do more than suggest
their identity.

REMARKS. As currently delimited, the ant
Dolichoderus scabridus does not occur in
Western Australia. This host record may refer to
what is now called D. ypsilon ypsilon, a Western
Australian species with D. scabridus ypsilon as a
synonym. Interestingly this species and the hosts
of the nontype records are in different formicid
subfamilies.

Chlamydopsis macmillani sp. nov.

(Figs 19H, 20H, 24)

MATERIAL. HOLOTYPE 8 : Perth, John Forrest
National Park, Darling Range, Western Australia, Sept.

1 974, GH. Lowe/' in nest of Rhytidoponera violaceum - in
midst of ants/ Western Australian Museum Entomology
Reg no. 27236/ Chlamydopsis , is close to C. latipes , Det.
R.P. McMillan; in WAM.

DIAGNOSIS. See above for group diagnosis of
the latipes subgroup. This species is distin-
guished by having the lateral portion of the
humeral trichome broadly rounded, and with
relatively inconspicuous internal shelf, such that

REVIEW OF CHLAMYDOPSIS

213

nearly the entire lumen of the trichome is visible
from above. The smaller size and reticulostrigose
elytra of C. macmillani distinguish it from C.
latipes. Its evenly impressed superficial humeral
groove, and glabrous pygidium and posterior
elytral disks distinguish it from C crowcrofti.

DESCRIPTION. L: 2.83; W: 1 .06; E/Pn L: 1 .68;
E/Pn W: 1 .47; Pn W/L: 1 .50; E L/W: 0.76; Pr/Py:
1.09; Sterna: 0.78, 0.28, 1.00; Tibiae: 1.37, 1.46,

1 .8 1 . As for C crowcrofti , above, except: dorsum
with few setae on pronotal disk, lateral pronotal
margins, and on anterior surfaces of humeri,
otherwise glabrous; sides of pronotum inwardly
arcuate, nearly as wide at apex as at base.
Humeral trichomes with openings C-shaped,
open mesally, with opposing dense bundles of
short setae closing the ‘C\ outer fringe of setae
occupying most of lateral arc, with arcuate
groove above and lateral to fringe, and short,
inconspicuous inner shelf beneath it; floor of
trichome lumen smooth, glabrous, continuous
between trichomes; superficial humeral groove
of humeral trichome evenly incised, with inner
and outer edges carinate. Metasternal disk with
sparse but conspicuous small punctures, these
denser along posterior half of longitudinal
metasternal suture; punctures of 1st visible
abdominal stemite not appreciably concentrated
along anterior margin, but fairly evenly
distributed, separated by l-2x their widths;
meso- and metatibiae not longer than their
respective femora, outer meso- and metatibial
punctures largely confined to basal one-third of
their lateral margins. Propygidium and pygidium
glabrous.

REMARKS. This species is named in honor of
Peter McMillan, a long-time chlamydopsine
collector and enthusiast, who recognised that this
species was undescribed.

Chlamvdopsis nullarbor sp. nov.

(Figs 191, 201,24, 29F)

MATERIAL. HOLOTYPE d: 32.08S 126.18E, 23km
ESE of Cocklebiddy WA, 25.x. 1977, J.F.Lawrence/
berlesed from leaf litter; in AN1C.

DIAGNOSIS. This member of the latipes
subgroup is most easily distinguished by
characters of the humeral trichome. The lateral
arc of setae arises in a discrete single row, with
conspicuous flat, impunctate shelves both mesal
and lateral to it. Approximately the outer half of
the lumen of the trichome is obscured by this
inner shelf, but no carina arises from the

transverse elytral depression to close the lumen
medially (as is the case in the following two
species). The most similar known species is C.
latipes , which may be immediately distinguished
by its impunctate elytra.

DESCRIPTION. L: 3.18: W: 1.15; E/Pn L: 1.76;
E/Pn W: 1 .52; Pn W/L: 1 .5 1 ; E L/W: 0.76; Pr/Py:
1.19; Sterna: 0.93, 0.34, 1.18; Tibiae: 1.50, 1.65,
2.02. As for C. crowcrofti , above, except:
Posterior half of elytra, and propygidium and
pygidium lacking setae; pronotum barely
notched at middle; most setae of pronotum
actually bundles of two setae; humeral trichome
broadly rounded, slightly narrowed mesally, with
anterior and posterior bundles of mesal setae
meeting; outer arc of setae projecting inward
above curved shelf, concealing slightly more
than outer half of lumen of trichome; also with
impunctate shelf outside of outer setal arc,
delimited laterally by an elevated outer margin;
transverse basal elytral depression glabrous
across middle, with a few punctures beneath
inner edge of trichome opening; posterior half of
elytral disk reticulostrigose; metasternal disk
evenly punctate along anterior and posterior
margins, and along median longitudinal suture,
lateral portions of disk much more sparely and
finely punctate; 1st visible abdominal stemite
sparsely punctate at middle, more densely so
towards metacoxae; pygidial punctures smaller
but no less dense toward apex.

REMARKS. This species name is the region of
the type locality. Latin reference to the lack of tall
trees in the area.

Chlamvdopsis rotunda sp. nov.

(Figs 22A, 23A, 24)

MATERIAL. HOLOTYPE (QMT108593) 9: 12.40S
142.39E Qld, 3km W Batavia Downs, 16 Sep - 24 Oct
1992, Flight Intercept Trap, PZborowski & T.Weir, in
QMB.

DIAGNOSIS. This species and the following can
be separated from the remainder of the latipes
subgroup (see above) by the transverse basal
elytral depression between the trichomes. In both
of these it rises from the middle to the sides,
ending, beneath the inner edges of each trichome,
as a short longitudinal carina which almost meets
an impunctate shelf extending inward from
beneath the outer setal arc of the trichome. The
lumen of the trichome is thus only visible as a
short notch between these, though it is more
broadly open beneath. In other species in this
subgroup the basal elytral depression is

214

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 22. Dorsal views of Chlamydopsis spp. A, C. rotunda. B, C. latipennis. C, C. carinota. D, C. inquilina. E, C.
detecti. F, C. storeyi. G, C. matthewsi. H, C. cavicollis.

REVIEW OF CHLAMYDOPSIS

215

continuous with the trichome lumen, and not
separated from it by a carina. The meso- and
metatibiae of these two species are also not as
distinctly widened as those of any of the above
species. Chlamydopsis rotunda is distinguished
from C. latipennis by the former’s: lack of
strigosity in the posteromedial half of the
pronotum, more lightly strigose elytra, lack of
propygidial and pygidial discal setae, and
densely granulose (rather than reticulopunctate)
pygidia.

DESCRIPTION. L: 3.55; W: 1.25; E/Pn L: 1.85;
E/PnW: 1.41; Pn W/L: 1.65; E LAV: 0.80; Pr/Py:
1.20; Sterna: 0.93, 0.34, 1.21; Tibiae: 1.37, 1.56,

1 .84. Body dark rufescent, broadly subquadrate;
sculpture varied from shallowly and finely
strigose (elytra) to coarsely reticulostrigose
(anterior half of pronotum); frons as wide as long,
sides rounded, deeply reticulopunctate, with two
prominent setae near anterior margin (number
may vary in other individuals); labrum rounded,
punctate; antennal scape arcuate, outer margin
sinuate, widest just before middle; antennal club
of female two-thirds length of scape.

Prothorax 1.6 x wider than median length,
sides margined, very slightly narrowed
anteriorly; anterior pronotal margin elevated, the
inner half of supra-antennal portions most
strongly, less so at middle and at sides; pronotal
disk depressed behind anterior margin,
particularly strongly behind junction of median
and lateral portions of margin, acutely projecting
at middle and faintly convex behind middle; disk
deeply reticulostrigose in anterior half, more
shallowly so posteriorly, nearly smooth in much
of posterior half.

Prosternum with anterior margin deeply
grooved, this groove curved posteriorly and
joining circumcoxal stria at sides; margin acutely
projecting on either side of middle; prostemal
keel narrowed posteriorly, acutely cmarginate at
apex; carina separating disk from protibial
depressions prominent; disk densely and
coarsely punctate.

Elytra 1.3 x as wide as pronotum, sides
parallel, approximately equally rounded to base
and apex; humeral trichomes low, transversely
ovoid, close to lateral margins, the opening itself
limited to a short longitudinal notch, largely
concealed; trichome with setose fringe around
outer half of edge, this fringe nearly meeting
elytral margin; inner edges of trichome
elevations with small bundles of obliquely
opposing setae (which do not meet in type); basal

elytral depression broad, with transverse carina
connecting anterior elevations of both trichomes;
elytral disk undulated in posterolateral comers,
coarsely strigose in anterolateral comers, much
more finely strigose elsewhere; posterior margins
of elytra not carinate.

Mesostemum projecting at middle, elevated
along midline, anterior and lateral edges raised,
otherwise depressed, coarsely punctate; meso-
metasternal suture crenulately impressed;
metastemum mostly smooth, very finely and
sparsely punctate, with faintly alutaceous
microsculpture, mesofemoral line complete to
metepistemum; 1st visible abdominal stemite
similar in texture to metastemum, metafemoral
line ending behind coxa; legs slender, not
unusually elongate; outer surfaces of all femora
impunctate; protibia angulate at basal third,
punctate along outer margin; meso- and
metatibiae moderately broad, bluntly angulate,
margins rounded, with faint setigerous punctures
along outer margin, otherwise impunctate.

Propygidium 1.25 X length of pygidium,
shallowly depressed along anterior margin,
otherwise strongly convex; pygidium flat along
posterior margins, but convex at middle; both
propygidium and pygidium unusually textured,
rather granulately rugose, the median apical third
of the pygidium slightly smoother, both with fine
erect setae.

REMARKS. The name of this species refers to its
rotund body shape.

Chlamydopsis latipennis Lea, 1912
(Figs 22B, 23B, 24)

Chlamydopsis latipennis Lea. 1912: 67; Lectotype, hereby
designated: NW AustnV On permanent loan from Macleay
Museum University of Sydney/ Chlamydopsis latipennis
Lea, Type, N.W. Australia, ANIC; examined, 2000.

RECORDS. ANIC: SA: Brookfield Con. Pk.,
3-12.ix.1991, site 1 JL,TW&Dressler, pitfalls. This 8
specimen is about 1.5 x the size of the type, but is
otherwise very similar.

DIAGNOSIS. L: 3.40; W: 1.25; E/Pn L: 1.73;
E/PnW: 1.48; Pn W/L: 1.53; E LAV: 0.77; Pr/Py:
1.28; Sterna: 0.93,0.34, 1.12; Tibiae: 1.50, 1.62,
1.93. See diagnosis under the preceding species.

Chlamydopsis carinota sp. nov.

(Figs 22C, 23C, 24)

MATERIAL. HOLOTYPE (QMT108594) 8: 12.40S
142.39E Qld, 3km W Batavia Downs, 16 Sep - 24 Oct
1 992, Flight Intercept Trap, P. Zborowski & T. Weir.

216

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 23. Lateral views of Chlamydopsis spp. A, C. rotunda. B, C. latipennis. C, C. carinota. D, C. inquilina. E, C.
detecti. F, C. storeyi. G, C. matthewsi. H, C. cavicollis. I, C. tuberculata.

REVIEW OF CHLAMYDOPSIS

217

DIAGNOSIS. This species is easily distinguished
by the strongly upturned anterior pronotal
margin, ubiquitous setae, punctatorugose meta-
stemal disk, and prominent trichomes, which at
their apices project laterally beyond the epipleuron.

DESCRIPTION. L: 2.80; W: 1.06; E/Pn L: 1.65;
E/Pn W: 1 .39; Pn W/L: 1 .65; E LAV: 0.72: Pr/Py :
1 .20; Sterna: 0.84, 0.25, 0.90; Tibiae: 1.31,1 .50,
[metatibia missing]. Body light rufescent brown,
all surfaces conspicuously and relatively densely
setose; frons as long as wide, sides rounded, their
edges slightly projecting, narrowed to base and
apex; disk deeply reticulopunctate, setose;
anterior frontal margin slightly rounded; labrum
broad, anterior margin sinuate, projecting at
middle, setose; outer bases of mandibles
punctate, setose; antennal scapes arcuate, widest
about one-third from base, narrowed to rounded
apex, reticulopunctate and setose; antennal club
of male 1.5 x length of scape.

Prothorax 1.6x as wide as median length;
sides rising from base, continuous with the
strongly elevated anterior margin, the upper
edges finely crenulate and setose; pronotal disk
depressed behind anterior margin and at sides,
convex along midline, more broadly so
posteriorly, depressed mediobasally, with a small
prescute liar tubercle; disk more or less evenly
covered with setiferous punctures separated by
about 3 x their widths, the punctures larger and
less widely separated in anterior comers.

Prosternum with anterior margin strongly
grooved, deflexed, sinuate, projecting on each
side; marginal groove curving away from margin
at sides and joining circumcoxal stria; prostemal
disk depressed behind anterior margin, slightly
elevated between procoxae, narrowed posteriorly,
acutely emarginate at apex, densely reticulo-
punctate and with sparse decumbent setae.

Elytra widest at humeri, with prominent,
rounded humeral trichomes; opening of trichome
oval, slightly oblique, transverse, open mesally,
lined with fringe of setae; superficial groove of
anterior elevation close to margin, with outer
edge prominent, slightly undercut, inner edge
poorly defined; trichome closed laterally, with
outer edge folded over, projecting laterally beyond
epipleuron; basal elytral depression broad,
confined to less than basal half, low transverse
carinae within; posterior half unevenly convex,
undulating at sides, densely reticulostrigose and
setose; posterior elytral margin finely carinate,
slightly elevated.

Mesostemum 3 x wider than median length,
anterior margin sinuate, projecting at middle,
disk more or less flat, reticulopunctate, setose;
mesometastemal suture finely impressed, largely
obscured by punctures; metastemal disk rather
convex, densely punctate; mesofemoral line
complete to, and continued on, metepistemum;
1st visible abdominal stemite entirely densely
punctate, metafemoral line ending just behind
metacoxa, not reaching side; legs all apparently
elongate, rather slender (both metathoracic legs
missing in unique type), densely punctate and
clothed with long setae on outer surfaces; tibiae
thickened along longitudinal axis, but abruptly
thinner towards outer edge on exposed side
(closely mirroring the tarsal groove on the
concealed side).

Propygidium slightly longer than pygidium,
depressed along basal margin but convex elsewhere.

REMARKS. The name of this species refers to its
prominent, upturned anterior pronotal margin.

Chlamydopsis inquilina Lewis, 1885
(Figs 22D, 23D, 27)

Chlamydopsis inquilina Lewis, 1885: 472; ?Type: Labelled:
‘Liverpool N.S. Wales (ants’ nests)’. The type locality
cited here is from a specimen labelled as Lewis’ type in
the NHM. However the original description noted only
‘Australia (Duboulay)’ as a type locality. Lea (1919) notes
that a later citation of the type from NSW (Lea, 1912) is
likely in error, and that F.H. du Boulay's specimens all
originated from Western Australia. Thus it is not certain
that the NHM specimen labelled as type was in fact part of
the original type series.

RECORDS. NHM: WA: Swan River; WA: Mt Lawley.
WAM: Cannington, in nest of lridomyrmex conifer a,
15.x. 1952. AMS: WA: Swan River. MCZ: WA: Swan
River. Lea (1919): WA: Swan River, lridomyrmex
conifera.

DIAGNOSIS. L: 3.05; W: 1.18; E/Pn L: 1.58;
E/Pn W: 1 .35; Pn W/L: 1 .37; E L/W: 0.86: Pr/Py:

1 .35; Sterna: 0.78, 0.25, 0.87; Tibiae: 1 .46, 1 .53,
1.74. Several characters distinguish this isolated
species: The pronotal and elytral disks lack
impressed punctures, but the conspicuously
alutaceous ground texture is peppered with small,
round, untextured ‘pseudopunctures’; the lateral
pronotal margin appears ‘doubled' by an
unusually prominent, carinate circumcoxal stria;
the outer wall of the humeral trichome is deeply
incised and lacking a dense setal fringe; the
medial portions of the pronotal and elytral disks
are glabrous (except on the humeral trichomes)
but bear long golden setae around their margins;
the dense setae of the propygidium and pygidium

218

MEMOIRS OF THE QUEENSLAND MUSEUM

are much more conspicuous in this than any other
species of the epipleuralis group.

REMARKS. Only one definite male of this
species has been studied. At least that specimen
was unusual among Chlamydopsis in lacking an
elongate antennal club. It would be very
interesting to determine whether this otherwise
ubiquitous chlamydopsine dimorphism is truly
lacking in this species. This male also possessed
small dentiform metastemal processes near the
posterior midline not observed in any other
Chlamydopsis.

Chlamydopsis detecti Lea, 1914
(Figs 22E, 23E, 27)

Chlamydopsis detecti Lea, 1914b: 215; Type: detecti Type
Lea, Dawson R./ Chlamydopsis detecti Lea Queensland
Type, 15575; SAM, examined, 2000.

DIAGNOSIS. Chlamydopsis detecti and the
following two new species form a distinctive and
closely related group. All are larger than average
for the genus, and share several characters:
strongly and continuously upturned anterior and
lateral pronotal margins bearing conspicuous
marginal setae; carinate and elevated posterior
elytral margins which continue anteriorly along
the lateral margin (in C. storeyi sp. nov. all the
way to the trichome); humeral trichomes very
prominent, dominated by rather narrow, strongly
elevated inner edges, which meet at a vertical
mesal cleft; mediobasal elytral depression large.

extending beyond middle of
elytra. Chlamydopsis matthewsi
sp. nov. is the most distinctive
of these, being deep red in
color and entirely glabrous
dorsal ly (except along elytral
and pronotal margins).
Chlamydopsis detecti and C.
storeyi are easily separated
from each other by the shape
of the pronotum (Fig. 22E vs
22F), which in the former is
proportionally shorter, and
rounded (rather than angulate)
where the lateral and anterior
margins meet. Chlamydopsis
storeyi also bears prominent
setae on the inner anterior
elevation of the humeral
trichome which are entirely
lacking in C. detecti. However,
it should be noted that C.
storeyi is quite sexually
dimorphic in surface setation
and sculpture, while C. detecti is known from
only one (undetermined) sex.

DESCRIPTION. A few additional characters of
this species are noteworthy. Body rufescent, not
yellow; Irons granulose, without reticulate
sculpture, with scattered setae; antennal scape
with few setae (or small setal bundles); pronotal
margins elevated, the anterior margin mostly
evenly rounded but shallowly notched at middle;
pronotal and elytral surfaces granulate; humeral
trichomes bare on anterior surface of inner
elevation; anterior and posterior inner elevations
rather broad, leaning slightly toward the outside
(in anterior view), arcuate around mediobasal
depression; outer elevations laterally carinate,
with ‘V'-shaped setose incision; anterior super-
ficial humeral groove very deeply impressed,
close to lateral margin, edges bare; mediobasal
depression large, with rounded, setose basal
tubercles; elytra with setae along posterior half of
suture; posterior elytral margin strongly carinate,
the carina continuous forward along posterior
one-fourth of lateral margin.

REMARKS. The type specimen was collected
from a nest of ‘Iridomyrmex detectus\ which has
since been split into several species, three of
which (/. purpureus (Smith), I. sanguineus Forel,
and /. viridaeneus Viehmeyer) apparently occur
in the range of C. detecti. This species is known
only from the holotype.

REVIEW OF CHLAMYDOPSIS

219

Chlamydopsis storeyi sp. nov.

(Figs 22F, 23 F, 27)

MATERIAL. HOLOTYPE (QMT 108595) 6 : Australia:
N WA, Kununurra, 22.XI1. 199 1-5.1. 1992, R. I. Storey, in
QMB. PARATYPE 9 : same data as type.

DIAGNOSIS. See diagnosis above under C.
detecti.

DESCRIPTION. L: 3.36; W: 1 .34; E/Pn L: 1 .5 1 ;
E/Pn W: 1 .37; Pn W/L: 1 .40; E L/W: 0.79; Pr/Py:

1 .39; Sterna: 0.87, 0.22, 0.84; Tibiae: 1 .25, 1 .34,
1.37. Body light yellow-orange, with legs,
trichome apices, and most major carinae darker,
rufescent; most normal body striae exaggerated,
carinate; female with pronotum and elytral disks
largely glabrous, with setae only around margins
and toward apices of trichomes; male with pro-
notum and elytral disks sparsely but evenly setose;
frons slightly wider than long, sides rounded,
narrowed gradually to apex and abruptly at base,
anterior margin slightly rounded; disk convex,
slightly projecting at bases of antennae,
irregularly reticulopunctate and with sparse,
elongate setae; labrum semicircular, with several
setae; outer bases of mandibles reticulopunctate
and setose; antennal scapes arcuate, widest about
one-third from base, but only slightly narrowed
to rounded apex, disk coarsely punctate on outer
edges, but only faintly punctured, micro-
sculptured medially; antennal club of both male
and female 0.6 x length of scape.

Prothorax 1 .3 x as wide as median length,
sides margined, widening towards the front;
lateral and anterior margins continuously and
strongly elevated, the lateral posteriorly
diminishing in height, joining posterior margin
which is finely carinate; anterior margin sinuate,
notched at middle, subacute on either side of
notch; pronotal disk strongly depressed behind
margins, convex in posterior medial half,
shallowly punctate on inner surfaces of marginal
elevations as well as along anterior one-third of
midline, elsewhere smooth; pronotum of female
glabrous except for a few setae along lateral and
anterior margins, that of male with sparse but
conspicuous setae on entire disk, most of them
curled over, forming a loop at their apices.

Prosternum with anterior margin deflexed,
very deeply grooved, the groove joining
circumcoxal stria at sides; lower edge of marginal
groove projecting on either side of middle;
circumcoxal stria strongly carinate, these
becoming doubled by keel carinae in posterior
half; keel narrowed posteriorly, acutely

emarginate at apex; prostemal disk sparsely
punctate along anterior margin, elsewhere
impunctate.

Elytra slightly wider than pronotum basally,
wider towards apex; humeral trichomes very
large; inner edges of anterior and posterior
elevations rising vertically nearly as high as the
body is deep, their opposing edges separate
basally, converging, then slightly separated at
apices, setose on inner margins and, less densely,
on outer surfaces; outer edges of anterior and
posterior trichome elevations rising only about
one-third as high as inner edges, separated from
them by longitudinal furrow, and at middle by
nearly circular opening; outer edge of anterior
superficial groove of trichome setose, prominent,
forming arcuate lateral margin from basal elytral
comer to trichome, inner edge of this groove well
developed and setose near base, but diminishing
before reaching opening of trichome; lateral
margin of trichome with broad, V-shaped notch;
epipleurae tuberculate and faintly punctate
beneath this notch, otherwise smooth; dorsum of
elytra smooth, sparsely setose in male, glabrous
(except along lateral and, especially, posterior
margins) in female.

Mesostemum about 4x wider than median
length; margins carinate; anterior mesostemal
margin projecting at middle; mesostemal disk
depressed, rugose in anterior half (depression’s
posterior edge parallel to sinuate anterior
margin), elevated and smooth in posterior half;
mesomctasternal suture deeply impressed;
metasternum short, about 4x length of
mesostemum along midline, disk smooth, with
only very fine, sparse punctures; median
longitudinal metastemal suture visible but not
impressed; mesofemoral lines strongly carinate,
complete to, and continued on, metepistemum;
1st visible abdominal sternite smooth;
metafemoral lines strongly carinate, complete;
legs fairly short, profemur and protibia sparsely
punctate on lower surfaces, meso- and
metafemora and tibiae smooth; all tibiae fairly
broad, with outer margins angulate about
one-third from base.

Propygidium 1.3 x median length of
pygidium, both nearly flat, vertical, only very
slightly convex, with inconspicuous fine setae.

REMARKS. The dimorphism in setation of the
pronotum and elytra in this species is unique
among Chlamydopsis , although the preceding
and the following species are closely related, and
may be found to share it when both sexes of these

220

MEMOIRS OF THE QUEENSLAND MUSEUM

are known. This species is named in honor of
Ross Storey, who collected and provided a
substantial fraction of the material for this study.

Chlamvdopsis matthewsi sp. nov.

(Figs 22a 23G, 27, 29H, 30C)

MATERIAL. HOLOTYPE 9: Australia N.S.W., Sturt
Nat. Pk., 21km W Fortville, P.J.M.Greenslade, 22/11/79,
sand dune, in SAM.

DIAGNOSIS. See diagnosis under C. detecti ,
above. This is the largest species of Chlam vdopsis
known. Its size and its deep red color are highly
distinctive. The preceding species is closely
related and generally similar, but in addition to
the color difference, it possesses epipleural
tubercles and setae near the apices of the inner
upper edges of the humeral trichomes, which C.
matthewsi lacks completely (at least in the
female).

DESCRIPTION. L: 4.36; W: 1.56; E/Pn L: 1.80;
E/Pn W: 1.38; Pn W/L: 1 .54; E LAV: 0.85; Pr/Py:
1.47; Sterna: 1.06,0.31, 1.03; Tibiae: 1.46, 1.62,

1 .65. Body dark red, large; frons 1 .4X as wide as
long, sides broadly rounded, gradually narrowed
to apex, abruptly narrowed at antennal bases;
frons acutely projecting on each side medial to
antennal insertions, shallowly depressed at sides,
and elevated along anterior margin; disk sparsely
punctate, slightly rugose at sides, with sparse,
elongate setae; labrum rounded, glabrous; outer
bases of mandibles finely punctate, glabrous;
antennal scapes arcuate, widest near middle,
faintly punctate, bearing scattered conspicuous
setae, outer margin bluntly angulate; antennal
club retracted and barely visible in type.

Prothorax 1.4x as wide as long, lateral and
anterior margins strongly and continuously
elevated; anterior margin somewhat uneven,
shallowly notched at middle, bearing sparse
fringe of curled setae; outer edge of supracoxal
groove visible from above outside of dorsal
lateral margin; pronotal disk strongly depressed
at sides, convex along midline, slightly more
prominently so posteriorly; disk finely and
shallowly punctate, with a few larger punctures
towards the front.

Prostemum as in the preceding species.

Elytra 1.8 x as long as pronotum along
midline, widest at shoulders, faintly sinuate and
slightly tapering posteriorly, apical margins
nearly transverse; humeral trichomes very
prominent, the inner edges of anterior and
posterior elevations strongly raised, almost

meeting at their apices, with vertical fringes of
opposing setae; lateral to these, the trichome
opening is rounded, with an outer fringe of longer
erect setae; lateral edge of anterior elevation
delimited by the outer edge of the anterior
superficial groove, which is very deeply
impressed, somewhat undercut toward outside;
inner edge of this groove only defined in anterior
two-thirds; trichome with relatively narrow,
V-shaped notch in lateral view; epipleuron
lacking the suprafemoral tubercle of the
preceding species; elytral disk impressed in an
approximate diamond shaped area in basal
two-thirds between trichomes; posterior elytral
margin elevated and strongly carinate, this carina
bearing setae at middle, curved anteriorly at sides
but diminishing one-sixth from apex.

Mesostemum about 4 X as wide as median
length, anterior margin sinuate, roundly
projecting at middle, grooved along anterior,
lateral and posterior margins, the groove broad in
anterolateral corners; mesosternal disk very
finely punctate; central part of metastemum
delimited on all edges by fine, continuous
groove, the mesofemoral lines defining the
anterolateral boundary; mesofemoral lines
reaching metepistemum, but continued on it only
by low, blunt ridge; median longitudinal
metastemal suture complete but not strongly
impressed; metastemal disk finely punctate; 1st
visible abdominal stemite with fine, but deeply
impressed anterior marginal groove, this groove
continuous at sides with metafemoral lines,
nearly reaching epipleuron; legs as in the
preceding species.

Propygidium 1.3 x as wide as long, 1.4 X as
long as pygidium along midline, both nearly
vertical; propygidial disk slightly flat along basal
margin, otherwise slightly convex, finely punctate
and faintly rugose in basal half; pygidium
slightly concave at sides, finely punctate, and
slightly rugose throughout.

REMARKS. This species is named in honor of
Eric Matthews, who has provided valuable
material and information throughout the course
of this study.

Chlamvdopsis cavicollis Lea, 1912
(Figs 22H, 23H, 27)

Chlamvdopsis cavicollis Lea, 1912: 65; Lectotype, hereby

designated: Sydney [NSW]/ On permanent loan from

Macleay Museum University of Sydney/ Chlamydopsis

cavicollis Lea, N.S. Wales; ANIC, seen, 2000.

REVIEW OF CHLAMYDOPSIS

221

FIG 25. Dorsal views of Chlamydopsis spp. A, C. tuberculata. B, C. mareeba. C, C. parallelus.

DIAGNOSIS. While its continuous, elevated
lateral and anterior pronotal margins ally it with
the preceding three species, this species is very
distinctive. The body shape (Fig. 22H) is
unusual, with the pronotum very small relative to
the elytra, and the elytra narrowing from the
humeri to the apex. The epipleural cleft of the
trichome is unique, forming a long, narrow,
posteriorly curving incision. Additional unusual
characters include: anterior and lateral pronotal
margins lined with prominent setal bundles;
anterior and posterior elevations of humeral
trichomes each with two separate bundles of
setae; posteromedial elevation of trichome only
weakly elevated; inner edge of anterior humeral
groove not well developed, the outer prominent,
scooplike; marginal epipleural carina not arcuate
over metathoracic leg; legs elongate and very
slender; elytral dorsum and pygidia with
numerous discal setae.

REMARKS. This species is known only from the
holotype, which has suffered some dermestid
damage, and is missing most of its legs.

INCERTAE SEDIS

The following three species are not obviously
related to any of the preceding species groups and
are left unafTiliatcd.

Chlamydopsis tuberculata Lea, 1912
(Figs 231, 25A, 27)

Chlamydopsis tuberculata Lea, 1912: 54; Type (9):
tuberculata Lea Type, Ballarat/ 14670, Chlamydopsis
tuberculata Lea, Victoria, mounted with separate card,

originally with two host individuals, only part of one
individual remaining; SAM; seen 2000.

MATERIAL. MCZ: Vic: Lome, x. 1918. Lea
(1919): Vic: Lome, ‘with a small black species of
Iridomyrmex \ Oke (1923): Vic: Grampians; Vic:
Macedon.

DIAGNOSIS. Chlamydopsis tuberculata is
easily separated from all other Chlamydopsis by
the pronotum. It not only bears a prominent,
blunt, transverse tubercle, but is more strongly
and continuously elevated along anterior and
lateral margins than any other species. The shape
of the pronotum, widening posteriorly (in dorsal
view), is also distinctive. Additional dis-
tinguishing characters include the position of the
humeral trichomes, situated very close to the
anterolateral elytral comers, the very shallow,
indistinct reticulation of the elytra, and the rather
slender but marginally rounded tibiae.

REMARKS. With regards to this species’ phylo-
genetic affiinities, one of the more significant
characters is the lack of marginal prostemal
groove (plesiomorphy), which excludes it from
the epipleuralis and ectatommae groups. There is
some similarity in trichome with species in the
striatipennis group. However, subtle differences
have prevented the recognition of these as
potential synapomorphies. No genitalia have
been examined due to the rarity of specimens.
The ovipositor will likely reveal additional clues
as to the relationships of the species.

222

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 26. Lateral views of Chlamydopsis spp. A, C. mareeba. B, C. parallelus.

Chlamydopsis mareeba sp. nov.

(Figs 25B, 26A, 27)

MATERIAL. HOLOTYPE (QMT 108596) 9 : 6km SE of
Mareeba, Qld., 1 6.xii-20.i. 1991, S.DeFaveri, F.I.T. site 36.

DIAGNOSIS. This species, although difficult to
place phylogenetically, is very distinctive. The
combination of prosternal marginal groove,
deeply transversely incised humeral trichome
bearing near continuous setal fringe, and
impunctate, plurisetose elytra are sufficient to
separate it from other known species. It should be
noted, however, that the unknown male may not
conform to this diagnosis.

DESCRIPTION. L: 2.55; W: 0.93; E/Pn L: 1 .73;
E/Pn W: 1.18; Pn W/L: 1.63; E L/W: 0.90; Pr/Py:
1.16; Sterna: 0.65, 0.19, 0.69;

Tibiae: 0.87, 1.00, 1.03. Body
rufescent, quadrate, dorsal
surface mostly impuncate,
with numerous elongate setal
bundles; frons about as long as
wide, sides rounded, disk
uniformly convex, with broad
shallow punctures slightly
separated by faintly
microsculptured areas, w r ith a
couple small setal bundles;
labrum rounded, weakly
punctate, glabrous; antennal
scape bluntly angulate near its
outer midpoint, abruptly
narrowed to base, more
gradually to apex; surface of
scape with elongate, shallowly
impressed punctures, a few
setal bundles; antennal club of
female about two-thirds length
of scape.

Pronotum 1.5 x as wide as median length;
sides margined, widened and slightly elevated
towards front; anterior margin elevated, median
and lateral portions continuous, and continuous
with lateral margins; disk depressed behind
anterior margin, convex posteriorly, nearly
smooth at middle, very faintly reticulopunctate at
sides and front; anterior and lateral margins, and
to a lesser extent the disk with conspicuous
punctures bearing bundles of elongate setae.

Prosternum with anterior margin deeply
grooved, sinuate and acutely projecting on either
side, marginal groove curving posteriorly at sides
and merging with circumcoxal stria; prosternal
keel narrowed posteriorly, widening slightly
behind procoxae, apex bluntly emarginae, with

FIG 27. Collecting records for species of Chlamydopsis epipleuralis group
and unplaced species.

REVIEW OF CHLAMYDOPSIS

223

deeply impressed marginal stria along edges of
leg depressions and posterior margin of keel;
prostemal disk with shallow punctures separated
by about their widths, faintly alutaceous
between, with sparse, irregularly scattered
bundles of setae.

strigicollis group above, the vertical superficial
humeral groove and deeply grooved anterior
prostemal margin of this species argue against a
close relationship. Discovery of the male would
be very helpful in placing it. Its name refers to the
Queensland town close to the type locality.

Elytra together about 1 .2 x as wide as base of
pronotum, more or less parallel-sided, narrowed
more gradually to apex than to base; humeri
strongly, narrowly elevated, the posterior
elevation extending back about two-thirds from
the elytral base; elevations deeply incised
transversely by trichome opening, setal fringe
consisting of anteriorly and posteriorly directed
tufts from inner edges of opening, these barely
discontinuous with outer fringe, which extends
from apices of elevations continuously around
lateral incision; apices of anterior and posterior
elevations both emarginate above opening, with
the setal fringe following emargination; anterior
elevation strongly convex, its anterior surface
nearly vertical, anterior superficial groove rather
shallowly impressed, extending from humeral
comer slightly inwardly to apical emargination
of anterior elevation; mediobasal depression
large, with transverse carinae, from suture
arching posteriorly to beneath trichome; dorsal
portion of elytral disk almost entirely impunctate,
with only faint punctures posteriorly, with
numerous elongate setae, most in bundles of 2-4
setae; epipleuron strigose, with strigae converg-
ing to trichome opening.

Mesostemum about 4x as wide as median
length; slightly elevated along midline,
depressed at anterior comers; texture like that of
prostemum; mesometastemal suture deeply im-
pressed, continuous with postmesocoxal groove;
longitudinal metasternal stria not impressed,
barely detectable; metstemal disk with alutaceous
microsculpture, with sparse, deep setigerous
punctures, otherwise impunctate; 1 st abdominal
stemite with denser, but smaller, punctures than
metastemum, most setae single; legs short,
slender with sparse, elongate punctures inter-
spersed with smaller setigerous ones, most setae
in bundles; pro- and mesotibiae acutely angulate
near basal one-third, metatibia more rounded.

Propygidium convex, with slightly elongate
punctures separated by about their widths
interspersed with sparser deeper punctures bearing
bundles of setae; pygidium similarly textured in
basal half, punctures fewer in apical half.

REMARKS. Although superficially similar to
the females of C. monteithi and C. setifera in the

Chlamydopsis parallelus sp. nov.
(Figs 25C, 26B, 27)

MATERIAL. HOLOTYPE (QMT 108597): 15.1 IS
143.52E Hann River Qld 15 Sep - 20 Oct 1993 Flight
Intercept Trap P.Zborowski & D.Rentz. PARATYPE 9 :
SEQ: 25°13’S 149^0 l'E, Expedition Ra. Nat. Pk., 5729
Amphitheatre yards, 440m 19.xii.97-4.iii.1998, DC&GM,
open for. FIT.

DIAGNOSIS. The elongate body form is very
distinctive. Its body length is slightly over twice
its width across the humeri, and the median
pronotal length is just about equal to its basal
width. In characters of phylogenetic significance,
however, the species is not particularly remarkable.
The anterior marginal prostemal groove is well
developed, diverging to meet the circumcoxal
stria at the sides. The humeral trichomes are
moderately well developed, with laterally
discontinuous, otherwise nearly circular setal
fringe. The carinae of the mediobasal depression
are unusual, forming laminate transverse peaks,
rising steeply from along the elytral suture behind
the scutellum, but extending only about
two-thirds of the way to the humeral trichome.

DESCRIPTION. L: 1.84; W: .065; E/Pn L: 1.81;
E/PnW: l.25;Pn W/L: 1.14; EL/W: 1.27; Pr/Py:
1.36; Sterna: 0.50, 0.12, 0.44; Tibiae: 0.47, 0.50,
0.53' Body narrow, elongate, light rufescent
brown; frons about as wide as long, weakly
convex, sides rounded, disk shallowly punctate,
the punctures larger and more conspicuous nearer
the anterior frontal margin, appearing granulate
within each puncture; anterior frontal margin
slightly outwardly arcuate, labrum rounded, with
a few small punctures; outer margin of antennal
scape bluntly angulate just basad of midpoint,
abruptly narrowed basally, more gradually to
apex, surface microsculptured but impunctate,
very finely setose; antennal club of female
two-thirds length of scape, sclerotised over much
of its surface, with only a couple small tomentose
patches on outer surface near apex.

Pronotal median length equal to basal width,
sides unmargined, inwardly arcuate, similar in
width basally and apically, narrowest about
one-third from front; anterior margin weakly
elevated, very shallowly emarginate across

224

MEMOIRS OF THE QUEENSLAND MUSEUM

middle, more deeply so above antennal cavities;
disk depressed in anterior corners, strongly
convex along midline, somewhat transversely
elevated in posterolateral comers; disk shallowly,
but more or less evenly punctate, the punctures
toward the sides slightly elongated.

Prosternum with anterior margin deeply
grooved, sinuate and acutely projecting on either
side, marginal groove curving posteriorly at sides
and merging with circumcoxal stria: prostemal
keel narrowed posteriorly, widening slightly
behind procoxae, apex weakly emarginate, with
fine marginal stria; prostemal disk with shallow
punctures separated by slightly less than their
widths, with very fine interspersed setae.

Elytra 1 .2 x as wide as pronotum, with sides
approximately parallel, though slightly sinuate at
middle; humeral trichomes moderately elevated,
close to humeral comers, trichome broadly open
dorsal ly, nearly circular, slightly wider than long,
fringed with short, dense setae which do not
completely obscure the opening, this fringe
briefly interrupted at sides, and, weakly, at the
junction with the anterior superficial humeral
groove; humeral groove well impressed, its edges
rounded, extending from humeri to inner apex of
trichome; mediobasal elytral depression
confined to basal one-third, w r ith prominent,
slightly oblique transverse basal carinae, these
bluntly triangular, laminate; elytral disk evenly
convex in apical half, apical margin not carinate;
disk shallowly and sparsely strigose, with
scattered fine setae.

Mesostemum about 3x as wide as median
length, anteriorly weakly projecting, with faintly
impressed lateral and anterior marginal stria, disk
with large shallow punctures, and smaller ones
interspersed; mesometasternal stria shallowly
impressed; longitudinal metastemal suture very
fine, barely visible; metastemal disk finely and
sparsely punctate, with scattered minute setae;
1st abdominal sternite similarly textured to
metasternum. Legs short, broad; meso- and
metatibiae nearly half as wide as long; protibia
bluntly angulate just beyond basal one-third;
outer margins of posterior tibiae, only faintly
angulate, very nearly rounded; all legs impunc-
tate, with fine setae; tarsi laterally compressed.

Propygidium strongly convex, faintly alutaceous,
with sparse, shallow punctures, fine setae;
pygidium weakly convex, faintly alutaceous,
with weak punctures only in basal one-third.

REMARKS. One specimen from Western
Australia ‘14.26S, 126.38E, CALM Site 13/4

12km S of Kalumburu Mission’ [ANIC] is
closely related to this species, and possibly the
same. However, its body is broader and less
elongate. Given the geographic distance, it seems
likely that this difference will be bridged by
intervening forms. Its status will need to
reassessed when additional material can be
studied. The name of this species refers to its
elongate, nearly parallel-sided body form.

SUMMARY OF KNOWN SPECIES AND
SPECIES GROUPS OF CHLAMYDOPSIS

striatipennis group

1 . C. striatipennis Lea

2. C. leai Oke

3. C. compress ipes Lea

4. C. pallida Lea

5. C. rana sp. nov.

6. C. antennata sp. nov.

7. C. trichonota sp. nov.
strigicollis group

8. C. reticulata Lea

9. C. dimorpha sp. nov.

1 0. C. strigicollis Oke

1 1 . C. mormolyce Lea

12. C. monteithi sp. nov.

13. C. setifera sp. nov.

14. C. lawrencei sp. nov.
pygidialis group

15. C. pygidialis Blackburn

1 6. C. carinicollis Lea

1 7. C. serricollis Lea

1 8. C. setipennis Oke

19. C. convergens sp. nov.

20. C. coronis sp. nov.

21. C. erupta sp. nov.

22. C transfers a sp. nov.
longipes group

23. C. longipes Lea

24. C. inaequalis Blackburn

25. C. agilis Lea
ectatommae group

26. C. ectatommae Lea

27. C. kununurra sp. nov.

28. C. acutricha sp. nov.

29. C. myrmecophila sp. nov.

30. C. variolosa Lea

3 1 . C. mallee sp. nov.

32. C. pecki sp. nov.

33. C. loculosa Lea

34. C. degallieri sp. nov.

35. C. papuae Lewis

36. C. jayawijaya sp. nov.

37. C. lucifer sp. nov.

38. C. bataviae sp. nov.

REVIEW OF CHLAMYDOPSIS

225

39. C. burnetta sp. nov.

40. C. zborowskii sp. nov.

41. C. pluriseta sp. nov.

42. C. contorta sp. nov.

43. C. pilosipes sp. nov.

44. C. bifovaecollis (Oke)

45. C. nielseni sp. nov.

46. C. australis sp. nov.

47. C. lepida sp. nov.

epipleuralis group

48. C. epipleuralis Lea

49. C. sculptus Oke

50. C. convexa sp. nov.

5 1 . C. striatella Westwood

52. C. formicicola (King)

53. C. dispersa sp. nov.

54. C. weiri sp. nov.

55. C. crowcrofti sp. nov.

56. C. latipes Lea

57. C. macmillani sp. nov.

58. C. nu llarbor sp. nov.

59. C. rotunda sp. nov.

60. C. laiipennis Lea

6 1 . C. carinota sp. nov.

62. C. inquilina Lewis

63. C. detecti Lea

64. C. storeyi sp. nov.

65. C. matthewsi sp. nov.

66. C. cavicollis Lea

incertae scdis

67. C. tuberculata Lea

68. C. mareeba sp. nov.

69. C. parallelus sp. nov.

PHYLOGENY

This analysis is based on external morph-
ological variation as well as some characters of
the ovipositor. While some variation in male
genitalia has been observed (primarily in setal
patterns and apical curvature), insufficient
dissectable males were available to include any
male characters in this study. The matrix of
female genitalic characters does contain
substantial missing data, but the states were much
more distinctive and obviously informative. This
analysis includes all known species of Chlamy -
dopsis , including one recently discovered New
Caledonian species, to be formally described
elsewhere. The outgroups represent a broad
selection of other Chlamydopsinae, as well as
three non-chlamydopsine histerids. These are:
Stictostix frontalis Lea (Tribalinae), a new
Malaysian genus near Peploglyptus (Onthophilinae;
description in preparation), Onthophilus flohri
Lewis (Onthophilinae) and the chlamydopsines

Pheidoliphila minuta Lea, Orectoscelis dumogae
Caterino, Orectoscelis obliquus Caterino,
Ceratohister sp., Eucurtia comata (Blackburn),
Ectatommiphila opaca (Lea), representatives of
two undescribed genera close to Orectoscelis ,
and a representative of one undescribed genus of
obscure affinities. The complete data set contains
82 taxa and 46 characters.

CHARACTERS.

1. Frons: /. without prominent tubercles; 2. with single broad
protuberance; 3. with multiple (usually 6) protuberances.

2. Labrum, apical margin: /. evenly rounded, nearly
semicircular; 2. broader, nearly straight for some distance
at middle.

3. Antennal insertion: 1. at the middle of the frons; 2. at the top
of the frons near the top of the eye. This character is
generally considered the defining synapomorphy of
Chlamydopsinae.

4. Antennal club: /. identical in length in both sexes, generally
less than twice as long as wide; 2. more elongate in 6,
usually three or more times as long as wide. The strongly
dimorphic antennal club, with that of the 6 extremely
elongated, is nearly ubiquitous in Chlamydopsinae.
However, in one species, a 6 with a short antennal club has
been seen. The small number of specimens examined
leaves some doubt that this represents more than an
aberration.

5. Medial portion of anterior pronotal margin: /. flat, neither
thickened nor elevated; 2. distinctly elevated, with some
anteriorly exposed surface below margin.

6. Anterior pronotal margin: /. median transverse and lateral
oblique portions continuous; 2. interrupted between
median and lateral portions, either by notch or by a carina.
State two encompasses considerable diversity that could
perhaps be more finely divided. In some of these species
the median segment of the anterior pronotal margin (that
portion above the vertex of the head) is elevated separately
from the lateral portions (those above the antennal
cavities). In others the median and lateral portions meet,
but the inner apex of the lateral portion extends for a very
short distance anteriorly beyond the median portion.
Grouping these conditions together as a state suggests that
the origin of discontinuity, in whatever form, was the
significant change, with diversity arising after.

7 Pronotum: 1. without a stria behind elevated lateral portion
* of anterior margin; 2. with an oblique stria extending along
posterior base of lateral portion of (usually) elevated
anterior margin. This stria, when present, extends from the
lateral pronotal margin, behind lateral portion of anterior
margin, to the anterior margin between its lateral and
median portions.

8. Anterior pronotal margin, median portion: /. even at
middle, whether elevated or not; 2. notched at middle.

9. Pronotum: /. pronotum margined laterally: 2, pronotum
without lateral margin.

10. Pronotum: /. lateral margins flat, not elevated; 2. lateral
margins elevated, angulately continuous with lateral
portions of anterior margin; 3. lateral and anterior margins
indistinguishable, forming a single oblique carina from
anterior midpoint to each posterolateral pronotal comers.
State three applies only to single unusual species from New
Guinea (new genus3). The relationships of this species are
unclear but it appears to be outside of Chlamydopsis.

226

MEMOIRS OF THE QUEENSLAND MUSEUM

11. Pronotal trichomes: 7. absent; 2. present (Figs 1E-G).
Pronotal trichomes occur in only a few Chlamydopsis ,
and, although somewhat similar, they are not identical in
form in those species possessing them. Nonetheless, due
to their rarity, they are considered potentially
homologous where they occur.

12. Anterior marginal or near marginal processes of
pronotal disk: 7. absent; 2. present as two distinct
swellings (or single bifid process) at or near anterior
margin. This character as scored mainly separates
Chlamydopsis from chlamydopsine outgroups, most of
which possess some form of paired marginal pronotal
processes.

13. Single median pronotal tubercle: 7. absent; 2. present.
This character had previously been combined with the
preceding character. However, although not observed it
appears possible for anterior marginal and median
processes to co-occur.

14. Median pronotal tubercle: 7. absent; 2. simple; 3.
expanded laterally to form transverse carina. This
character is dependent on the presence of a pronotal
tubercle in the preceding character.

15. Pronotum: 7. without longitudinal carina; 2. with
longitudinal carina along midline in anterior half
(sometimes also with posterolateral extensions).

16. Anterolateral groove from antennal cavity to
pronotum; 7. absent; 2. present, simple, merging with
prostemal circumcoxal stria at side (Fig. 28D). 3.
present and leading to dorsal pronotal pit (Fig. 29A).
State 2 indicates a groove running from the upper edge
of the antennal cavity upward, intersecting the lateral
portion of the anterior pronotal margin, thence
extending posterolateral ly to the lateral pronotal margin
below which it meets the circumcoxal stria. The groove
exhibited in state 3 is potentially homologous with this
one (although this is not asserted by the present coding
scheme). However, in those taxa exhibiting state 3 the
groove extends straight back from the antennal cavity
and terminates in conspicuous dorsal pronotal pits. This
state is diagnostic of the bifovaeeollis subgroup of the
ectatommae group.

17. Prostemal disk: 7. length anterior to profemoral carina
less than that posterior to carina; 2. length anterior to
carina equal to or greater than length posterior to carina.
State 2 is characteristic of a large section of
non-Chlamydopsis chlamydopsines.

18. Anterior marginal stria of prostemal lobe: 7. not deeply
grooved (Fig. 28A); 2. deeply grooved, the groove

- running along margin all the way to side, not continuous
with the circumcoxal stria (Fig. 28B); 3. deeply
grooved, departing from margin at sides and curving
posteriorly to meet the circumcoxal stria (Fig. 28C).
This character has proven one of the most useful for
sorting out preliminary affinities within Chlamydopsis.
These grooves would appear to constitute some kind of
conduit system on the beetles’ surface. In several species
these grooves meet elytral grooves to form a continuous
series extending all the way to the humeral elytral
trichomes. Perhaps these serve to disperse recognition
substances from a site of origin to elsewhere on the body.

19. Prostemal disk: 7. without transverse stria behind
anterior margin: 2. Prostemum with transverse stria just
behind anterior margin. This stria, when present, is
reminiscent of the ‘presternal stria' of many
non-chlamydopsine Histeridae, and may conceivably be
homologous, it being present in a couple of the
outgroups.

FIG 28. A-C, Ventral view of left half of prostemum of
Chlamydopsis spp. A, C. striatipennis. B, C.
monteithi. C, C. burnetta. D, Dorsal view of left half
of pronotum of C. kununurra , showing anterolateral
groove.

20. Scutellum: 7. visible dorsally; 2. hidden. This character
is informative only with respect to outgroups.

21. Humeral elytral trichome: 7. absent; 2. present. This
and the following several characters refer to the
structure of the humeral trichome, or ‘epaulette’ of most
previous authors. This structure is extremely varied, and
unquestionably informative at some levels. However, it
is very difficult to determine homologies among the
various components. Some of the characters below are
admittedly interdependent, and other scoring schemes
could be justified, but no wholly satisfactory schemes
have yet been found.

22. Position of humeral trichome: 7. absent; 2. behind
humeri; 3. limited to humeri; 4. trichome largely
posthumeral, but extending forward mesally to anterior
elytral margin or even to humerus. The position of the
trichome is primarily informative with respect to
non-Chlamvdopsis outgroups. However, in a few
Chlamydopsis the degree to which the trichome extends
forward toward the anterior elytral margin may be
informative.

23. Anterior superficial groove of trichome: 7. with
anterior groove up the middle of the anterior elevation,
approximately evenly dividing it into inner and outer
prominences (Figs 29A-B, E-G); 2. with anterior groove
oblique or horizontal, entering the lumen of the trichome
at its mesal base (Figs 29C-D); 3. without any visible
groove on anterior elevation of trichome (Fig. 291); 4.
with anterior groove displaced laterally, forming a
lateral margin of the anterior elytral comer (Fig. 29H).
This groove, nearly always present in Chlamydopsis , is
quite varied in form. Its absence in a few species is
almost certainly due to loss, as in most cases it can be
seen in apparent relatives. The orientation of this
groove, when present, varies considerably. It may
appear almost perfectly longitudinal, forming a
marginal groove and entering the trichome laterally (e.g.

C. detecti and relatives). Or it may take the opposite
extreme, being directed strongly medially, in some cases

REVIEW OF CHLAMYDOPSIS

227

FIG 29. Humeral trichomes of right elytron of Chlamydopsis spp. A, C.
bifovaecollis, also showing pronotal pit. B, C my nnecophila. C, C.
inaequalis. D, C. reticulata. E. C. burnetta. F, C. nullarbor. G C.
dispersa. H, C. matthewsi. I, C. sp. nr. striatipenms.

lying nearly parallel to the anterior
elytral margin (e.g. C. mormolyce and
other members of the strigicollis and
pygidialis groups), entering the
opening of the trichome from its inner
side. These changes of position may be
viewed as the results of differential
development on either side of a
(putatively) symmetric plesiomorphic
state (state 1 ).

24. Setal fringe of humeral tnchome: /. as
a single continuous marginal fringe,
usually encircling fairly simple
trichome opening (e.g. C. bifovaecollis;

Figs 29A-C); 2. with anterior marginal
and posterior marginal fringes,
discontinuous laterally and mesally
(Figs 29D-E); 3. with three distinct
origins, semicircular fringe along outer
edge separated from anterior and
posterior inner bunches of setae (Fig.

29F-H): 4. tnchome setae arising from
two origins, one in an elongate (usually
sinuate) fringe along the inner edge of
the dorsal ‘roof of the trichome, and
one within the small mesal opening of
the trichome, mostly hidden by the
dorsal fringe (Fig. 291; inner fringe not
visible in Figure); 5. with a single small
setal origin on the mesal surface of
incurved anterolateral trichome
elevation (e.g. C. pallida ); 6. trichome
absent, character inapplicable.

Although this character focuses on the
origin points of the trichome setae, it in
fact captures much of the variation in
overall shape of the tnchome itself.

25. Inner edges of anterior and posterior
trichome elevations: /. well developed
but not meeting, the lumen of the
trichome open to mediobasal
depression (Figs 29A-D, I) ; 2. well
developed and nearly or fully meeting,
closing trichome mesally (full closure
may be achieved by setal fringes on
their opposing surfaces; Figs 29E-H);

3. inner edges joined by a thin lamina,
closing trichome mesally (e.g. C.
pallida ); 4. trichome unelevated or
absent, character inapplicable.

26. Outer edges of anterior and posterior trichome elevations:
/. not closing the trichome laterally; 2. meeting, closing
trichome laterally (a notch may be present as long as it is
not continuous with trichome lumen as in, e.g. C.
myrmecophila ); 3. trichome unelevated or absent,
character inapplicable. Sexual dimorphism is responsible
for the one scored polymorphism in this character (in C.
mallee). Other species, once both sexes are discovered,
may need to be rescored for this character.

27. Trichome lumen: 1. lumen (central cavity) broadly open
dorsal ly; 2. lumen covered dorsally, with only a small
mesal opening leading to internal cavity (detectable via a
‘window’ of thin cuticle on trichome ’s lateral surface; see,
e.g. C. striatipenms; Figs 29C, I); 3. trichome absent,
character inapplicable.

28. Carinae of midbasal elytral depression: /. absent,
depression flat (or not depressed); 2. with transverse (in
some slightly arcuate) transverse carinae.

29 Carinae of midbasal elytral depression: 1. absent; 2. bare
or sparsely setose; 3. bearing dense bundles of setae (Fig.
29E). This character refers to a specialisation of the preceding.

30. Elytral marginal stria: I. continuous along elytral suture;
2. absent at least along suture.

31. Elytron: /. apical margin flat; 2. with apical marginal
carina (which is separate from marginal stria); 3. with
apical marginal carina extending forward along lateral
edge to humeral trichome.

32. Elytra of 6 and 9 : /. identical in surface texture; 2.
differing substantially in texture such that females are
smooth and males are reticulostrigose.

33. Prostcrnal/mesosternal junction: /. mesosternum
projecting, prosternum emarginate; 2. prosternum
posteriorly truncate to rounded, projecting over anterior
margin of mesosternum.

228

MEMOIRS OF THE QUEENSLAND MUSEUM

34. Meso- and metatibiae: 7. without grooves for retraction
of tarsi; 2. with grooves for retraction of tarsi. This
character is only informative with respect to outgroups.

35. Ventral cavities for retraction of legs: /. poorly
developed, not margined all the way around; 2. well
developed, completely delimited by carinae (femoral
lines). This character is related to character 38, below, in
that elongate legs tend to be accompanied by loss of
well-defined cavities.

36. Mesofemur: 7. not clavate, less than twice as thick at
apex as at base; 2. clavate, twice as thick or more at apex
than base. This and the following character, as defined,
pertain mainly to the longipes group, although there is a
tendency towards the femora becoming clavate with
increasing length throughout the group.

37. Metafemur: 7. not clavate, less than twice as thick at
apex as at base; 2. clavate, twice as thick or more at apex
than base.

38. Mctatibia: 7. short, slender, outer margin angulatc; 2.
short, broad, outer margin angulate; 3. short, broad,
outer margin rounded; 4. elongate (in practical terms,
extending above epipleuron when held vertically); 5.
short, slender, outer margins even.

39. Mctatibia, if elongate: 7. not elongate; 2. broad,
angulate; 5. broad, rounded; 4. slender

40. Tarsal claws: 7. evenly curving to apex (particularly
note inner edge); 2. inner edge straight for
approximately basal two-thirds, then curved just at
apex; 3. short, nearly perpendicularly bent at base, then
straight in apical two-thirds; 4. long, thin, nearly straight
througout.

41. Propygidium: 7. flat to convex, at least in 6:2. with
transverse carina or series of protuberances in both
sexes.

42. Body setae: 7. simple; 2. scale-like; 3. absent; 4. simple,
in bundles. There is considerably more variation in setal
types in chlamydopsine outgroups. Within
Chlamydopsis this character is mainly informative with
respect to whether the simple setae are borne singly or in
‘bundles’ of multiple setae.

43. Gonocoxite of ovipositor: 7. less than half as long as
valvifer (Figs 30A, C); 2. coxite half or more as long as
valvifer (Fig. 30B).

44. Gonocoxite of ovipositor: 7. apex bidentate (Figs
30A-B); 2. apex untoothed, simple, scooplike (Fig. 30C).

45. Gonostyle of ovipositor: 7. small, linear, not projecting
beyond apex of coxite (Fig. 30A,C); 2. elongate,
frequently clavate, projecting bevond apex of coxite
(Fig. 30B).

46. Struts of $ 9th stemite: 7. divergent towards base; 2.
convergent towards base.

PHYLOGENETIC METHODS. Analysis of this
large dataset relied heavily on heuristic
parsimony approaches. PAUP* (Swofford, 1998)
was used for all tree searches, with characters
treated as unordered throughout. An initial search
saved no more than 50 shortest trees for each of
500 random taxon addition replicates. Restricting
tree number and increasing addition replicates
allowed a substantial amount of treespace to be
explored preliminarily. This restricted search
resulted in 800 trees (i.e. 16 of the 500 replicates

A

FIG 30. Valvifers and gonocoxites of Chlamydopsis
spp. A, C. epipleuralis. B, C. rana. C, C. matthewsi.

found trees of equal, shortest length), which were
then used as the basis of unrestricted branch-
swapping. This unrestricted search was allowed
to proceed for several hours, but was terminated
due to memory restrictions when trees in memory
hit 76,000, with nearly 20,000 trees still to swap.
No trees shorter than the initial 800 were found.

The smaller set of 800 trees was used as the
basis for character reweighting, according to
character rescaled consistency indices. A single
reweighted search was carried out, with the
restriction of 50 trees described above implemented.
Although for both equally weighted and reweight-
ed searches additional equally parsimonious
arrangements might lead to a slight reduction in
resolution of the consensus trees presented here,
this search strategy should ensure that no shorter
trees will be found. Decay indices were
calculated ( with all characters weighted equally)
using the program TreeRot (Sorenson, 1999).

RESULTS. Two trees are presented from these
analyses, the strict consensus of 76,000 trees
resulting from the equally weighted, unrestricted
search (C.I. = 0.2906, R.I. = 0.7593; Fig. 3 1 ), and
the strict consensus of 1 0762 trees resulting from
the reweighted, unrestricted search (Fig. 32)
These two trees offer a relatively consistent picture
of the broadest relationships in Chlamydopsis. At
the basalmost levels of the tree, monophyly of
Chlamydopsinae is strongly supported (6 decay
steps). Within Chlamydopsinae a clade comprising
Chlamydopsis , Eucurtia , and Ectatommiphila is
supported by 2 decay steps, with an unusual, as
yet undescribed, species from New Guinea as its
sister group. Monophyly of Chlamydopsis itself
is supported by a single decay step, with
Ectatommiphila as its sister group. Relationships
within Chlamydopsis are mostly supported by

REVIEW OF CHLAMYDOPSIS

229

TABLE 2. Character state data and consistency indices (calculated over the unweighted tree topology), a = 1 ,2;
B — 1,3.

Character

1

2

3

4

5

6

7

8

9

10 11 12

13

14

15

16

17 18 19 20 21

22 23 24 25 26 27 28 29 30 31

32 33 34 35 36 37 38 39 40 41 42 43 44 45 46

C.I.

8

r~

00 1

o

•r

o

-t

8 8

o o rs

e * wi

o

8 8

£

9

m

o

v,

£

s

©

»-■

5

© ©

•O

sO

04

o

04

"»

o

8 8

2 8 8

o*

IQ

n

©

Pi

"

~

-

d

o

o

o

d

d

o

o

o

d

d

o

© d

o

©

o

o

©

c

d

— d — ©

d

o

o

©

d

d

©

Stic tost ix frontalis

i

1

2

l

1

i

1

1

1 1 1

1

1

1

1

1

1

2

1

1

1

3

6

4 3

3

l

1

2

i

1

!

|

1 1 1

5

|

1

nr Peploglyptus

i

2

2

7

1

i

1

1

1 2 2

1

1

1

1

1

1

?

1

1

1

3

6

4 3

3

l

1

2

i

1

1

1

1 1 1

5

1

4

1

2

1

|

?

Onthophilus flohn

i

a

1

1

1

i

1

1

1 1 1

1

1

1

1

1

2

1

1

1

3

6

4 3

3

1

1

2

I

l

I

1

1 1 1

5

1

l

1

3

1

3

7

new genus 1

i

1

2

?

1

i

l

2

1 1 1

1

1

1

1

2

1

1

2

1

1

3

6

4 3

3

1

1

l

l

2

2

2 1 1

3

1

1

1

I

?

9

?

9

new genus 2

i

?

2

?

l

i

l

2

1 1 2

1

1

1

1

2

1

1

2

1

1

3

6

4 3

3

1

1

l

l

2

2

2 1 1

7

9

9

1

1

?

?

?

?

Phridoltphila minuta

i

1

2

2

1

i

1

2

1 1 2

|

1

1

1

2

1

1

2

2

3

3

3

4 1

1

1

1

l

1

2

2

2 1 1

2

1

1

1

2

1

I

7

Orectoscelis dumogae

i

1

2

2

1

i

I

2

1 1 2

I

1

1

1

2

1

1

2

2

3

3

3

4 1

1

1

1

l

1

2

2

2 1 1

2

1

1

1

3

?

?

?

?

Orectoscelu obliquus

i

1

2

2

1

i

1

2

1 1 2

1

1

1

1

2

1

1

2

2

3

l

2

2 1

1

1

I

1

1

?

2

2 1 1

2

1

1

1

1

9

9

?

?

Ceratohister n . sp

i

1

2

2

1

l

1

2

1 1 2

1

1

1

1

2

1

1

2

1

1

3

6

4 3

3

1

1

I

1

2

2

2 1 1

2

1

1

2

9

7

?

7

new gmus 3

i

1

2

2

I

2

i

1

1

3 1 1

2

2

1

1

1

1

1

2

2

2

1

1

2 1

1

1

1

1

1

2

2 1 1

2

1

1

1

7

?

?

9

Eucurtia comata

i

?

2

?

1

i

1

1

2 1 1

l

1

1

!

1

1

I

1

2

2

1

2

2 1

1

1

l

2

i

1

1

2

2 1 1

2

1

7

1

1

7

7

?

9

Ectatommiphila opaca 1

1

2

2

2

i

1

1

1 1 1

1

1

1

1

1

1

1

1

2

2

I

2

4 3

1

1

1

2

3

1

1

2

2 1 I

2

1

7

1

1

1

1

2

?

CM . striatipennis

i

2

2

2

2

2

2

1

1

1 1 1

1

1

1

1

1

1

1

1

3

4

3

4

1 2

2

2

2

2

1

1

1

2

1 1 1

4

4

2

1

1

7

7

7

7

Chi . leai

i

2

7

2

2

7

7

l

1 1 1

?

2

1

I

1

1

7

1

2

2

3

4

1 2

2

2

2

2

1

1

1

2

1 1 1

2

1

9

1

1

2

1

2

7

Chi . compressors

i

2

2

2

2

2

2

1

1

1 1 1

2

2

1

1

1

1

1

1

2

2

1

4

3 2

2

2

2

1

1

1

1

2

1 1 1

4

3

2

1

7

1

1

2

2

Chi pallida

i

2

2

2

2

2

2

2

1

2 1 1

2

2

1

1

1

1

1

1

2

4

3

5

3 2

2

2

2

2

1

1

2

I 1 1

4

3

2

1

1

1

1

2

2

Chi rana

i

2

2

2

2

1

2

1

1

1 2 1

1

1

1

1

1

2

1

1

2

4

1

4

1 2

2

2

2

1

1

1

1

2

1 1 1

4

3

2

1

1

2

1

2

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Chi antennata

i

2

2

3

2

1

1

1

?

1 2 1

1

1

1

1

l

2

1

1

2

4

3

4

1 2

2

2

2

2

1

9

1

2

1 1 1

4

3

1

1

1

7

9

7

?

Chi trichonota

i

2

2

3

2

1

1

1

?

1 2 1

1

1

1

1

1

2

1

1

2

4

3

4

1 2

2

2

2

2

1

7

1

2

1 1 1

4

3

1

1

1

7

9

9

?

Chi reticulata

i

2

2

2

2

2

2

2

1

1 1 1

1

1

1

1

1

1

2

1

2

2

2

2

1 1

1

1

2

2

2

I

2

1 1 1

1

2

1

1

2

1

2

2

Chi dimorpha

i

1

2

2

2

2

2

2

1

1 1 1

1

1

1

1

1

1

2

2

2

2

1 1

1

1

1

1

1

2

1

2

2 1 1

2

1

2

1

1

2

1

2

2

Chi . strigicollis

i

1

2

7

2

2

1

1

1

1 1 l

1

1

1

1

1

1

2

2

2

2

2

1 1

1

2

2

2

2

1

1

2

1 1 1

3

1

2

1

1

7

2

9

Chi mormolyrr

i

1

2

7

2

2

1

2

1

1 t 1

1

1

1

I

I

I

I

|

2

2

2

2

1 2

1

2

2

1

1

1

1

2

1 1 2

4

4

2

1

1

7

9

9

9

Chi monieithi

i

1

2

2

2

2

1

1

2

1 1 1

I

1

1

1

1

2

1

1

2

2

2

2

1 2

1

2

2

2

1

2

1

2

2 1 1

1

1

2

1

1

7

7

7

7

Chi . setifera

i

1

2

7

2

2

l

1

2

1 1 1

1

1

l

1

1

2

1

1

2

2

2

2

1 2

1

2

2

2

1

9

1

2

2 1 1

1

1

2

1

1

1

1

2

2

Chi . lawrencei

i

1

2

2

2

2

1

1

2

1 1 1

2

2

I

1

1

2

l

1

2

2

2

1

1 1

1

2

2

2

l

9

1

2

2 1 1

1

1

2

1

1

7

7

7

7

Chi pygidialis

3

7

2

2

2

2

1

2

1

2 1 1

2

3

2

1

1

1

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1

2

2

2

2

1 |

|

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1

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3

1

1

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2

1

1

2

l

2

7

7

2

Chi . cannicoHis

3

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2

2

2

2

1

2

1

2 1 1

2

3

2

1

1

1

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1

2

2

2

2

1 1

1

2

2

2

3

1

1

2

1 1 1

2

1

7

2

1

7

7

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?

Chi serrtcollis

3

?

2

2

2

2

1

2

1

2 1 1

2

3

2

1

1

1

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1

2

2

2

2

1 1

1

1

l

2

3

1

1

2

1 1 1

2

1

7

2

1

7

9

9

7

Chi setipennis

•

7

2

7

2

2

1

2

1

2 1 1

2

?

1

1

1

1

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1

2

2

2

2

1 1

1

1

1

2

2

1

1

2

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4

2

7

2

4

7

7

7

7

Chi . convergens

3

1

2

2

2

2

I

1

I

2 1 1

2

2

2

1

1

1

1

1

2

4

2

I

1 1

l

2

2

2

2

1

1

2

1 1 1

2

1

2

2

1

2

1

2

1

Chi . coroms

3

1

2

2

2

1

2

1

2 1 1

2

3

2

1

1

I

2

l

2

4

2

1

1 1

1

2

2

2

2

1

2

1 1 1

2

1

2

2

1

7

7

9

9

Chi erupta

3

1

2

2

2

2

1

2

1

2 1 1

2

3

1

1

1

1

2

1

2

4

2

1

1 1

1

2

2

2

2

1

1

2

1 l 1

2

1

2

2

1

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7

7

7

CM . transversa

3

1

2

2

2

2

I

2

1

1 1 1

2

3

2

1

1

1

1

1

2

4

2

1

1 |

1

2

2

2

2

1

1

2

1 1 1

1

1

2

2

1

7

7

7

7

Chi . longtpes

2

1

2

1

2

2

7

7

2

1 1 1

2

7

1

1

2

1

2

1

2

4

2

9

1 1

9

2

2

2

1

|

1

2

1 2 2

4

4

9

1

1

2

1

2

7

Chi inaequalts

2

1

2

7

2

2

1

2

1

1 1 1

1

1

1

1

1

2

1

2

4

2

1

1 2

2

2

2

2

2

1

2

1 2 2

4

2

2

1

1

7

7

7

9

CM . agllis

9

2

2

2

2

1

2

1

2 1 1

2

7

1

1

I

1

9

1

2

4

2

9

1 1

2

2

3

2

2

1

1

2

1 1 2

4

2

7

1

1

7

9

7

?

CM ectatommae

2

2

2

2

2

1

1

1

1 1 1

2

2

1

2

1

3

1

1

2

2

1

1

1 1

1

2

2

2

2

1

I

2

2 1 1

1

1

2

1

1

7

7

7

7

CM . kmunurra

1

2

2

2

2

1

2

1

I I 1

2

2

1

2

1

3

1

1

2

2

1

|

1 I

I

2

2

2

2

9

1

2

2 1 1

1

1

2

1

t

7

?

?

9

Chi acutricha

1

2

7

2

1

1

2

2

1 1 1

2

2

1

2

1

3

1

2

2

2

1

1 2

1

2

2

2

2

1

1

2

2 1 1

1

1

2

1

1

7

7

?

?

CM . myrmecophila

1

2

2

1

2

1

1

2

1 1 1

1

1

1

2

1

3

1

1

2

2

I

1

l 2

|

2

2

2

|

|

1

2

2 1 1

I

1

2

1

1

1

1

7

1

Chi . variolosa

1

2

2

2

1

2

I

1

1 1 1

1

1

l

2

1

3

7

1

2

2

1

1

1 2

1

2

2

2

1

2

1

2

2 1 1

1

1

2

1

1

7

7

7

7

Chi mailer

1

2

2

2

1

2

1

1 1 1

1

1

1

1

1

3

1

1

2

2

r

1

1 a

1

2

2

2

1

2

1

2

2 1 1

1

1

2

1

1

7

9

7

9

Chi peeki

1

2

2

2

1

1

a

1

1 1 1

1

1

1

2

1

3

1

1

2

2

i

1

1 2

1

2

2

2

|

2

1

2

2 1 1

1

1

2

1

1

7

9

9

?

Chi loculosa

1

2

2

2

2

1

1

1

1 1 1

1

1

1

2

1

3

1

1

2

2

i

1

1

I

|

2

|

|

2

1 1 1

4

4

3

1

1

I

1

2

1

CM . degalheri

1

2

2

2

2

1

a

1

2 1 1

1

l

1

2

1

3

1

1

2

2

i

1

1 2

1

2

2

2

1

9

1

2

1 1 1

1

1

2

1

1

9

?

7

?

Chi papuae

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2

2

2

2

7

7

1

1 1 1

2

2

1

2

1

3

1

1

2

2

i

9

1 2

I

2

2

|

|

?

1

2

1 9 9

1

9

9

1

1

9

9

9

9

Chi Jayawijaya

1

2

2

2

2

1

2

1

1 1 1

2

2

1

2

1

3

?

1

2

2

i

2

1 |

1

2

2

2

1

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1

2

1 1 1

1

1

2

1

1

9

9

7

7

Chi lunfer

1

2

2

1

2

I

2

1

1 1 1

2

2

1

2

1

3

1

1

2

2

2

1

1 2

1

I

I

2

2

I

1

2

2 I 1

1

1

2

1

1

9

?

7

7

CM . bataviae

1

2

2

1

2

1

2

1

1 1 1

2

2

1

2

1

3

1

1

2

2

2

1

1 2

1

1

1

2

2

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1

9

2 1 1

1

1

2

1

1

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?

9

CM burneua

CM . tborowskii

1

1

2

2

2

2

2

2

2

2

1

1

1

1

1

1

1 1 1

1 1 1

2

2

2

2

1

1

2

2

1

1

3

3

1

1

1

1

2

2

2

2

1

1

2

2

2 2
2 2

\

2

2

3

3

2

2

2

2

1

1

1

1

2

2

1 1 1

1 1 1

4

4

4

4

2

2

1

1

4

4

I

7

1

?

2

7

2

7

Chi pluriseta

1

2

2

2

2

1

1

1

1 1 1

2

2

1

2

1

3

1

1

2

2

1

2

2 2

1

2

3

2

2

1

I

2

1 1 1

4

4

2

1

4

7

7

?

7

Chi contoria

1

2

2

2

2

1

1

1

1 1 1

1

7

1

2

1

3

1

1

2

2

1

2

2 1

1

2

3

2

2

7

1

2

1 1 1

4

4

2

1

4

7

7

?

7

Chi pUosipes

1

2

2

2

2

1

1

1

1 1 I

2

2

1

2

I

3

1

1

2

2

1

2

2 2

1

2

3

2

2

1

1

2

1 1 1

1

1

2

1

4

7

1

2

7

CM . bifovaecollis

?

2

?

1

2

1

?

1

1 1 I

2

2

1

3

1

2

1

1

2

2

1

l

1 2

1

1

1

2

1

7

1

2

2 1 1

1

1

7

1

1

7

7

7

7

CM . ntelseni

1

2

2

2

1

I

1

1

1 1 1

2

2

1

3

1

2

1

2

2

1

1

1 2

1

1

1

2

1

7

1

2

7 1 1

3

1

1

1

1

7

7

7

7

CM australis

1

2

2

1

2

1

1

7

1 1 1

?

1

1

3

1

2

1

I

2

2

1

1

1 2

1

2

2

2

1

7

I

2

1 1 1

l

1

2

I

1

7

7

7

7

Chi tepida

1

2

2

1

2

1

1

?

1 1 1

1

1

1

3

2

1

1

2

2

1

1

1 2

1

1

1

2

1

7

1

2

1 1 1

1

1

2

1

1

7

7

7

?

Chi epipleuralis

1

2

2

2

1

1

1

1

2 1 1

1

1

1

1

3

1

1

2

2

1

3

2 1

1

2

2

2

2

1

1

2

2 1 1

1

1

1

1

1

1

1

1

7

CM srulptus

1

2

2

2

l

1

1

1

2 I 1

1

1

1

1

3

1

1

2

2

1

3

2 1

l

2

2

2

2

1

1

2

2 1 1

1

1

1

1

1

7

7

7

7

Chi convexa

1

2

2

2

1

1

2

1

2 1 1

1

1

1

1

3

1

1

2

2

7

2

2 2

1

2

2

2

2

1

1

2

2 1 1

1

1

2

1

1

7

7

7

7

CM . striatella

7

2

2

2

1

1

2

1

2 1 1

2

2

1

1

3

9

1

2

2

1

3

2 2

l

2

2

2

1

1

1

2

1 1 1

I

1

?

1

1

7

7

7

7

CM . formicicola

1

2

2

2

1

1

2

1

2 1 1

2

2

I

1

3

1

1

2

2

1

3

2 2

1

2

2

2

1

1

1

2

1 1 1

1

1

2

1

1

2

1

2

2

CM dispersa

1

2

2

2

|

1

a

1

1 1 1

2

2

1

1

3

1

1

2

2

1

3

2 2

1

2

2

2

3

1

1

2

1 1 1

1

1

2

1

1

7

7

7

7

Chi weiri

1

2

7

2

2

1

2

1

1 1 1

2

2

1

1

3

1

1

2

2

1

3

2 2

1

2

2

2

2

7

1

2

1 1 1

1

1

2

1

1

7

7

7

7

CM . crawcrofti

1

2

2

2

2

1

2

1

1 1 1

2

2

1

1

3

1

1

2

2

1

3

2 1

1

2

2

2

2

7

1

2

1 1 1

4

2

2

1

1

7

7

7

7

CM latipes

1

2

2

2

2

1

2

1

1 1 1

2

2

1

1

3

l

2

2

1

3

2 1

1

2

2

2

1

1

1

2

1 1 1

4

2

2

1

1

2

1

2

2

CM mannillani

1

2

2

2

2

1

1

1

1 1 1

2

2

1

3

1

1

2

2

1

3

2 1

1

2

2

2

2

7

1

2

1 1 1

4

2

2

1

1

7

7

7

7

Chi nuUarbor

1

2

2

2

2

1

2

1

1 1 1

2

2

1

1

3

1

1

2

2

1

3

2 1

1

2

2

2

2

7

1

2

1 1 1

4

3

2

1

1

7

7

7

7

CM . rotunda

1

2

?

2

2

|

2

1

1 1 1

2

2

1

1

3

1

1

2

2

4

3

2 1

1

2

2

2

2

1

1

2

I 1 1

1

1

2

1

1

2

1

2

2

CM . latipennis

2

2

?

2

2

1

2

1

1 1 1

2

2

1

1

3

1

1

2

2

1

3

2 1

1

2

2

2

1

1

1

2

1 1 1

4

4

2

1

1

7

7

7

7

CM . cannota

2

2

2

2

1

1

2

1

2 1 1

1

1

1

3

1

1

3

2

4

3

2 2

1

2

2

2

2

?

1

2

1 1 7

3

7

2

1

1

7

7

7

7

CM tnquilina

1

2

1

2

1

I

2

1

2 1 1

1

1

1

3

1

1

2

2

1

3

2 1

1

2

2

2

1

1

1

2

1 1 1

4

I

1

1

1

2

1

2

1

CM detect !

?

2

?

2

1

1

2

1

2 1 1

1

1

1

3

?

1

2

2

4

3

2 1

1

2

2

2

3

7

1

2

2 1 1

2

1

7

1

b

7

7

7

7

Chi storey 1

1

2

1

2

1

1

2

l

2 1 1

1

1

1

3

I

I

2

2

4

3

2 1

1

2

2

2

3

1

1

2

2 1 1

2

1

1

l

1

1

2

1

7

CM rnatthewsi

1

2

?

2

1

1

2

t

2 1 1

1

1

1

3

1

1

2

2

4

3

2 1

1

1

1

2

2

?

1

2

2 1 1

2

1

1

1

1

1

2

1

2

Chi cavicolUs

7

2

?

2

1

1

?

1

2 1 1

1

1

1

3

7

1

2

2

4

3

2 1

1

2

2

2

2

7

1

2

7 1 1

4

7

7

1

3

7

?

7

7

Chi tuberculata

7

2

?

2

2

1

1

1

2 1 1

2

1

1

l

7

2

?

7

7

2 7

1

2

2

2

1

7

2

1 1 1

3

1

7

1

?

7

7

?

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Chi mareeba

1

2

7

2

1

1

I

1

1 1 1

1

1

1

3

1

2

2

l

3

2 1

1

2

2

2

1

7

2

2 1 1

1

1

1

1

3

7

7

7

?

Chi parallelus

1

2

?

2

1

1

1

2

1 1 1

1

1

1

3

1

1

2

2

1

2

2 1

1

2

2

2

1

1

2

2 1 1

2

1

7

1

1

7

1

2

?

Chi NewCalcdonia'

1

2

2

2

2

2

1

1

1 1 1

1

1

1

1

1

1

2

2

4

3

1 2

1

1

1

2

1

1

2

2 1 1

3

1

1

1

2

l

1

7

230

MEMOIRS OF THE QUEENSLAND MUSEUM

Chi bifovaecollis
Chl.niolsem
Chi australis
Chi lepida
Chi montoithi
CW sedfera
Chi lawrencei
Chi variolosa
Chlpeckl
Chl.myrmocophila
Chi mallee
Chi loculcsa
Chl.dwgallion
CW.luafer
Chl.bataviao
Chi acutricha
Chi ©ctnlommae
Chl.kununurra
Chi papuae
Chi |ayawijaya
Chi burnolta
Chi zborowskii

Chl plunseta

Chi contorts
Chi pibsipes
Chl.latipos
Chi labponnis
Chi crowcrolti
Chi macmillani
Chi null arbor
Chi. rotunda
Chi weiri
Chi disporsa
Chi stnatolla
Chi formicicota
Chi longtpos
Chi inaoquolw
Chi agios
Chi convergans
Chi trans versa
Chi pygidialis
ChLcannicollis
Chi serricoliis
Chi setipennis
Chi coronls
Chi erupta
Chl.roticulata
Chi dimorpha
Chl.stngicollis
ChFantonnala
Chi tnchonota
Chi rana

Chi compress*pes
Chi pallida
Chi stnaliponnts
Chlleai

Chi mormolyce —
Chi ’NewCaledoma'
Chl.tuberculata
Chi cnrinota
Chi convexa

Chi matlhewsi
Chi cavicollis
Chi inquiUna
Chl.epipleurahs
Chi sculptus
Chi mareeba
Chi parallels
Ed opaca
Euc comat . 1
new genus3
Of«»ct obliquus
Pheid.minuta
Orect dumogae
genus 1
new genus2
Cerat.n sp
nr.Peploglyptus
Onth flohri
Stict. front

Jjifovaecollis

subgroup

_ strigicollis
group (part)

ectatommae

group

# latipes
subgroup
__ epipleuralis
group (part)

> longipes
' group

_ pygidialis
’ group

strigicollis
‘ group (part)

_ striatipennii
‘ group

t strigicollis
’ group (part)

epipleuralis
group (part)

FIG 31. Strict consensus of 76,000 equally parsimonious trees based on equally weighted analysis.

single decay steps, with a few species pairs and
trios supported more strongly.

The species groups of Chlamydopsis proposed
above are only roughly recovered. The
striatipennis group is recovered in the equally
weighted analysis, but in the reweighted analysis,
the invariably monophyletic longipes group
appears within it. In the equally weighted
analysis the longipes group arises from within a

mixed strigicollis group (partial) + pygidialis
group clade. These two groups resolve together
in both trees, with the pygidialis group
consitituting a distinct clade only in the
reweighted analysis. Three species tentatively
suggested as related to the strigicollis group (C.
monteithi, C. setifera, and C. Iawrencei) do not
resolve with this group in either analysis, but
instead appear within the ectatommae group, in

REVIEW OF CHLAMYDOPSIS

231

rC^

rC

Chi bifovaecollis
Chl.nielseni
Chl.lopida
Chi australis
i — Chi monteithi
i — Chl.setifera

Chl.lawrencai

Chl.mallee

i — Chi variolosa
1 — Chl.pecki
Chl.loculosa
Chl.degallien
Chi myrmecophila
Chi lucifer
Chl.bataviae
Chlacutricha
I — Chl.ectatommae
I — Chl.kununurra

— Chl.papuae

— Chljayawijaya

— Chi bumettu

— Chi zborowskii

— Chl.pluriseta
Chl.contorta
Chi pilosipes

. — Chl.labpes i

' — Chi latipennis 1
Chl.crowcrofli 1

r

Chl.macmillani I

Chl.dispersa

. — Chl.striatella
“I — Chl.formicicota

m bifovaecollis

'subgroup

.... strigicollis
group (part)

> eclatommae
' group

# talipes

* subgroup

__ epipleuralis
group (part)

Chl.antonnata
Chi trichonota
Chl.rana
Chl.compressipes
Chi pallida
Chl.stnatiponnis
Chl.longipes
Chl.matKiualis
Chl.agilis
Chi Inai

Chi mormolycc
Chi convergens
Chi. trans vena
Chl.coronis
Chi erupta
Chl.cannicollis
Chi pygidiahs
Chi semcollis
Chl.setipenms
Chi. reticulata
Chi dimorpha
Chl.stngicollis
Chi "NewCaledonla -
Chi tuberculata
Chi carinota
Chi convexa
Chi detecti
Chi storeyi
Chi matthewsi
Chi cavicollis
Chi inquilina
Chl.epiplouralis
Chl.sculptus
Chi mareeba
Chi parallels
Ect.opaca
Euc.comata
new gonus3
Orect obliquus

s

B

striatipennis
' group

^ longipes
* group
_ strigicollis
' group (part)

pygidialis
" group

strigicollis
" group (part)

epipleuralis
’ group (part)

Phoid minuta
Orect.dumogae
new genus 1
new genus2
Cerat n sp
nr.Poploglyptus
Onth.flohri
Stict.front

FIG. 32. Strict consensus of 10,762 equally parsimonious trees based on reweighted analysis.

the reweightcd analysis as sister group to the four
species of the bifovaecollis clade. While this
alternative obviously merits closer investigation,
it does require the loss of some significant
features on the branch leading to these three,
notably the divergent prostemal groove and the
anterolateral pronotal groove. The ectatommae
group itself appears as a coherent lineage in both

equally weighted and reweightcd trees (apart
from, in both, the inclusion of the three
strigicollis group species mentioned above). It is
also worth noting that this clade includes the
bifovaecollis subgroup (the inclusion of which
here was suggested with some reservation).

A large clade, comprising most members of the
strigicollis group, and the striatipennis , longipes ,

232

MEMOIRS OF THE QUEENSLAND MUSEUM

and pygidialis groups, is recovered in both
analyses, with several unplaced and epipleuralis
group species at its base. The species designated
as epipleuralis group are not recovered as a clade
in either analysis. This group is scattered, with the
latipes subgroup and a few others as a grade basal
to the ectatommae group, and the remainder as a
grade basal to nearly all other Chlamydopsis. Of
the species not placed in groups above, no
relationships to other particular groups are
strongly supported. Chlamydopsis parallelus and
C. mareeba are resolved (sequentially) at the very
base of Chlamydopsis , while C. tuberculata and
the New Caledonian species appear near the base
of the ( strigicollis + pygidialis + longipes +
striatipennis group) clade.

DISCUSSION

The forty new species of Chlamydopsis
described in this treatment more than double the
known species diversity, and greatly increase the
known morphological and geographical range of
the group. There are now 69 described species of
Chlamydopsis , with representatives from every
Australian state and both Papuan and Irian New
Guinea (and New Caledonia). The species
diversity in Queensland has emerged as clearly
exceeding any other region, whereas species
from near the populated areas of Victoria and
New South Wales previously predominated. It is
important to note, however, that while
Queensland harbours the bulk of species
diversity, phylogenetic diversity is more evenly
distributed, with several species groups occuring
primarily elsewhere (e.g. longipes group,
bifovaecollis subgroup, latipes subgroup).

The previously unappreciated sexual di-
-morphism in Chlamydopsis is very interesting.
Obvious sexual dimorphisms are generally rare
in histerids. Those that have been documented
have been primarily attributed to courtship
(Caterino, 2002). The antennal club dimorphism
(male antennal club twice or more the length of
the female’s — apparently first noted by Oke,
1923), nearly ubiquitous in Chlamydopsinae,
obviously must have some olfactory sig-
nificance. But it is impossible to say whether this
relates to mate location, ant nest location (and
perhaps differing dispersal tendencies between
males and females), or some other factor. The
significance of elytral and other textural
dimorphisms is even more obscure. Body texture
in myrmecophiles is often attributed to Was-
mannian mimicry (Wasmann, 1889), in which
guests’ surface sculpturing (and, in some, shape

and color) mimics that of the host, presumably a
tactile disguise. Accepting that the underlying
causes of this similarity have been controversial
(Wilson, 1971), it is nonetheless observed in
many species of Chlamydopsinae. Regardless
whether this mimicry is directed at the hosts or at
potential predators (Mclver, 1987), the fact that
intersexual morphological differences are
observed in some Chlamydopsinae suggests that
some differences in host relationships or activity
patterns exist between them.

Outlines of the phylogeny of Chylamydopsis
have begun to emerge from this study. Several
apparently monophyletic groups of species have
been identified, and although relationships
among them need additional study, relationships
within them are relatively consistent across
analyses. Outgroup relationships to Chlamydopsis
are in greatest need of additional analysis. The
exact relationships of Chlamydopsis to Eucurtia
and Ectatommiphila , in particular, are unclear. It
was considered initially likely that Chlamydopsis
would prove paraphyletic with respect to these
other two genera. However, with existing data,
the monophyly of Chlamydopsis is supported.
The relationships among more distant outgroups
yield additional uncertainty with respect to
relationships within Chlamydopsis. Apart from
the New Guinean species (‘new genus3’), all of
the chlamydopsine outgroups here almost
certainly constitute a clade (lacking a dorsally
visible scutellum, and having a substantially
elongated prothorax). It is unclear why these
were not resolved as such in either analysis, and
what effect this might have on resolutions
elsewhere in the tree.

While the phylogenetic results obtained here
do not justify a great deal of evolutionary
exploration, one character reconstruction,
especially, merits some discussion. These trees
agree in reconstructing a deep prostemal groove,
which departs from the margin laterally, as basal
within Chlamydopsis. This groove is then
subsequently weakened and lost in various other
groups. This well developed and divergent
groove is one of the most distinctive and unusual
characters in Chlamydopsis , primarily of the
epipleuralis group, and its evolution according to
this scenario would be very surprising. This
single result casts a shadow of doubt over much
of the basal resolution in these trees.

One of the primary impediments to resolving
relationships here has been the representation of
so many species by only a single sex (or in some

REVIEW OF CHLAMYDOPSIS

233

TABLE 3. Published host records for species of Chlamydopsis. Only valid host species names are listed. See
Table 1 for equivalence with originally published host names. Letters refer to literature cited: a = Lea, 1910; b =
Lea, 1912; c = Lea, 1914b; d=Lea, 1918; e = Lea, 1919; f= Lea, 1925; g = Oke, 1923; h - King, 1869; i=this
study. The previously reported host has been split into these three species potentially sympatric with the beetle.
2 Lea reports that this species was collected in the vicinity of three species of ant; it was not possible to determine
which was the host: Myrmecia pyriform is, Ectatomma metallicum , Pheidole conflicts 3 These records were
reported for the now synonymised C. excavata Lea.

Host subfamily

Ponerinae

Dolichoderinae

Formicinae

Myrmecinae

Valid host
species

|

"a

6

5

§

|

*

Rhytidoponera sp.

Rhytidoponera violacea

Rhytidoponera punctata

Pachycondyla lutea

Iridomyrmex gracilis

gracilis

Iridomyrmex rufoniger

ssp.

I. purpureus

I. sanguineus /. foreli 1

Iridomyrmex conifer

Iridomyrmex sp.

Dolichoderus ypsilon

\ypsilon

Notoncus

ectatommoides

d

E

<A

11

If

5 Si

5 §

U a

Crematogaster sp.

Meranoplus minor

i

it

9 8.

i a
Is

Aphaenogaster sp.

C. striatipennis

f ’P

g

e,f,g

C. leai

g

C. pallida

d

C. reticulata 2

f

e\ g 3

C. strigicollis

g

C. mormolyce

f

C. pygidialis

g

f

C. carinicollis

£

e

C. serricollis

c

C. setipennis

C. longipes

a,b,g

C. agilis

c,e,f

C. ectatommae

b,d,p

C. myrmecophila

i

C. loculosa

f

f

C. bifovaecollis

g

g

C. epipleuralis

g

b,f

C. sculptus

g

C. formicicola

h,b,f

C. latipes

f

e

C. inquilina

e

C. detecti

c

C. tuberculata

e

cases, perhaps, the inability to associate males
with females). The sexual dimorphism in
integumental texture, and, in some, trichome
morphology, may prove to be important
phylogenetic markers. But at present too many
species have had to be scored as ‘unknown’ for
them to have had much positive effect.
Ovipositor morphology, likewise, shows
interesting variation, but is missing in too many
taxa to be as informative as it might. The fact that
many species remain known only from types

(which I have rarely risked to dissect) contributes
further ambiguity to the dataset and results.

A summary of known host associations is
presented in Table 3. These represent a broad
phylogenetic range of ants, with hosts from four
different subfamilies. Hosts in the Ponerinae
predominate (13 species of Chlamydopsis ), with
Dolichoderinae a close second (9 species). It is
perhaps surprising to note that several beetles use
multiple hosts, even hosts in different subfamilies
(e.g., C. striatipennis with Rhytidoponera and

234

MEMOIRS OF THE QUEENSLAND MUSEUM

Iridomymxex ; C. latipes with Rhytidoponera and
Dolichoderus ; C. carinicollis with
Rhytidoponera and Aphaenogaster). Some of
this may reflect local differentiation, and the host
identifications given in the literature, for the most
part, cannot be verified, but it appears that host
specificity may be low for some species. This
would have obvious implications in terms of
chemical and behavioural integrating mechanisms.
Similarly, little in the way of phylogenetic
structure is evident with respect to host use, with
apparently closely related species (e.g., C.
striatipennis , C. leai, C. pallida) ranging across
multiple host species and subfamilies. One
possible phylogenetic distinction worth noting is
the lack of host records for the ant Pheidole .
Species of this ant genus are the preferred hosts
for several species of Pheidoliphila , and this may
represent a deep divergence between these two,
broadly sympatric chlamydopsine genera.

While more large scale surveys will un-
doubtedly turn up additional interesting species
of Chlamydopsis , the most pressing need at this
stage is for more specific collecting. Locating
these species in their natural environments will
allow identification of hosts (known now for only
a small fraction of species), facilitate association
of sexes, and allow preservation of specimens for
molecular work. At present only a single
specimen of Chlamydopsinae adequate for DNA
study has been obtained. It is unlikely that full
phylogenetic resolution for the group will be
achieved without combining morphological and
molecular data. These ambiguities and
limitations notwithstanding, the study of
Chlamydopsinae is advancing rapidly. This is a
fascinating and wonderful group of insects, and
undoubtedly their continued study will yield
many evolutionary insights.

ACKNOWLEDGEMENTS

I am indebted to many collectors and curators
for seeking, preparing, and providing the
specimens that formed the basis of this study. The
efforts of Ross Storey (QDPI) and Geoff
Monteith (QMB), in particular, made this work
possible (not to mention necessary). For
providing additional material, as well as
answering numerous questions about localities,
habitats, etc., I am also very grateful to Tom Weir
(ANIC), Max Moulds (AMS), Henry Howden
(HAHC), Phil Perkins (MCZ), Catriona McPhee
(MVM), Martin Brendell and Stuart Hine
(NHM), Eric Matthews (SAM), Wolfgang
Schawaller and Alex Riedel (SMNS), Dave Furth

and Nancy Adams (USNM), Terry Houston and

Brian Hanich (WAM), and Nicolas Degallier.

LITERATURE CITED

BLACKBURN, T. 1891. Further notes on Australian
Coleoptera, with descriptions of new genera and
species.IX. Transactions and Proceedings of the
Royal Society of South Australia 14: 65-154.

1900. On some new genera and species of
Australian Coleoptera. Proceedings of the Royal
Society of Victoria 12(n.s.): 206-233.

1901 . Further notes on Australian Coleoptera, with
descriptions of new genera and species.
Transactions and Proceedings of the Royal
Society of South Australia 25(11): 99-131.

CATERINO, M.S. 2000. Descriptions of the first
Chlamydopsinae (Coleoptera: Histeridae) from
Wallacea. Tijdschrift voorEntomologie 143: 267-278.

2002. Revision of the Mister militaris group
(Coleoptera: Histeridae). Annals of the
Entomological Society of America 95: 323-334.

DEGALLIER, N, 1984. Gomyopsis kuscheli , nouveau
genre et nouvelle especc de Chlamydopsinae
(Coleoptera: Histeridae). Nouvelle Revue
Entomologie (n.s.) 1: 55-59.

KING, R.L. 1869. On the Byrrhides of Australia.
Transactions of the Entomological Society of
New South Wales 2: 71-75.

LEA, A. 1910. Australian and Tasmanian Coleoptera
inhabiting or resorting to the nests of ants, bees
and termites. Proceedings of the Royal Society of
Victoria 23: 116-230.

1912. Australian and Tasmanian Coleoptera
inhabiting or resorting to the nests of ants, bees
and termites. Supplement. Proceedings of the
Royal Society of Victoria 25: 31-78.

1914a. Notes on some miscellaneous Coleoptera,
w ith descriptions of new species. Transactions
and Proceedings of the Royal Society of South
Australia 38: 249-344.

1914b. On Australian and Tasmanian Coleoptera
with descriptions of new species. Part II.
Proceedings of the Royal Society of Victoria 26
(n.s.): 211-227.

1918. On Australian Coleoptera. Part I. Records of
the South Australian Museum 1: 83-104.

1919. Notes on some miscellaneous Coleoptera,
with descriptions of new species. Part V.
Transactions and Proceedings of the Royal
Society of South Australia 43: 166-261.

1925. XIII. On Australian Histeridae (Coleoptera).
Transactions of the Entomological Society of
London 57: 239-263.

LEWIS, G 1885. New species of Histeridae, with
synonymical notes. Annals and Magazine of
Natural History 15(5): 456-473.

1903. On new species of Histeridae and notices of
others. Annals and Magazine of Natural History
12(7): 417-429.

REVIEW OF CHLAMYDOPSIS

235

1910. On new species of Histeridae and notices of
others. Annals and Magazine of Natural History
6(8): 43-58.

1913. On new species of Histeridae and notices of
others. Annals and Magazine of Natural History
12(8): 81-87.

MAZUR, S. 1984. A world catalogue of the Histeridae
(Coleoptera: Histeridae). Polskie Pismo
Entomoiogiczne 54: 1-376.

1997. A world catalogue of Histeridae. Genus
(Supplement): 1-373.

McIVER, J.D. 1987. On the myrmecomorph
Coquillettia insignis Uhler (Hemiptera):
arthropod predators as operators in an
ant-mimetic system. Zoological Journal of the
Linnaean Society 90: 133-144.

McMILLAN. R.P. 1950. Observations on the jockey
beetle (Chlamydopsis duboulayi ) [actually C.
loculosa - see McMillan, 1954]. The West
Australian Naturalist 2: 132-133.

1954. Name of a Jockey Beetle a correction. The
West Australian Naturalist 4.

NISHIKAWA, M. 1995. Notes on chlamydopsine
histerid beetles of Japan, with description of a new
species. Elytra 23: 257-261.

1 996. On the genus Eucurtiopsis Silvestri occurring
in Japan and Taiwan. Kanagawa-Chuho,
Odawara 113: 7-1 1.

OKE, C. 1923. Notes on the Victorian Chlamydopsini
(Coleoptera), with descriptions of new species.
Victorian Naturalist 49: 152-162.

SORENSON, M.D. 1999. TrecRot, version 2. (Boston
University).

SWOFFORD, D.L. 1998. PAUP*: Phylogenetic Analysis
Using Parsimony, version 4.0b4a. (Sinaucr).

SHATTUCK, S.O. & BARNETT, N.J. 2001 . Australian
Ants Online. (CSIRO-Entomology,
www.cnto.csiro.au/science/ants/default.htm).

WASMANN, E. 1889. Nachtragliche Bemerkungen zu
Ecitochara und Ecitomorpha. Deutsche
Entomologische Zeitschrift 83: 414.

WESTWOOD, J.O. 1869. Remarks on the genus
Ectrephes , and descriptions of new exotic
Coleoptera. Transactions of the Entomological
Society of London 1869: 315-320.

1874. Thesaurus Entomologicus Oxoniensis, or,
Illustrations of New, Rare, and Interesting
Insects, for the Most Part Contained in the
Collections Presented to the University of
Oxford by the Rev. F.W. Hope, M.A., D.C.L.,
F.R.S., Oxford. XXIV, 205.

WILSON, E.O. 1971. The Insect Societies. (Belknap
Press of Harvard University Press: Cambridge).

*

BARAHNA, A NEW SPIDER GENUS FROM EASTERN AUSTRALIA (ARANEAE:

AMAUROBIOIDEA)

VALERIE TODD DAVIES

Davies, V.T. 2003 06 30: Barahna , a new spider genus from Eastern Australia (Araneae:
Amaurobioidea). Memoirs of the Queensland Museum 49(1): 237-250. Brisbane. ISSN
0079-8835.

Eight new species of Barahna gen. nov. are diagnosed, described and illustrated; these are
Barahna booloumba , the types species and B. brooyar , B. yeppoon , B. taroom , B. scoria , B.
toonumbar , B. myall and B. glenelg. The last is described from a single female. The genus
ranges from mid-eastern Queensland, through northern and eastern New South Wales to
southern Victoria. □ Araneae, Amaurobioidea, Barahna, new genus, eastern Australia.

Valerie Todd Davies, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia;

10 September 2002.

This genus is unique among described
Australian amaurobioids in having an embolus
with obvious ‘pars pendula\ a name coined by
Comstock (1913). It refers to the membraneous
part of the embolus in which the sperm duct is
carried; it is reduced and not seen in most emboli.
A further feature is the retrolateral apophysis on
the male palpal patella. This occurs in at least two
other, probably unrelated, Australian amauro-
bioids, Paramatachia and Dardurus.

Barahna spp. have been found in wet
sclerophyll, vine scrub and open eucalypt forests
from mid-east Queensland and eastern New
South Wales. A single female specimen has been
collected from southern Victoria suggesting that
it also occurs in the coastal forests of that State.
The spiders were found in sheet webs with a
small retreat under rocks, logs or near the base of
trees; they run on the underside of the web. Egg
sacs containing 8 or more eggs are carried in the
chelicerae. All species are named for their type
locality.

METHODS

Most ot the spiders were collected from pitfall
traps (PF) or sieved litter samples. Notation of
spines follows Platnick & Shadab (1975);
measurements are in millimetres; the left palp is
described and illustrated unless noted.

Abbreviations. Museums: AM, Australian
Museum, Sydney; QM, Queensland Museum,
Brisbane; SAM, South Australian Museum,
Adelaide; WAM, West Australian Museum, Perth.

Collectors: DC, D. Cook; DW, D. Wallace;
GBM, G B. Monteith; GT. G Thompson; MRG,
M. R. Gray; RJR, R. J. Raven; SRM, S. R.
Monteith; VED, V. E. Davies.

Location data: NP, National Park; SF, State
Forest.

Anatomy: The usual abbreviations are used for
body measurements, eyes and spinnerets.
Abbreviations on illustrations are explained in
the legends;. others in the text.

SYSTEMATICS

Barahna gen. nov.

TYPE SPECIES. Barahna booloumba sp. nov.

ETYMOLOGY. From the Aboriginal ‘baran-barahn’,
spider in the Bundjalung language of northern New South
Wales-Queensland border region.

DIAGNOSIS. Females may be distinguished
from other cribellate amaurobioids by the
presence of a pair of small dorso-posterior pro-
tuberances in the foveal region (Fig. IB); these
each have a small pale (unsclerotised) area (Fig.
7). Males may be distinguished by a membraneous
pars pendula on the concave side of the embolus;
as well, the palpal patella has a retrolateral
apophysis (Fig. II).

DESCRIPTION. Small, 3-clawed cribcllates
(most less than 4.0). Coloration of the abdomen
varies from pale to dark grey with a pattern of
light spots in a vague chevron pattern (Fig. 1C).
Carapace highest behind eyes; eyes directed
forwards. From above, posterior eye row
straight-slightly recurved, anterior row recurved.
AME reduced. Two retromarginal and 4-6
promarginal cheliceral teeth (Fig. IE); prolateral
filamentous seta at base of fang longer than other
setae. In the male the retromarginal teeth may be
reduced in size and form a diagonal line with the
promarginals. Labium about as long as wide;
sternum longer than wide, pointed posteriorly

238

MEMOIRS OF THE QUEENSLAND MUSEUM

(Fig. 1 D). Legs 4 1 23 with bands of darker colour.
Feathery hairs on legs and cephalothorax (Fig.
3D). Tarsal trichobothria in a single row, in-
creasing in length distally; bothrium collariform;
tarsal organ oval. Female genital atrium with
medial thickening which is sclerotised basally
(Fig. 1G); this sclerotisation provides a useful
diagnostic character; it is not delineated in
micrographs (Fig. 3A). Tiny lateral teeth, visible
at high magnification, on posterior lateral edges
of atrium. Long, coiled insemination ducts to
posterior spermathecae (Fig. 3B). Male palp with
oval tegulum, circular or semi-circular embolus
with pars pendula; membraneous conductor. A
small, thin apophysis (? median) obscured by
conductor (Figs IK, 2D) may be present.
Retrolateral tibial and patellal apophyses.
Cribellum with two spinning fields in female:
large broad colulus in male. Proximal calamistrum
with one row of setae. Large ALS with two MAP
in female, one and a nubbin in male (Fig. 4B).
PMS with paracribellar spigots in female. Four
simple tracheal tubes in abdomen, medians
longer than laterals.

KEY TO BARAHNA SPP.

1. <3, very long embolus encircling tegulum to terminate

postero-retrolaterally. 9, basal sclerite of epigynal
atrium pointed (Fig. 1G) or rounded 2

<3, embolus describing about three-quarters of a circle to
terminate mid-retrolaterally (Fig. 6A). 9, basal sclerite
of atrium flat-topped (Fig. 6D) 6

2. <3, embolus arising postero-retrolaterally on tegulum
(Fig. 2G). 9, basal sclerite short, rounded (Fig. 2J)

yeppoon

d, embolus arising antero- or mid-retrolaterally on
tegulum. 9, basal sclerite pointed, or slender and
rounded (Fig. 5 D) ♦ 3

3. <3 , embolus curled distally. 9 , basal sclerite pointed . 4

(3, embolus slightly curved distally. 9, basal sclerite
slender or rounded. 5

4. (3, embolus arising on mid-retrolateral tegulum

, booloumba

8, embolus arising on antero-retrolateral tegulum (Fig.

2 A) . brooyar

5. 8 , palpal patellal apophysis truncated (Fig. 5B). 9 , basal
sclerite long, slender with rounded apex .... taroom

8, palpal patellal apophysis digitiform. 9, basal sclerite

short, with small pointed apex (Fig. 5F) scoria

6. 9, basal sclerite as long as wide (Fig. 6D) 7

9, basal sclerite longer than wide (Fig. 6L) . . . glenelg

7. 8 , patellal apophysis simple,digitiform. 9 , spermathecae

small toonumbar

8, patellal apophysis long and bifid (Fig. 6G).9,
spermathecae large myall

Barahna booloumba sp. nov.

(Figs 1 A-K, 3A-C, 4A-C, 7 A, 8B; Table 1)

ETYMOLOGY. From the type locality Booloumba Ck,
Conondale NP.

MATERIAL. HOLOTYPE. <3, Booloumba Ck,
Conondale Ra, southeast Queensland, 26°38’S, 152°39’E,
200m. rainforest, PF, 29 Nov. 1991-8 Jan. 1992, DC (QM
S42241). PARATYPES. Southeast Queensland: 9, same
locality as holotype, PF, 13-18 Apr. 1976, RJR (QM
S42242); c3, 29 (QM S42243); 4<3, 29 (QM S42244);
28 , 9, Sunday Ck, Conondale Ra, 26°43'S, 152°34'E,
900m, rainforest, PF, 7 Jan.-2 Mar. 1992, DC (QM
S42245); 28, 29, same data as holotype (QM S42247);

9 , same locality as holotype, PF, 29 Nov. 1974-Feb. 1 975,
G Maywald (QM S42248); 8, Little Yabba Ck via
Kenilworth, 26°36’S, 152 0 35'E,PF, 10 Aug.-9Nov. 1974,
GBM, SRM (QM S42249); 28, same data (QM S42250);
2d, Sunday Ck, 900m, PF, 29 Nov. 1996-7 Jan. 1997, DC
(QM S4225 1 ); 8 . Mapleton Falls NP, 26°38’S, 1 52°5 1 'S,
500m, PF, 8 Jan.-3 Mar. 1992, DC(QM S47 136); c3,Ewan
Maddoek Dam, 26°48’S, 152 0 59'E. PF M. Glover (QM
S28512); 39. 38, Mt Coolum, 26°34’S, 153°05 , E, PF,
Jan. 1984, B. R. Jahnke (QM S42252); 9, Kenilworth SF,
26°38’S, 152°39 , E, wet sclerophyll forest, 5 May 1998, G
Milledge(AM KS7 1029); 2 (3, same data (AM KS71030):
c3, 9 (AM KS71031); 29, rainforest (AM KS71032); <3,
9, Conondale NP, 26°42 , S. 152°37’E, wet sclerophyll
forest, G Milledge (AM KS71033); 8 , Little Yabba Ck,
via Kenilworth. PF, 29 Dec. 1974-22 Mar. 1975, GBM,
SRM (QM S45688); 8, Cold Ck via lmbil, 26°27’S,
152°37’E, PF, 31 Dec. 1974-27 Mar. 1975, GBM. SRM
(QM S45689); <3, Sunday Ck, 26°40*S, 152°34’E,
rainforest, PF, 18 Dec. 1996-20 Jan. 1997, GBM (QM
S45690); <3, Mary Caimcross Pk via Maleny, 26°43'S,
152°33’E, 240m, PF, 29 Dec. 1974-27 Mar. 1975, GBM,
SRM (QM S4569 1 ); 8 , Dingo Ck via Traveston, 26° 1 9'S,
152°48’E, 1 00m, PF 5, 9 Nov. -3 1 Dec. 1974, GBM. SRM
(QM S45692); 2(3, Camerons Scrub, 27°31’S, 152°44'E,
50m, PF 7562. 11 Nov. 1998-13 Jan. 1999, GBM, DC, GT
(QM S45693), 8, 9, Camerons Scrub, top of ridge,
27°30’S, 152°44'E, 160m, rainforest, sieved litter, 21 Nov.
1998, GBM, GT (QM S45718); 2 8, Mt Coolum, open
forest, Jan. 1984, B.R. Jahnke (QM S45694); 98, 9,
Enoggera Reservoir, 27°27 , S, 152°55’E, site 3. 100m,
rainforest, PF 9214, 27 Jan.-15 Mar. 2000, GBM, J.Holt
(QM S45728); 4c3, 9, same locality, site 8, rainforest, PF
9068, 10 Nov -21 Dec. 1999,GBM(QM S45729):7<3,site
3,PF9063, 160ct.-21 Dec. 1999,GBM(QM S45730); 9,
PF 9433, 1 5 Mar.- 1 8 May 2000, GBM (QM S4573 1 ); 4<3 ,
9, PF 9098, 21 Dec. 1999-27 Jan. 2000. GBM, J. Holt
(QM S45732); 2c3, Mt Cabinet via Jimna, 26°43’S,
152°34’E, PF 29, 30 Nov.-29 Dec. 1 974, GBM, SRM (QM
S45733); 8, same data, 29 Mar.- 16 June 1975 (QM
S45734); <3, Deer Reserve, Kilcoy, 26°59’S, 152°29, PF
30, 1 1 Jan.-29 Mar. 1975, GBM, SRM (QM S45735); 9,
same locality, 1 June-24 Aug. 1975 (QM S45736); 3(3,
Tungi Ck, Jimna, 26 o 40’S. 152°28'E, PF, rainforest, 18
Dec. 1996-20 Jan. 1997, GBM (QM S45737); 3 <3, same
locality, PF28, 29 Dec. 1974-29 Mar. 1975, GBM, SRM
(QM S45738); 2 <3, Wrattens Camp Forestry via Widgee,

NEW SPIDER FROM EASTERN AUSTRALIA

239

FIG. 1. A-K, Barahna booloumba sp. nov. A-H 9; A, eyes, chelicerae (frontal); B, cephalothorax (lateral); C,
body (dorsal); D, sternum, labium, maxillae; E, cheliccra (ventral); F-H, epigynum (posterior, ventral, dorsal
cleared). I-K, 6 palp (ventral, retrolateral, dorsal), bs = basal sclerite, c = conductor, pa - patellal apophysis, pp
= pars pcndula, ta = tegular apohysis.

26°18’S, 152°20’E, PF 10, 11 Aug.-lONov. 1974, GBM,
SRM (QM S45696); 9 , same data (QM S45697); 26 , 8 9 ,
Perrys Knob, 27°36’S, 1 52°36’E, vine scrub, 200m, sieved
litter, 15 May 1999, GBM (QM S45695); 9, <3, same
locality, PF 7564, 1 1 Nov. 1998-13 Jan. 1999, GBM, DC,
GT (QM S45709); 9, Mt Cotton, 27 t> 37’S. 153°13’E,
200m. sieved litter, 13 Sept. 1997, GBM (QM S45714); 9,
Dandabah, Bunya Mts NP, 26°53*S, 151°36 T E, notophyll
vine forest with Araucaria emergents (hoop pine
rainforest), 960m, litter, 29 Feb. 1 976, VED (QM S45667);
9 Paul Lentz Plain, Bunya Mts NP, 26°50’S, 151°33’E,
hoop pine rainforest , PF, 7 Nov. 1994, Queensland
National Parks (QM S45668); 9, Bunya Mts NP,
rainforest sieved litter, 2 Oct. 1979, GBM (QM S45669);

Dandabah, Bunya Mts NP, under log with sheet web,
4 Sept. 1974, RJR (QM S45670); 6. Bunya MLs NP,
26°54’S, 15l°37’E, 1006m, rainforest, PF19, 1974-1975,
GBM, SRM (QM S45671); 6, same locality, 960m,
rainforest, sieved litter, 2 Oct. 1979, GBM (QM S45674);
46, 9, Belthorpe, 26°50'S, 152°41’E, rainforest , PF,18
Dec. 1996-20 Jan. 1997, GBM (QM S55185); 26, 39,
Redwood Pk,nr, Toowoomba, 27°34’S, 152°00’E, 7 Jan.
1986, GBM, SRM (QM S55186); 29, 6, Cunninghams
Gap NP. 28°03 , S, 152°23’E, dry forest, sieved litter, 28
June 1 99 1 , D. Black (WAM 98/208 1 -3); 6 . Ravensboume
NP 27°22’S, 152°I FE, 731m, rainforest, PF38, 10 Nov.
1974-12 Jan. 1975, GBM, SRM (QM S55188) ; 99,
juv.d, Upper Brookfield nr Brisbane, 27°28’S, 152°52’E,

240

MEMOIRS OF THE QUEENSLAND MUSEUM

notophyll rainforest with Araucaria , litter, 22 May- 11
Aug. 1976, RJR. VED (QM S55190); 3 <5, 29, same
location, 28 Jan. 1 982, RJR, VED (QM S55 1 9 1 ); 6 , 9 , PF,
23 Dec. 1 980-22 Jan. 1981, RJR, VED (QM S55 1 94); 4 9 ,
17 July 1981 (QM S55195); 9 with egg sac under log,
same data, 2 Sept 1981 (QM S55198); 39, 2<5, PF, 8-23
Apr. 1981 (QM S55199); 3d, Gold Ck nr Brisbane,
27°28'S, I52°53’E, PF, 31 July-1 Sept. 1981, RJR, VED
(QM S55200); <5, Flinton Hill nr Ipswich, 1 20m, PF 52, 20
May- 11 Aug. 1976, GBM, SRM (QM S55201); <3, Mt
Glorious, 27°20’S, 152°45’E, 29 Sept. 1973, RJR (QM
S55202); 29, Jollys Lookout, Boombana NP, open forest,
litter, 30 June 1991, D. Black (WAM 98/2090-2); 9,
Brisbane Forest Pk nr Boombana, closed forest PF, 17-24
July 1 987, Rhonda Grundy (QM S55203); 2 6 , Mt Mee via
Samford, 27 a 05 , S, 152°41’E, rainforest, PF14, 9 Nov.
1974-11 Jan. 1975, GBM. SRM (QM S55204); 29, Mt
Mee SF, 27°06’S, 152°42*E, 550m, rainforest stick
brushing, 28 Feb. 1979, GBM (QM S55205); c?, 9, Mt
Mee. Upper Neunim Ck, 27°05'S, 152°42 , E, rainforest,
sieved litter, 3 Sept 1979, GBM (QM S55206); c5, Mt
Mee, Plateau Site, rainforest PF 89, 8 Oct. 1977-20 Jan.
1978, GBM (QM S55207); <5, Cainbable via Lamington
NP, 28°10’S, 153°07 , E, 762m. PF 57, 28 Sept 1975-31
Jan. 1976, GBM. SRM (QM S 55208); 2 <5 , Bahrs Scrub,
27°46'S, 153°10’E, 100m. rainforest, PF, 10 Dec. 1991-21
Jan. 1992, DC (QM S25032). OTHER MATERIAL. New
South Wales: 6, New Brighton Beach via Brunswick
Heads, 28°3FS, 153°33 , E, 10m, rainforest PF 39, 3
Aug.- 16 Nov. 1975, GBM, SRM (QM S55189).

DIAGNOSIS. Retrolateral origin of embolus;
palpal patellal apophysis digitiform (Fig. IK).
Sclerotised base of epigynal atrium pointed
anteriorly. Dorso-posterior protuberances in
foveal region rounded with small posterior
unsclerotised areas (Fig. 7A).

DESCRIPTION. Male. CL 1.9, AL 1.9. Ratio of
AME: ALE: PME: PLE is 6: 1 0: 1 0: 1 0. Legs 4 1 23
(Table 1). Notation of spines. Femora: I, D110,
P001; II, Dill. P001, R001; III, D101, P101,
R00 1 ; I V, D 1 0 1 , POO 1 , R00 1 . Patellae: III, D00 1 .
Tibiae: I, P011, V102, ROM; II, D001, P101,
VI 02, R 1 01 ; III, D001, P101, V012, R101; IV.
P101, V112, R101. Metatarsi spined with distal
whorl 4-5 spines.

6 palp (Fig. 1I-K). Embolus with
membraneous pars pendula arising retrolateral ly

TABLE 1. Barahna booloumba (5(9) leg lengths.

======

Leg I

Leg II

Leg III

Leg IV

Femur

1-7 (1-5)

1.5(1.3)

1.3(1.2)

1.9(1.5)

Patella X
Tibia

2.3 (1.9)

1.8 (1.4)

1.6 (1.3)

2.3 (1.8)

Metatarsus

1.6(1.3)

1.3(1.0)

1.3 (0.9)

1.9(1.5)

Tarsus

1.0 (0.8)

0.8 (0.7)

0.6 (0.6)

0.8 (0.8)

Total

6.6 (5.5)

5.4 (4.4)

4.8 (4.0)

6.9 (5.6) |

and encircling tegulum clockwise, ending in a
distal curl postero-retrolaterally. Membraneous
conductor arising just anterior to embolus,
broadening distally; small membraneous thin
finger-like tegular apophysis arising dorsal to
embolus and obscured by conductor (Fig.2D).
Retrolateral tibial apophysis (RTA) with curved
blunt ventro-retrolateral and digitiform
retrolateral branches. Smaller digitiform patellal
apophysis.

Other males ranged from 3. 0-3. 8 in length.
Female. CL 1.6, AL 1.8. Ratio of AME: ALE:
PME: PLE is 6:9:9:9. Legs 4123 (Table 1).
Carapace with two small dorso-posterior
protuberances on fovea. Notation of spines.
Femora: l, D101, P001; II, D101, POOL R001;
III, D101, P001, R001; IV, D101, POOL V010,
R001. Patellae: III, D001. Tibiae: I, P01 1, VI 02;
II, P01LV010, R101; III, DOOl, P011, V012,
R011; IV, D 1 01, P011 , V 112, ROIL Metatarsi
spined with distal whorl of 4-5 spines.

Epigynum (Figs 1F-H, 3A,B). Gonopores at
base of atrium; insemination ducts proceed
laterally then coil (two or three coils) forward to
turn back to enter spermathecae. Tiny lateral
teeth on posterior lateral edges of atrium;
sclerotised basal part of atrium pointed.

Spinnerets (Fig. 4A,C). ALS with two major
ampullate spigots, about 14 piriform spigots and
at least two tartipores. PMS with large anterior
spigot (minor ampullate), two posterior spigots
(cylindrical) and eight others. Four of these have
one or two strobilate shafts (paracribellars) and
four have single plain shafts (aciniform). PLS
with two large (cylindrical) and four aciniform
spigots.

Females ranged in length from 3. 5-3.9.

DISTRIBUTION. B. booloumba (Fig. 8B) has
been collected from many localities in
southeastern Queensland and from Brunswick
Heads in coastal New South Wales.

Barahna brooyar sp. nov.

(Figs 2A-F, 8B)

ETYMOLOGY. From the type locality Brooyar SF.

MATERIAL. HOLOTYPE. c5, Brooyar Fire Tower via
Gympie, southeast Queensland, 26° 1 0*S, 1 52°28’E, 457m,
rainforest, PF54, 23 Aug. 1 975-29 Feb. 1976, GBM, SRM
(QM S45722). PARATYPES. Southeast Queensland:
9,c3 same data (QM S45723); c5, same locality and
collectors, 29 Feb.-25 Apr. 1 976 (QM S45724); 7 6 , Marys
Ck, Foresdy via Gympie, 26°16’S, 152°33’E, 240m, PF9,
11 Aug.- 10 Nov. 1974, GBM, SRM (QM S45725); d,
same locality and collectors, 16 June-28 Aug. 1975 (QM

NEW SPIDER FROM EASTERN AUSTRALIA

241

FIG. 2. A-K., Bar ahn a spp. nov. A-F, B. broovar sp. nov. A-D, 6 palp (ventral, retrolateral. dorsal, expanded in
KOM); E,F, 9 epigynum (ventral, dorsal cleared). G-K, B. yeppoon. G-I, 6 palp (ventral, retrolateral, dorsal);
J,K, 9 epigynum (ventral, dorsal cleared), ta = tegular apophysis.

S45726); 9, 28 Mai.- 16 June 1975 (QM S45727); (5,11
Aug.- 10 Nov. 1974 (QM S55209).

DIAGNOSIS. In males the embolus arises near
the anterior retrolateral edge of tegulum (Fig. 2A)
obscuring the anterior coil of the sperm duct,
unlike B.booloumba where it arises
mid-retrolaterally (Fig. II) and the sperm duct is
clearly seen.

DESCRIPTION. Male. CL 2.0, AL 1.8. Ratio of
AME: ALE: PME: PLE is 6: 9: 8: 9. Legs 41 23. 1,
6.9; II, 5.6; III, 4.6; IV, 7.3. Notation of spines
somewhat variable within left and right sides of
spider and between specimens. Femora: I, D01 1 ,

POOL II, D001, POOL HI, DUO, POOL R00(1);

I V, D1 00, R00( 1 ). Tibiae: I, P0 1 1, V2(2)2. R0 1 1 ;
II, D00L P0(0)1, VI 12, R 101 ; HI- D( 1)0(0),
P 1 0 1 . V( 1)12, ROM; IV, D00(1).P0(0)1,V( 1)12,
R(0) 1(1). Metatarsi spined with dorsal whorl of
4-5spines.

d palp (Fig. 2A-D). Embolus arising near
anterior edge of tegulum obscuring coil of sperm
duct; embolus curled distally. RTA digitiform,
ventro-RTA blunt, sclerotised. Patellal apophysis
digitiform.

Males ranged from 2.7-3.9 in length.

Female. CL 1 .9, AL 2. 1 . Legs: I, 6.7; II, 5.3; III,
5.0; IV -. Notation of spines similar to male with

242

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. A-D, Batalina spp. nov. A-C, B. booioumba; A, 9 epigynum (ventral, arrow to lateral teeth); B, epigynum
(dorsal cleared; photo-micrograph showing embolus in insemination duct); C, protuberances on fovea; D, B.
yeppoon, 2 foveal area.

fewer spines on ventral tibiae: HI, 012; IV, VI 1 1 .
Epigynum (Fig. 2E,F). Sclerotised base of atrium
long and pointed.

The other female was 4.0 long.

DISTRIBUTION. B. brooyar( Fig. 8B) has been
collected from only two localities in southeastern
Queensland.

Barahna yeppoon sp. nov.

(Figs 2G-K, 3D,4D, 8A)

ETYMOLOGY. From the type locality, Yeppoon.

MATERIAL. HOLOTYPE. d, Yeppoon. Meikleville
Hill. Queensland 23°06'S. 150"43’E. vine thicket PF, 21
Apr.-l 8 July 1990, DW, RJR (QM S34450).
PARATYPES. Queensland: 39, Yeppoon. Iwasaki Rd,
vine thicket, PF. 18 July-23 Oct 1990, DW, RJR, K.
Williams (QM S33577); d, same locality as holotype, PF,
18 July-23 Oct. 1990; DW, RJR, K. Williams (QM
S34096); d, 79, 6.2km W Yeppoon, 23" 10'S. I50°43'E,
semi-dry scrub, 27 May 2000, G Milledge, H. Smith (AM
KS67481); d, same data (AM KS67482); d, same data

(AM KS67483): 2d, Frenchville, 23°20’S, 150°34’E,
open forest PF. 21 Apr.-l 8 July 1990, DW. RJR (QM
S34443); 9. Rosslyn Mead, 23°10’S, 150°47’E, vine
thicket, PF, 18 July-23 Oct. 1990. DW, RJR. K. Williams
(QM S25580); 2d, same data (QM S25589); 2d, same
locality, PF, 20 Apr.- 18 July 1990, DW, RJR (QM
S34246); 2d, Bondoola. 23°1 PS, 150°4PE, open forest
PF. 20 Apr.-July 1990, DW, RJR (QM S22I96); 9, same
locality, PF, 18 July-23 Oct. 1990, DW, RJR. K. Williams
(QM S46237): d. Nob Ck, 23 Sept. 1993, DW (QM
S34427); d, Olsen’s Caverns, 23"10’S, 150°28’E, open
forest PF, DW. RJR. K. Williams (QM S41239); 2, Mt
Archer, 23“20’S, I50“35’E, 650m, open forest. PF, 4
Sept.- 11 Nov. 1991. DW. RJR (QM S46022); 9, Mt
Chalmers, 23°I0'S. 150°38’E. open forest, PF, Oct.

1 990-2 1 Mar. 1991, D W. RJR. K. Wiliams (QM S2455 1 );
d. Fairlies Knob, 25"32’S, 152°19'£. 120m. vine scrub,
PF, 21 July-20 Oct. 2000, DC, J. Wright E. Vanderduys,
9463 (QM S55257); 9, Teewah Ck. Cooloola. 25°57’S,
153“06'E, 14 Sept 1973, RJR (QMS55245); 9, Cooloola,
rainforest, under rotten log, 21 Aug. 1970, E.C. Dahms
(QM S55246); d, 9, Cooloola, high dune, PF, Oct 1978,
C. Plowman (QM S55248); 9, 7d, Orchid Beach, Fraser
I„ 24°58’S, 153°19’E, PF, 20 Aug.- 17 Dec. 1997, RJR. P.

NEW SPIDER FROM EASTERN AUSTRALIA

243

Fishbum, P. Lawless (QM S43420); 3, Fraser I., Feb.
1971. ANU (AM KS3484I); <3, 39, Fraser I., Orchid
Beach, 24°58’S, 153°18 , E, PF, 20 Aug.- 17 Dec. 1997,
RJR. P. Lawless (QM S43453); 3 <3, 9, same data (QM
S4 1 776); 3 3 , 3 9 , Fraser I., heath, Banksia , eucalypt, PF, 7
Mar.- 1 Oct. 1996, RJR (QM S3 1275); 3, Mt Coot-tha,
27°29’S, 152°57’E, PF, 17Dcc. 1996. RJR(QM S41181);
9, Slaughter Pk, Mt Coot-tha, 22 May 1978, VED. RJR
(QM S55261); 9, Mt Coot-tha, 1 Sept. 1973, RJR (QM
S55262); 3. Mt Nebo, 27°24'S, 152°47 , E, PF, mixed
sclerophyH 16 Oct. 1978, A. Rozefelds (QM S55263); (3,
Mt Glorious, 2?*20*S, 152°45'E, sieved litter, 20 June
1991, D. Black (WAM 98/2101); 43. Cooloola Village,
26°00'S, 153WE, PF, 8 July-4 Dec. 1998, RJR, P.
Lawless (QM S53390); 3, Camerons Scrub. 27°30'S,
152°44 , E, 90m. PF. 7557, vine scrub, 1 1 Nov. 98-13 Jan.
1999. GBM, DC, GT (QM S45715); 3, Bunya Mts,
Paradise, 26°52‘S, 151°35*E, pyrethrum pine trunks 5026,
GT (QM S45672); 3, Marlaybrook. Bunya Mts,
semi-evergreen thicket, 1-6 Mar. *1976, RJR, VED (QM
S45673); 9 , Kroombit Tops, open forest with Casuarimi ,
on sand 23-25 Feb. 1982, GBM, RJR. GT (QM S45686);
3, Calliope Ra. Kroombit Tops, 24°22’S, 151WE, PF 13.
open forest, 12-18 Dec. 1983, GBM, VED, GT, J. Gallon
(QM S55264); 23,9, same data, PF 1 4 (QM S55265); 9 ,
Jimna SF, 26‘'40'S, 152°28’E, under logs. 31 May 1978, K.
McDonald, S. Crafter (QM S45706); 103, top of
Blackbutt Ra., via Benarkin, 26 U 52’S, 152°12’E, PF16,
rainforest 396m, 1 7 Aug.- 1 0 Nov. 1 974, GBM, SRM (QM
S55268); 43, same data (QM S55269); 3, Base of
Blackbutt Ra., 26°52'S, 1 52°1 1 'E, 240m, PF 1 5, rainforest,
17 Aug.- 10 Nov. 1974, GBM, SRM (QM S55270); 29,
Fairlies Knob. 25°31'S, I52°17 , E, 300m, PF9974, hoop
pine scrub, 20 Dec. 2(XX)-22 Mar. 2001. DC. GBM (QM
S55271 ); 33,49, same locality, PF9464, vine scrub, 21
July-20 Oct. 2000, DC, J. Wright, E. Vanderduys (QM
S55272); 3 . same data, 120m, PF9463 (QM S55273); 23,
One Tree Hill, 25°17 > S, 151°55’E. 18()m, vine scrub,
PF9000, 27 Sept.- 14 Dec. 1999, DC (QM S55274); 9,
Nipping Gully, 25°4LS, 151°25’E, 150m, PF7259. open
forest site 3, 27 Aug.-9 Oct. 1 998, GBM (QM S55275); 3 ,
same locality, 240m, PF 7707, 26 Jan.-2 June 1999, GBM,
GT (QM S55276); 2 3 , “Stockhaven” 25°48'S, 151°59'E.
450m, rainforest, PF 75 1 5. 1 0 Oct.- 1 9 Dec. 1 998, GBM, C.
Gough (QM S55277); 3, Nangur SF, 26°08'S, 151°59’E.
site 2. 320m, PF, rainforest, 29 July-23 Oct. 1995, GBM
(QM S42320); 3, Yarraman, 26°50’S, 152°03'E, 520m,
rainforest, PF 17, 17 Aug.-lO Nov. 1974, GBM, SRM
(QM S55278); 33, McAfees Lookout 27°26 , S,152 0 52'E.
1 50m, PF7825, wet scleroplyll, 6 July- 1 8 Oct. 1 999, GBM
(QM S55280); 23,39, Camira, 27°38 , S,152°55'E, open
forest, PF, 30 July-Oct. 1990. RJR (QM S19622-3); 3,
Camira, under log, 22 Jan. 1997, RJR (QM S55256); 9.
Rochedale SF, 2734 > S,153°08 , E. PF. 1-23 Oct. 1979.
VED, RJR (QM S55282); 3, same locality and collectors,
PF, 20 Sept.- 1 1 Oct 1979 (QM S55283); 9, 17 Jan. 1980
(QM S55285); 9 , PF, 30 Aug.-6 Sept. 1979(QMS55286);
3, Bulburin Barracks, 24°32 , S.l 51°28*E, 580m,
rainforest pyrethrum 7816. 8 Oct. 1999, GBM (QM
S45680); 2 9 , Bulburin SF, 540m, small sheet webs, 25-28
March 1977, RJR, VED (QM S45676); 3, top of range.

Bulburin SF, 24°33 , S,15I°33'E, 600m, PF33, 5 Oct-30
Dec. 1 974, GBM, SRM (QM S45679); 9 , with egg sac,
Bulburin Forest nursery, 24 0 3TS,15 1°29'E, 580m,
rainforest under log, 21 Mar. 1975, MRG, C. Horseman
(AM KS7I038); 3, Rundle Ra., 23°39 , S,150°59’E,
semi-evergreen vine thicket, 30m, 24-31 Mar. 1975, R
Kohout, VED (QM S45704); 123, 129, Mt
Goonancman. 25°26'S,152°18'E, litter, 3-7 Nov. 1980,
RJR, VED (QM S45701); 43, 9, Rocky Pt, 10km S
Round Hill l id, 24°1 4’S, 1 5 1 °56'E, 60m, PF7 1 , 28 Aug.- 1 5
Dec. 1976, GBM, SRM (QM S45699). OTHER
MATERIAL. New South Wales: 9,3, Carrai SF nrCarrai
Bat Cave. 31 o 01’S, 152°20'E, Jan. 1993, MRG (AM
KS34522).

DIAGNOSIS. In males the embolus arises
retro-posteriorly whereas it is retrolateral in B.
booloumba and antero-retrolateral in B.brooyar,
palpal patellal apophysis blunt, intumed and
shorter than B. booloumba and B.broovar in
which it is digitiform. Sclerotised base of
epigynal atrium is bluntly rounded whereas it is
sharply pointed in B. booloumba.

DESCRIPTION. Male . CL 2.0, AL 2.0. Ratio of
AME: ALE: PME: PLE is 6: 10:9:10. Legs 4123.

I, 8.4; II, 7.0; III. 6.3; IV, 8.5. Notation of spines.
Femora:!, D110, POOL II. D1 1 1, POOL R001 ; III,
D110, P011, R001; IV, D110, POOL R001.
Patellae: 1 1 1, D00 1 . Tibiae: I, P 1 1 L V222, R0 1 1;

II, D001, POOL V221, ROIL III, P011, V222,
R011; IV, Pill, V212. R 1 0 1 . Metatarsi spined
with distal whorl of 5 spines.

3 palp (Figs 2G-I. 4D). Embolus arises
retro-posteriorly encircles tegulum to a distal
curve postero-retro laterally. Patellal apophysis
short, blunt, slightly incurved.

Males ranged in size from 3. 1-4.0.

Female CL 2.2 AL 2.3, Ratio AME: ALE: PME:
PLE is 7:11:10:11. Legs 4123. 1, 8.2; 11,6.9; III,
6.3; IV, 8.3. Notation of spines similar to 3 but
fewer on ventral tibiae: III, VI 20; IV, VI 20.
Epigynum (Fig. 2J,K) sclerotised base of atrium
short with blunt-rounded apex. Small lateral teeth
present.

Females ranged from 3. 1-4.9 (average 4.0).

Scanning electron micrograph of the foveal
region (Fig. 2D) revealed a pattern of dark spots
which suggested pheromone production,
however these were also found in the male.
Further investigation of the pale areas near the
fovea did not show any pores.

DISTRIBUTION. B. yeppoon (Fig. 8 A) has been
found from mid-eastern to southeastern
Queensland. A male and female were collected

244

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. A-D, Barahna spp. nov. A-C, B. booloumba spinnerets; A, 9 ALS (left); B, d ALS (right), arrow to
nubbin; C, 9 PMS, arrows to paracribellar spigots; D, B. yeppoon, d palp, arrow to tegular apophysis.

from near the bat caves in Carrai SF, New South
Wales.

Barahna taroom sp. nov.

(Figs 5A-E, 8C)

ETYMOLOGY. From the type locality, Taroom.

MATERIAL. HOLOTYPE. d , Taroom, 9km N, southeast
Queensland, 25°27’S, 150°08’E, PF, June-1 1 Sept. 1996. P.
Lawless, H. Janetzki, D. Potter (QM S36452).
PARATYPES. Queensland: 9, Taroom. Dawson R.,
Nathan Gorge, riverine forest, PF. 1 2 Sept.- 1 3 Jan. 1 997, P.
Lawless (QM S370I9); d 9, same data (QM S37024); 9,
same data (QM S37034); 4 d (QM S37049); 3d, Taroom
District. Nathan Gorge, riverine forest. PF107. 13 Nov.
96-13 Jan. 1997, P. Lawless (QM S36615); 2 <3, same
locality, PF010. 15 June- 12 Sept. 1996, P. Lawless (QM
S37265); 4 d , same data (QM S36860); 2d , same locality,
PF 1 07, 1 3 Nov. 96- 1 3 Jan. 1 997, P. Lawless (QM S37076);
58, 9, same data (QM S37096); 8. Taroom District,
25°25’S, 150°0rE, Boggomoss no. 19, PF105, 18 Junc-9
Sept. 1996. P Lawless (QM S368I1). 5, Mt Zamia,
24°06’S, 148°05’E, vine scrub, 360m. PF9818, 27 Oct.-17
Dec. 2000, DC, GBM (QM S55215); d, Mt Gavial,
23°36’S, 150°29’E, open forest, 400m, PF7486, 17 Dec.

98-14 Mar. 1999. DC (QM S55217); d, Mt Hopeful,
23°45'S, 1 50°32’E, vine scrub, 440m. PF901 2, 27 Sept.- 1 5
Dec. 1 999, DC (QM S552 1 8); 2 d . Mt Gavial, 450m, open
forest; PF7488, 18 Dec. 98-14 Mar. 1999, DC (QM
S55224); 9, Mt Gavial, 320m. vine forest, PF7489, 17
Dec. 98-14 Mar. 1999, DC (QM S55223); 9. Mt Gavial,
400m, open forest, PF7687, 14 Mar.-28 June 1999, DC
(QM S55222); 4d, 9, Nipping Gully, site 1, 25°41’S,
1 5 1 °26'E, 200m, rainforest PF7397, 9 Oct.- 1 8 Dec. 1 998,
GBM. C. Gough (QM S55225); d. Nipping Gully, site 5,
200m, rainforest, PF7503, 9 Oct-18 Dec. 1998, GBM, C.
Gough (QM S55230); d. Nipping Gully, site 5, 300m,
rainforest intercept trap 7705, 26 Jan.-2 June 1 999, GBM,
GT (QM S55229); d, Wctheron, 25°34'S, 151°4PE,
150m, vine scrub. PF7509, 10 Oct- 1 9 Dec. 1998, GBM,
C. Gough (QM S55228); d. East Woodmillar, 25°4LS,
151°36’E, 250m, vine scrub, intercept trap, GBM, C.
Gough (QM S55231); d, 9, Yeppoon, Meudeville Hill,
23°06'S. 150°43'E, vine thicket PF. I8Julv-23 0ct 1990,
DW, RJR, K. Williams (QM S55279); d. same locality
and collectors, PF, 21 April- 18 July 1990 (QM S34444);
3d, 9, same data (QM S34448); 39, Olsens Caverns,
23° 1 0'S, 1 50°28’E, open forest PF, 21 Apr.- 18 July 1990,
DW. RJR (QM S40767); 9, Wonga Hills, 26°03’S,
150°49’E, site 4, 470m, brigalow, PF 10249, 10 Oct.- 11
Dec. 2002, GBM, DC (QM S55288). OTHER

NEW SPIDER FROM EASTERN AUSTRALIA

245

FIG 5. A-K, Barahna spp. nov. A-E, B. taroom; A-C, 6 palp (ventral, retrolateral, dorsal); D,E, 2 epigynum
(ventral, dorsal cleared); F-K, B. scoria ; F-H, 2 epigynum (ventral, dorsal cleared, lateral); I-K, o palp (ventral,
retrolateral, dorsal).

MATERIAL: 4(5, 22, Lower Dry Ck, Kroombit Tops,

45km SSW Calliope, 24°23’S, lSO^'E, 720m,
microphyll rainforest, PF, site 10, 13-18 Dec. 1983, GBM,
VED, J. Gallon, GT (QM S55242); 5 c5 , 2 2 , same locality
and collectors, 700m, PF, site 9, 11-18 Dec. 1983 (QM
S45685); 39, Calliope Ra., Kroombit Tops, 24°22’S,
151°00’E, 880m, open forest, site 13, 12-18 Dec. 1983,
GBM, VED, J. Gallon, GT (QM S55244); 2c5, The Caves,
23° 1 1 ’S,150°28'E, open forest, PF, 21 Apr.-18 July 1990,
RJR, DW (QM S40757); 9, same locality, PF, 21 Mar.
1991, RJR, DW, K. Williams (QM SI 9579).

DIAGNOSIS. Embolus arises antero-
retrolaterally, unlike B. yeppoon, which has
posterior origin, distally it is gently curved (Fig.
5B), unlike B booloumba which is curled. <3
palpal patellal apophysis is truncated (Fig. 5C.),
unlike the other species. Basal sclerotisation of
epigynal atrium is rounded anteriorly and longer
than B . yeppoon , not pointed like B. booloumba ;
dorsal foveal protuberances reduced.

DESCRIPTION. Male. CL 2.2, AL 2.2. Ratio of
eyes AME: ALE: PME: PLE is 6:8:9:10. Legs
4=123. 1, 9.8; II, 8.0; III, 7.2; IV, 9.8. Notation of
spines differs from B. booloumba : Femora: II,
P011, ROIL Tibia: III, V221, R011; IV, D001,
VI 11, Rill.

6 palp (Fig. 5A-C). Embolus arising
antero-retrolaterally; membraneous conductor
broadening distally, tegular apophysis obscured
by conductor. RTA digit i form; ventro-retrolateral
apophysis blunt, sclerotised, intumed. Patellal
apophysis short, truncate.

Other males were smaller, ranging from 3. 2-3. 8
in length.

Female . CL 1 .5, AL 1 .7. Legs 4123. 1, 4.9; II, 3.9;
III, 3.6; IV, 5.0. Notation of spines differs from B.
booloumba. Femora: I, D100; III, D001; IV,
D100. Tibiae: P001, V010; III, P101, V011,
R 1 0 1 ; IV, P101, V011, R101. Epigynum (Fig.

246

MEMOIRS OF THE QUEENSLAND MUSEUM

5D,E). Basal sclerotisation of atrium is long and
rounded anteriorly. Insemination ducts and
spermathecae similar to B booloumba.

Females ranged in size from 3.2-3. 6.

DISTRIBUTION. B. taroom (Fig. 8C) has been
collected from mid-eastern to southeastern
Queensland.

Barahna scoria sp. nov.

(Figs 5F-K, 8C)

ETYMOLOGY. From the type locality Mt Scoria.

MATERIAL. HOLOTYPE. <J, Mt Scoria, southeastern
Queensland, 24°32’S, 150°36’E, 200m, vine scrub, PF
9468. 29 June-2 1 Oct. 2000, DC. J. Wright, E. Vanderduys
(QM S55212), PARATYPES. 29, same locality as
holotype, PF 9980, 20 Dec. 2000-23 Mar. 2001, DC, GBM
(QM S55214); d, same data (QM S55213); 3d, same
locality, PF9452, 28 Mar.-29 June 2000, DC (QM
S55216).

DIAGNOSIS. Embolus arises antero-
retrolaterally, encircles tegulum and is gently
curved distally like B. taroom and unlike other
species. The sclerotised atrial base is low with a
small pointed apex unlike B. taroom where the
apex is thin and rounded.

DESCRIPTION. Male . CL 1.3, AL 1.3. Ratio of
AME: ALE: PME: PLE is 5:8:7:8. Legs 4123. 1,
5.0; II, 4.0; III, 3.6; IV, 5.2. Notation of spines.
Femora: I, Did, P001; II, D100. POOL R001;
III, D100, POOL R001; IV, D100, POOL ROOL
Patellae: I, D001; II, D001; IU, D001; IV, D001.
Tibiae. I. P011, V121, R001; II, P011, VI 12,
R01 1 ; III, D001, P011, V012, ROIL IV, D001,
P0 1 1, V 1 1 2, R0 1 1. Metatarsi spined with whorl
of 4-5 spines distally.

8 palp (Fig. 5I-K). Embolus emerging
antero-retrolaterally from loop of sperm duct;
slightly curved distally. Ventro-retrolateral tibial
apophysis blunt and sclerotised, RTA short,
digitiform. Patellal apophysis short, diagonally
truncated.

Other males ranged from 2. 5-2.9.

Female. CL 1 .3, AL 1 .5. Legs 41 23. 1, 4.3; II, 3.5;
III, 3.2; IV. 4.6. Notation of spines. Femora: I,
POOL II, D 1 00; III, D100; IV, D100, ROOL
Tibiae: I, V020; II, V010; III, P011, ROIL IV,
D001, P001, ROOL Metatarsi spined with whorl
of 4-5 spines distally. Epigynum (Fig. 5F-H)
sclerotised base of atrium low with small pointed
apex.

The other female was 2.7 in length.

DISTRIBUTION. B. scoria (Fig. 8C) has been
collected from only one locality, Mt Scoria.

Barahna toonumbar sp. nov.

(Figs 6A-E, 8B)

ETYMOLOGY. From the type locality, Toonumbar SF.

MATERIAL. HOLOTYPE. d, Toonumbar SF via
Grevillea, northern New South Wales, 28°28'S, 1 52°43’E,
PF 23, 31 Mar.-2 Aug. 1975, GBM, SRM (QM S55259).
PARATYPES. New South Wales: d, same data (QM
S55260); d, 9. Border Ranges NP, Tweed Range Rd,
28°25'S. 153°02'E. 470m, 4 Feb.-9 Apr. 1993, M, Gray, G
Cassis (AM KS35945); 9,samedatn(AM KS35950);39,
Whian Whian SF, Running Rd. 28°36'S, l53°2rE, 250m,
PF, 4 Feb.-9 Apr. 1993, MRG G Cassis (AM KS35917);
29, Whian Whian SF, Mackays Rd, 420m. PF, MRG G
Cassis (AM KS35918).

DIAGNOSIS. Embolus shorter than that of
previously described species; it arises
postero-prolaterally ending mid-retrolaterally.
Sclerotisation of base ofepigynal atrium differs
from previously described species; it is squarish
and flat-topped.

DESCRIPTION. Male. CL 1.5, AL 1.4. Ratio of
AME: ALE: PME: PLE is 6:9:8:8. Legs 4123. 1,
4.3; II, 3.6; III, 3.3; IV, 4.9. Notation of spines.
Femora: I, D101, POOL II. DO 1 L POOL R001;
III. D110, POOL R00L IV, D100. P001, ROOL
Patellae: III, D001; IV, D001. Tibiae: 1, D001,
P011, R011; II, D00L PI01, Vlll, R101; III,
D00 1, P 1 0 1, V002, R 1 0 1 ; IV, D 1 0 1, P 1 0 1, V 1 1 2,
R0 1 1 . Metatarsi spined with whorl of 4-5 spines.

8 palp (Fig. 6A-C). Embolus emerges postero-
prolaterally; it describes about three-quarters of a
circle around the tegulum to rest on the
membraneous conductor; tegular apophysis
absent. The ventro-retrolateral apophysis intums
acutely to a point. RTA pointed. Patellal
apophysis short.

Other two males 2.5 and 2.8 in length.

Female. CL 1. 1, AL 1 .5. Legs 4123. 1, 3.4; II, 2.9;
III. 2.6; IV, 3.6. Notation of spines. Femora: I,
D100, POO 1 ; IL D 1 00; III, D 1 00, ROOL IV, D100,
ROOL Patellae: II, D001; III. D001; IV, D001.
Tibiae: I, V00 1 ; II, POO 1, V001 , R00 1 ; III, D00 1,
P101, V01 1, R101; IV, D100, P100, V00LR001.
Metatarsi spined with whorl of 4-5 spines
distally. Epigynum (Fig. 6D,E) basal sclerotised
area flat-topped.

Other females ranged from 2. 3-3.5.

DISTRIBUTION. B. toonumbar (Fig. 8B) has
been collected from the Border Range area of
northern New South Wales.

NEW SPIDER FROM EASTERN AUSTRALIA

247

FIG 6. A-L. Barahna spp. nov. A-E, B. toonumbar, A-C, d palp (ventral, retro lateral, dor )>•’ P '

(ventral, dorsal cleared); F-K. B. myall, F-H, d palp (ventral, retrolateral, dorsal); I-K. $ epigynum (posterior,
ventral, dorsal cleared); L,M, Barahna glenelg, 9 epigynum (ventral, dorsal cleared).

Barahna myall sp. nov.

(Figs 6F-K." 7B,C, 8B)

ETYMOLOGY. From type locality. Myall Lakes NP.

MATERIAL. HOLOTYPE. d, Mvall Lakes NP. New
South Wales, 32°38’S, 152"12’E. PF, 10 Oct. 1997. L.
Wilkie (AM KS71037). PARATYPES. N.S.W: 2d, 22,
Smiths Lake. Myall Lakes, 32°23'S. 152“30’E, dry
sclcrophyll. 8 May 1974, L. Blyton (AM KS34835): 2
(tracheal tubes dissected) same data (AM KS34835); d.
Myall Lakes, same data as holotype(AM K.S71036); 9, 1
June 1997 (AM KS71035); 3d, 2 , same data as holotype
(AM KS62448); 9 (AM K.S62449); 9, with egg sac.
Upper Allyn, 32°22’S, 151°32 , E, MRG (AM KS34837);

Gloucester, 56km W on Barrington Tops Forest Rd,
r oi 'S 1 51'22'E, 1 7 Mar. 1982, MRG H. Pamaby (AM
S8829) <J, Kerewong SF nr Lome, 31“36’S, 152°34'E,
F coll. 1 5 Julv 1 979, D. Milledgel AM KS5404); 2 , same
ala (AM KS7 1034). d , Lome SF, 31 0 35’S. 152°57'E, PF,
all 4 Nov. 1979, D. Milledge (AM KS5599); o, same
ata (AM K.S5641 ); 29, same data (AM KS5667); 2d,
tine location, coll. 30 July 1 979 (AM KS 162 14); 9 same
ata (AM KSI6219); 9. Dorrigo NP, Domgo-Belhngen
d , 30"24'S. 152 n 45’E, PF, 18 Feb. 1993, MRG G Cassis
\M KS35767). 2, Kuring-gai NP. Bobbin Head ( W side),
3‘39'S. 151"09‘E, 10 Apr. 1974, MRG (AM KS34832);
? , same locality. 10 May 1974, MRG (AM KS34833), 2,
ame locality, 2 Nov. 1993, MRG (AM KS34834); 2d, 2 ,
jvs. Maria SF nr Kempsey, 3 HI’S, 152°50’E, in logs,

248

MEMOIRS OF THE QUEENSLAND MUSEUM

FIQ 7. A-D, Barahna spp. nov. foveal areas. A, 9 B. booloumba ; B, 9 B. myall ; C, 6 B. myall ; D, 9 B. glenelg.

Eucalyptus gummifera (Bloodwood forest), 23 Apr. 1974,
MRG (AM KS34836); 9 , Bulls Ground SF, nr Wauchope,
31°35 , S, 152°4rE, PF, 10 Feb. 1991. A. York, NSW
Forestry (AM KS43484); 6. Kiwarrak SF, nr Taree,
31°58’S, 152°26’E,PF coll. 28 Sept. 1979, W.C. King (AM
KS3943). 29, Glen Dhu Cave Timor, 31°34 t S, 15l°27’E,
cave, 25 May 1995, S. Eberhard (AM KS5701 1).

DIAGNOSIS. Larger than other species, seldom
less than 5.0 in length. Embolus arises
postero-pro laterally and terminates
mid-retrolaterally as in B. toonumbar and unlike
other species. Patellal apophysis very long and
bifid unlike B. toonumbar. Sclerotised base of
atrium is flat and square like B. toonumbar.
Spermathecae larger than other species.

DESCRIPTION. Male. CL 3.0 AL 2.4. Ratio of
AME: ALE: PME: PLE is 10:13:12:13. Legs
4123. 1, 10.5; II, 8.8; III, 8.0; IV 1 1 .0. Notation of
spines. Femora: I, D101, P002, R001; II, D101,
P011, R011; III, Dill, P011, R011; IV, D110,
POOL Patellae: II, D001; III, D001. Tibiae: I,
PI 10, V202, R011; II. P011, V222, R011; III.
D00 1 , P0 1 1 , V222, R0 1 1 ; I V, D 100, P 1 0 1 , V222,
ROIL Metatarsi spined with distal whorl of 5
spines.

6 palp (Fig. 6F-H). Embolus arises posteriorly
describing about three quarters of a circle round
the tegulum to rest on membraneous conductor.
Tegular apophysis absent. Short, blunt
ventro-RTA, digitform RTA. Bifid patellal
apophysis, longer than RTA.

Other males ranged from 3.6-5. 6.

Female. CL 2.4 AL 2.8. Ratio of AME: ALE:
PME: PLE is 9:13:11:14. Legs 4123. I, 7.9; II,
6.4; III, 5.7; IV, 8.5. Notation of spines. Femora:
I, D101, P002, R001; II, D101, P011, R011; III,
Dill, P011, R01 1; IV, D110, POOL Patellae. II,

D00 1 ; III, D00 1 . Tibiae: I. P 1 1 0, V202, R0 1 1 ; II,
P011, V222, R0 1 1 ; 1 1 1 , D00 1 , P0 1 1 , V222, R0 1 1 ;
IV, D100, P101, V222, ROIL Metatarsi spined
w'ith whorl of 5 spines distally.

The foveal area is almost flat and the associated
pale areas dorsal rather posterior (Fig.7B);
further investigation did not show any pores. The
male lacks the swellings and pale areas (Fig. 1C).

Epigynum (Fig. 6I-K). Basal sclerotised area
of atrium flat-square; spermathecae large. Four
simple tracheal tubes in abdomen, medians
longer than laterals (dissection courtesy of
M.R.Gray).

Other females ranged from 5.2-6.5.

DISTRIBUTION. B. myall <Tig.8B) was
collected from eastern New South Wales.
Females from Kuring-gai NP near Sydney which
are placed in this species need further
confirmation when males are available.

Barahna glenelg sp. nov.

(Figs 6L,M, 7D, 8D)

ETYMOLOGY. From the type locality, Glenelg R.

MATERIAL. HOLOTYPE. 9, Glenelg R, Dartmoor,
Victoria, 37°55’S, 141°16’E, under logs at base of
limestone outcrop, 25 Mar. 1 974, MRG (AM KS034839).

DIAGNOSIS. Foveal protuberances broader
than other species. Basal sclerite flat-topped like
B. toonumbar and myall but differing from these
in being longer than wide.

DESCRIPTION. Female. CL 2.5, AL 2.9. Ratio
of AME: ALE: PME: PLE is 8: 1 0:9:9. Coloration
and pattern similar to other species. Foveal
protuberances occupying a wider area separating
foveal groove (Fig. 7D). Legs 4123. 1, 7.2; II, 6.5;

NEW SPIDER FROM EASTERN AUSTRALIA

249

FIG. 8. A-D, distribution maps of Barahna spp. nov. A, B. yeppoon; B, B. booloumba, B. brooyar , B. myall , B.
toonumbar . ; C, B. scoria , B. taroom\ D, B. glenelg.

250

MEMOIRS OF THE QUEENSLAND MUSEUM

III, 5.9; IV, 7.8. Notation of spines. Femora: I,
D101, P002, R001; II, D10I, POOL R00I; III,
Dill, P011, R001; IV, D 1 0 1 , POOL R001.
Tibiae: I, P011, V221, R000; II, P011, V202,
ROI 1; III, D001, P101, VI 22, R101; IV, D001,
P 101, VI 12, RIO 1. Metatarsi spined with distal
whorl of 4-5 spines.

Epigynum (Fig. 6L.M). Basal sclerite of atrium
longer than wide, flat-topped.

The male is unknown.

DISTRIBUTION. One female B. glenelg (Fig.
8D) has been collected from Glenelg R., near
Dartmoor, Victoria.

ACKNOWLEDGEMENTS

I am indebted to curators and collection
managers of the Australian, West Australian and
South Australian Museums for loans of material

for this study, and especially to colleague Dr G.B.
Monteith and his co-workers for collections from
mid-eastern and southeastern Queensland and
northern New South Wales. I thank the Council of
the Australian Biological Resources Study for its
funding of rainforest surveys during which some
of these spiders were collected and for financial
support for illustrators Catherine Harvey and Dr
Christine Lambkin and SEM technician Kylie
Stumkat. I am indebted to Dr Robert Raven for
construction of the distributional maps.

LITERATURE CITED

COMSTOCK, J.H. 1 9 1 3. The spider book. (Doubleday,
Page & Company: New York).

PLATNICK, N.I. & SHADAB, M.U. 1975. A revision
of the spider genus Gnaphosa (Arancae:
Gnaphosidae) in America. Bulletin of the
American Museum of Natural History 155: 1-16.

TANGANA, A NEW SPIDER GENUS FROM AUSTRALIA (AMAUROBIOIDEA:
AMPHINECTIDAE: TASMARUBRIINAE)

VALERIE TODD DAVIES

Davies, V.T. 2003 06 30: Tangana , a new spider genus from Australia (Amaurobioidea:
Amphincctidae: Tasmarubriinae). Memoirs of the Queensland Museum 49(1): 251-259.
Brisbane. ISSN 0079-8835.

Six species of Tangana gen. nov. are described. They arc T. greeni (type species), T. collina ,

T. mepartlan , T. acuta and T. clarkei all from Tasmania, the island state off the southeast coast
of Australia and T. harveyi from Victoria, mainland Australia. They are placed with
Tasmarubrius and Tasmabrochus in the subfamily Tasmarubriinae. □ Amaurobioidea,
Amphincctidae, Tangana, new genus, Australia.

Valerie Todd Davies, Queensland Museum, PO Box 3300 South Brisbane 41 01, Australia;

9 May 2003.

The Tasmarubriinae was diagnosed and
described by Davies (2002). Like the other
genera, Tangana spp. are large (8.0 or longer)
ecribellates found in small retreats under logs or
rocks. Some species were sympatric, even found
in the same pitfall sample. The only modification
from the description (loc. cit.) concerns the
curvature of the eye rows — view r ed from above
the anterior row is slightly recurved, the posterior
row straight to slightly procurved (Fig. ID). It is
thought that the broad embolus conveys not only
the sperm duct but also the duct carrying the
acellular glandular secretion that plugs the female
gonopores (Suhm, Thaler & Alberti, 1996).

MATERIAL AND METHODS

All but one of the following taxa were collected
from sites in Tasmania, the exception being a
single female from Victoria. Much of the
material, comprising about 50 vials from Maggs
Mt, NW Tasmania was collected by R.H. Green.
To avoid repetition in listing these, after the first
record the collector’s name and locality name are
omitted. Pitfalls (PF) were the main trapping
method, with some hand-collecting. Notation of
spines follows Platnick & Shadab (1975). The
left male palp is described and illustrated.
Measurements are in millimetres. The epigyna
vary little between species thus there is an
emphasis on male characters and males only are
used in the key.

ABBREVIATIONS. Museums: AM, Australian
Museum, Sydney; QM, Queensland Museum,
Brisbane; QVM, Queen Victoria Museum,
Launceston; SAM, South Australian Museum,
Adelaide; TM, Tasmanian Museum and Art
Gallery, Hobart; WAM, West Australian
Museum, Perth. Morphology: The usual

abbreviations are used for body measurements
and eyes. MA, median apophysis; RTA,
retrolateral tibial apophysis. Other abbreviations
on illustrations are explained in the text or in the
legends.

SYSTEMATICS
Tangana gen. nov.

TYPE SPECIES. T. greeni sp. nov.

ETYMOLOGY. From the Aboriginal ‘tangana’, meaning
spider in the Tasmanian language.

DIAGNOSIS. Males differ from other tas-
marubriines ( Tasmarubrius and Tasmabrochus)
in lacking a large fixed prolateral tegular
apophysis and in having a terminally branched
median apophysis. Lateral teeth are present on
epigynum. Spines on both male and lemale tarsi
III and IV, smaller than those on other leg
segments and not present in other tasmarubriines.

DESCRIPTION. Large, 3-clawed ecribellates
with geniculate chelicerae, 2 retromarginal and 2
promarginal cheliceral teeth with an intermediate
row of tiny thin denticles. Preening combs on
distal metatarsi I (a few tines), II, III and IV. Short
thick embolus, small membraneous conductor
and movable median apophysis. A small tegular
bulge may be present between embolic base and
median apophysis. Small proximal paracymbium
usually present. Palpal tibia with long ridge-like
distal retroventral apophysis and an excavated
retrolateral apophysis with two or more process-
es. Epigynum with lateral teeth. Gonopores often
blocked with acellular material, insemination
ducts heavily sclerotised, large spermathecae.

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MEMOIRS OF THE QUEENSLAND MUSEUM

KEY TO MALE TANGANA. SPP.

1 . Small membraneous tegular apophysis present (Fig. 2C)

greeni

Small membraneous tegular apophysis absent 2

2. Median apophysis trifurcate (with small ventral branch)

acuta

Median apophysis bifurcate (prolateral and dorsal
branches only). * 3

3. Retrodorsal branch of RTA rounded (Fig. 3C) . . collina

Retrodorsal branch of RTA pointed 4

4. Retrolateral branch of RTA with proximal spur (Fig. 4C)

mcpartlan

Retrolateral branch of RTA without proximal spur

clarkei

Male of T. harveyi is unknown.

Tangana greeni sp. nov.

(Figs 1, 2. 6A; Table 1)

ETYMOLOGY. In honour of Robert H. Green, collector of
many Tasmanian spiders.

MATERIAL. HOLOTYPE. d , Maggs Mt NW Tasmania,
41°45’S, 146°12’E, PF. 13 June-19 Sept. 1979, R.H.Green
(QVM 13:39937). PARATYPF.S: Tasmania. 49, same
data as holotype (QVM 1 3:39938); 1 7 d . same data (QVM
13:39939); 3 (QVM 13:39940); d,9, same data, 15 Oct.
1979 (QVM 13:7076); 9, 2d, juvs, 17 Oct. 1979 (QVM
13:7075); 23, 10 Oct. 1979 (QVM 13:7072); 3 , 23 Sept.
1981 (QVM 13:42035); 8 3, ex Poa tussock, dry
sclerophyll 14 Oct. 1980 (QVM 13:42081); 23, 17
Oct. 1979 (QVM 13:42133); 2d, 2, (QVM 13:42137); 3,
42, 14Nov. 1984 (QVM 13:42238); 2.500m from camp,
dry sclerophyll, 19 Nov. 1980 (QVM 13:42274); 3, 26
Nov. 1987 (QVM 13:42427); 2$, W field station, dry
sclerophvU, PF, 8 Oct.-6 Nov. 1 979 (QVM 13:42522); 2,
Aim R Camp, wattle. 5 Feb. 1980 (QVM 13:42719); 2,26
Nov 1987 (QVM 13:42723); 3, 23 Sept. 1985 (QVM
1 3 :42838); 8 <5, 26 Nov. 1987 (QVM 13:42842); <5, 2. wet
sclerophyll, 16 OcL 1979, Foreslry (QVM 13:43635);
142, wet sclerophyll, 800m, 14 Oct. 1980 (QVM
13:43638); 42 3, same data (QVM 13:43640); 23, 26
Sept. 1989 (QVM 13:43641); 23, 7 Sept.1987, Forestry
(QVM 13:43642); 3d, 500m, 14 Oct. 1980 (QVM
13:43643); tJ,25Nov. 1986(QVM 1 3:43650); 43 , 9, 13
Oct. 1982 (QVM 13:43634); 9, 25 Nov. 1986 (QVM
13:43652); 2, 130ct. 1982(QVM 13:43715); 73, lOOct.
1983 (QVM 13:42600); 3, Cradle Mt, Pencil Pine Ck,
41°38’S 145°54’. 16 - 24 April 1980, R.H.Green (QVM
13:43655); 9, Mt Field, 42°41’S, 146°43’E. 200m, under
log, 20 May 1996, LJ. Boutin (TM J3312); 9, 6 , juv.,
8km SW Waratah, NW Tasmania, 4U29'S. I45°27’E,
690m, PF. 17 Aug.-3 Dec. 1990, R. Mesibov (QVM
13:23308); 2, same data, 680m (QVM 13:23315); 3,
670m (QVM 13:23316); d,(QVM 13:23322): 9, (QVM
13:23323); 2, Mt Barrow, NE Tasmania, 41°23'S,
147°25’E, 25 Apr. 1971. R.H.Green (QVM 13:42359); 3,
samedata(QVM 13:42360); 3 , (QVM 13:42364): 2d. nr
Pioneer, NE Tasmania, 41°03’S,148°0rE, Forestry (TM
J3313); 2, NE Tasmania, Nothofagus forest nr
Weldborough, 4 1 ° 1 2 ’ S, 147°54’E, Aug. 1993, P. Cranston,

J. Trueman (QM S55289); 2d, 8km SW Waratah, NW
Tasmania, 41°29'S, 145°27'E, 680m. 20 Sept. 1990, R.
Mesibov (QVM 13:43654); d Black Bog Ck. 870m. 8
July 1997. R. Mesibov (QVM 13:43656); 3, same data,
83()m(QVM 13:43657); d, Central Plateau, Butlers Rd nr
Tarraleah, 700m, 20 Oct. 1992, R. Brereton (QVM
13:43658); d, same data, 18 Aug. 1992 (QVM 1 3:43659);
d , Central Plateau. Homes Dam nr Tarraleah, 750m, 1 992
(QVM 13:43694). OTHER MATERIAL: d, Maatsuyker

I. , S Tasmania. 43°39’S, I46°16’E, Site 3, 10-22 Dec.
1996, S.J. Cronin (QVM 13;23836).

DIAGNOSIS. Median apophysis trifurcate
distally with spur-like prolateral, long dorsal
(largely obscured by trunk) and very short retro-
lateral branches. Small membraneous tegular
projection (apophysis) at retrolateral base of
embolus. Palpal tibia with long blunt ridge-like
distal retroventral apophysis; excavated retro-
lateral apophysis with long digitiform retrolateral
and curved retrodorsal apophyses. The latter with
a flat rounded centro-retrolateral projection (Figs
1C, 2D) connected by a low ridge to the
retrodorsal apophysis.

DESCRIPTION. Male. CL 6. 1 . CW 4.8, AL 5.8,
AW 3.5. Light brown cephalothorax, almost
glabrous. Dorsal abdomen pale with six dark
chevrons with pairs of light spots between them.
Pale venter. From above posterior row of eyes
straight, anterior row slightly recurved. AME
smallest Ratio of AME: ALE: PME: PLE is
7:12:10:11. Chelicerae slightly geniculate, two
retromarginal and two promarginal teeth,
intermediate row of tiny thin denticles (Fig.
2A,B). Labium as wide as long. Sternum a little
longer than wide, 1:0.9. Legs 4123 (Table 1).
Nofation of spines. Femora: I, D 1 1 1 , P0 1 2. R 1 1 1 ;

II, D1 11, Pill, R01 1; III, Dill, P101, R01 1; IV,
D110, P010, R001. Tibiae: I, Pill, V221. R001;

II, Pill. V222, R001; III, D011, Pill, V222,
R 1 1 1 ; I V, D00 1 , P 1 1 1 , V222, Rill. Metatarsi : I,
P001, V222, R001; II, D010, PI 1 1, V221, R01 1 ;

III, D101, Pill, V221, Rill; IV, D110, PI 12,
V221, R 1 1 2. Tarsi: III, P011, ROM; IV, P011,
R010.

TABLE 1: T. greeni d(2) leg lengths.

Leg I

Leg II

Leg III

Leg IV

Femur

6.3 (5.4)

6.0 (4.7)

5.6 (4.5)

6.4 (5.8)

. Patella &
Tibia

8.1 (6.8)

7.2 (6.2)

6.7 (5.4)

8.1 (6.8)

Metatarsus

5.8 (4.4)

5.6 (4.3)

5.8 (4.2)

8.2 (6.1)

Tarsus

3.9 (2.8)

3.6 (2.6)

3.1 (2.1)

3.5 (2.6)

Total

24.1 (19.4)

22.4(17.8)

21.2(16.2)

26.2 (21.3)

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253

FIG 1 . A-F, Tangana greeni sp. nov. A-C, 6 palp (ventral, retrolateral, dorsal);
(ventral, dorsal), cp = central process of RTA; rda = retrodorsal apophysis,
retroventral apophysis of tibia.

D, dorsal carapace; E-F, epigynum
rla = retrolateral apophysis; rva =

6 palp (Figs 1A-C, 2C,D). Cymbium with
bulge on retrolateral edge, narrow neck region
with small rounded proximal projection (para-
cymbium). Thick embolus, small membraneous
conductor, short terminally trifurcate median
apophysis and small blunt membraneous tegular
projection prolateral to it. Tibia with distal
retroventral apophysis and excavated retrolateral

apophysis consisting of digitiform retrolateral
process and curved retrodorsal apophysis. The
latter with a rounded central process connected
by a low ridge to the blunt retrodorsal apophysis.

Males varied in length from 9.2 - 1 1 .5.

Female . CL 5.8, CW 4.0, AL 5.3, AW 3.4. Similar
pattern to male. Ratio of AME: ALE: PME: PLE
is 7:10:9:10. Legs 4123 (Tablel). Notation of

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MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. A-D, Tangana greeni sp. nov. A,B, chelicera; CD, 6 palp (ventral, rctrolateral). c = conductor; e =
embolus; ma = median apophysis; rla = retrolateral apophysis of RTA; rva = retroventral apophysis; ta - tegular
apophysis.

spines. Femora: I, D1 10. P01 1, R01 1; II, Dill,
Pill, R001; III, Dill, P 1 0 1 , ROM; IV, Dill,
P101, R001 . Tibiae: I, Pill, V222; II, P011,
V222; III. D101, Pill, V222, Rill; IV, Dill,
Pill, V222, Rill. Metatarsi: I, POU, V221,
R011; II, Pill, V221, Rill; III. D100, Pill,
V221, Rill; IV, D110. PI 12, V221, R112.Tarsi:
III, P010, R010; IV, P010, ROM.

Epigynum (Fig. 1E,F) about a fifth the length
of venter. Postero-lateral gonopores leading

through heavily sclerotised insemination ducts to
spcrmathecae thence by fertilisation ducts to
uterus. Irregular shaped plugs in some gonopores.

Females varied in length from 9.0 - 14.0.

DISTRIBUTION (Fig. 6A). Found over a wide
area in NW Tasmania and Central Highlands
region and to a lesser extent in NE Tasmania. One
male from Maatsuyker I., in the south, is not
listed as a paratype.

NEW SPIDERS FROM SOUTHEASTERN AUSTRALIA

255

Tangana collina sp. nov.

(Figs 3A-E,6B)

ETYMOLOGY. From Latin ‘collis’, high ground.

MATERIAL. HOLOTYPE. 6 , Maggs Mt. NW Tasmania.
41°45’S, I46°I2’E, PF, 13 June- 19 Sept. 1979, R.H. Green
(QVM 13:39941). PARATYPES: Tasmania. 9 , same data
as holotype (QVM 13:39942): 8cl, same data (QVM
13:39943); 6. 9.7Nov. 1987 (QVM 13: 42722).; 2d, 26
Nov. 1987 (QVM 13: 42843); 26, 20 Sept.1988 (QVM
13: 43636) ; 6. 18 Sept.1984 (QVM 13: 43637): 9, 14
Nov. 1984 (QVM 13: 43644); 9, 23 Feb. 1988 (QVM
13:43645); 9 , 4 Feb. 1 980 (QVM 13:43647); 9, 22 Feb.
1983 (QVM 13: 43648): 29, 9 Feb. 1982 (QVM 13:
43651); 9, II Feb. 1987 (QVM 13:43691); 9, 25 Nov
1986 (QVM 13:43692); 6. 15 June 1984 (QVM 13:
43693); 96, 9, 14 Oct 1980 (QVM 13:39944); 6, 23
Sept. 1981 (QVM 13:39945); 6. 10 Oct. 1983 (QVM
13:39946); 6 , Maggs Mt .Ami R„ Camp, wattle, 7 Sept
1987, Forestry, (QVM 13:39947); 26, Cradle Mountain,
Pencil Pine Creek. 41°38'S, 145°54'E, 16-24 Apr. 1980,
R.H.Green (QVM 13:39948); 9, Upper Natone Forest
Reserve, 4 1 6 * * * * * * 13 1 4’S, 145°53’E, under log, 6 June 1989, J.L.
Robinson ( TM J2825); 9, along Obsidian Ck, Savage
R.,W Tasmania, 41°35’S, 145°08’E, 6 Aug. 1974, R.
Mesibov (TM J102).

DIAGNOSIS. A larger spider than T. greeni. 6
palp with flange-shaped paracymbium (Fig. 3B).
MA terminally bifurcate, lacking the retrolateral
branch. Without tegular projection. Palpal tibia
with ridge-like retroventral apophysis, excavated
RTA with a rounded retrolateral and curved
retrodorsal branch, lacking the central process.
Gonopores slit-like.

DESCRIPTION. Male. CL 7.3, CW 5.3, AL 6.7,
AW 3.7. Similar colour and pattern to D. greeni.
Ratio of AME; ALE: PME: PLE is 9; 12: 12: 14.
Legs 4123. I, 26.3; II, 24.0; III, 22.7; IV, 29.9.
Notation of spines. Femora: I, D 1 1 1 , P0 1 1 , R0 1 1 ;
II, Dill, PI 11, R011; III. Dill, Pill. R021; IV.
Dill, P101, R011. Tibiae: I. PI 1 1, V222, Rill;
II, PUL V 222, Rill; III, Dill, Pill, V222,
Rill; IV, Dill, Pill, V222. Rill. Metatarsi: I,
Pill, V221, R01 1; 11, P011, V 221, Rill; 111.
D110, PI 12, V 221, R 112; IV, D110, PI 12, V
221, R 1 1 2. Tarsi: III, P010, R 010: IV,
POOl.ROll.

6 palp (Fig. 3A-C). Cymbium with bulge on

retrolateral edge, narrow neck region and

proximal paracymbium. Thick embolus,

membraneous conductor, bifurcate MA with

prolateral and dorsal branches only. Tibia with

blunt retroventral apophysis, excavated RTA

with rounded retrolateral and curved retrodorsal

branches.

Males varied in length from 1 1.4 - 14.0.
Female. CL 7.9, CW 5.5, AL 8.8, AW 5.4. Ratio
of AME: ALE: PME: PLE is 9: 12:1 1:12. Legs
4123. 1, 23.3; II, 20.6; III, 20.4; IV, 26.4. Notation
of spines. Femora: 1, D 1 1 1 , POO 1 , R0 1 1 ; II, D 1 1 1 ,
Pill, R011; III, Dill, R012; IV, Dill. P011,
R01 1 . Tibiae: I. PI 1 1 , V 222; II, PI 1 1, V 222; III,
Dill, PI 11, V222, Rill; IV, Dill, Pill. V222,
Rill. Metatarsi: I, P011, V221, R011; II, Pill,
V221. R011: III, D100, Pill, V221, Rill; IV,
D110, PI 12, V221, R112.Tarsi:III, P0I1, R011;
IV, P0II. R011.

Epigynum (Fig. 3D,E ) about an eighth the
length of venter. Slit-like gonopores.

Females ranged in length from 13.5 - 20.0.

DISTRIBUTION (Fig. 6B). Sympatric with T.
greeni at Maggs Mt, NW Tasmania and found
further west.

Tangana mcpartlan sp. nov.

(Figs 4A-E, 6B)

ETYMOLOGY. From the type locality, McPartlan Pass,
SW Tasmania.

MATERIAL. HOLOTYPE: 6, McPartlan Pass, SW
Tasmania, 42°5rS,146°H’E, 200m , PF, Mar. 1999,
M.Dricssen (QM S55290). PARATYPES: Tasmania. 9,
same data as holotype (QM S55291); 9, same data, 12
Mar. 1999 (QM S55292); 6. same data, 21 Mar. 2001,
(QM S55318).

DIAGNOSIS. Large spider. MA bifurcate
distally with long prolateral and dorsal branches,
lacking retrolateral branch. Tegular projection
absent. Palpal tibia with ridge-like retroventral
apophysis, excavated RTA with a rounded
digitiform retrolateral branch with small
proximal process (Fig. 4C): curved retrodorsal
branch with flat rounded process curving to
dorsal spur.

DESCRIPTION. Male. CL 7.1, CW 5.4, AL 6.3,
AW 3.8. Red brown cephalothorax. Abdomen
brown with six paired pale spots. Labium longer
than wide 1:0.7. Sternum longer than wide 1:0.9.
Ratio of AME: ALE: PME: PLE is 8:9:9: 1 0. Legs
4123; 1, 24.5; II, 22.8; III, 22.3; IV, 28. 1 . Notation
of spines. Femora: I, D1 1 1, P01 1, R022; II, Dill,
PI 12, R012; III, Dill, Pill, R021; IV, Dill,
Pill, R011. Patellae: III ,D001; IV, D001.
Tibiae: I, PHI. V222, Rill; II, Pill, V222,
Rill. Ill, Dill, Pill ,V222, Rill; IV, Dill,
Pill, V222, Rill. Metatarsi: I, D110, PI 12,
V221, R112; II, Dill. PI 12, V221, R112; III,
Dill, Pill, V221, R1 12; IV, D1 10, P212,V221,
R212. Tarsi: III, P010, R010; IV, P011, R011.

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MEMOIRS OF THE QUEENSLAND MUSEUM

o

FIG 3. A-E, Tangana collina sp. nov. A-C, 6 palp (ventral, retrolateral, dorsal); D,E, epigynum (ventral, dorsal)
fd = fertilisation duct; pc = paracymbium; sp = spermathccae.

cT palp (Fig. 4A-C). Cymbium with bulge on
retrolateral edge, narrow neck region with small
rounded paracymbium. Thick embolus, small
membraneous conductor, MA terminally
bifurcate. Tibia with ridge-like retroventral
apophysis, excavated RTA.

Female . CL 8.1, CW 5.4, AL 7.3, AW4.5.
Coloration similar to male. Ratio of AME: ALE:
PME: PLE is 7:9:10:11. Notation of spines.
Femora: I, Dill, P011, ROM; II, Dill, Pill,
Rill; III, Dill, Pill, Rill; IV, Dill, Pill,
R011. Patellae: III, D001; IV, D001. Tibiae: I,
P011, V222; II, Pill, V222; III, Dill, Pill.
V222, Rill; IV, Dill, Pill, V222, Rill.
Metatarsi: I, PI 1 1, V221, R001; II, P012, V221,
R012; III, D100, PI 12, V221, R112; IV, D101,

PI 12, V221, R 1 1 2. Tarsi: III, P010, R010; IV
P010, R010.

Epigynum (Fig. 4D,E) about a sixth the length
of venter. Curved, pointed lateral teeth.

Other 9 CL 7.8, CW 5.3, AL 9.7, AW 6.3.

DISTRIBUTION (Fig. 6B). From McPartlan
Pass, SW Tasmania.

Tangana acuta sp. nov.

(Figs 5A-C, 6B)

ETYMOLOGY. From the Latin ‘acutus’, pointed referring
to the sharply pointed tip of the embolus.

MATERIAL. HOLOTYPE: 6 , nr Pioneer, NE Tasmania
41°03’S, 148°01 ’E, PF, Forestry (TM J3314)!
PARATYPES: 2d, same data as holotype (TM J3315).

NEW SPIDERS FROM SOUTHEASTERN AUSTRALIA

257

FIG. 4. A-E, Tangana mcpartlan sp. nov. A-C, <3 palp (ventral, retrolateral, dorsal), D,E, epigynum (ventral,
dorsal). It = lateral teeth.

DIAGNOSIS. Smaller spider with flange-shaped
paracymbium. Short MA, trifurcate distally.
Sclerotised tegular area at base of embolus.
Palpal tibia with retrodorsal apophysis
terminating in a long sinuous spur (Fig. 5B).

DESCRIPTION. Male. CL 5.7, CW 3.7, AL 4.4,
AW 2.9. Red brown cephalothorax, pale
abdomen. Ratio of AME: ALE: PME: PLE is
7: 10:8; 11. Legs 4123. 1, 20.3; II, 18.3; III, 16.7;
IV, 22.9

Notation of spines. Femora: I, D110, P011,
R011; II, Dill, P012, ROM; III, Dill, P011,
R0 1 1 ; I V, D 1 1 1 , P0 1 1 , R0 1 1 . Patellae: III, D00 1 .
Tibiae: I, POO 1 ,V222; II, P 1 1 1 , V222; III, Dill,
Pill, V222, R 1 1 1 ; I V, D 1 1 1 , P 1 1 1 , V222, Rill.

Metatarsi: I, P010, V222, R ^10; II, POU,

ROIL III, DUO, PI 12, V221, R112; IV, Dill,
P112, V221, R112. Tarsi: III, R001; IV, ROIL

<y palp (Fig. 5A-C). Small flange-shaped
paracymbium. MA trifurcate distally with
prolateral, dorsal and small ventral branch. Tibia
with distal retroventral apophysis and excavated
RTA with long digitiform retrolateral branch,
small central process and curved retrodorsal
apophysis terminating in long sinuous spur.

Other males were slightly smaller, 8.8 and 9.3.

Female unknown.

DISTRIBUTION (Fig. 6B). Known only from
one sample collected near Pioneer, NE Tasmania.

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MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 5. A-H, Tangana spp. nov. A-C, I acuta , 6 palp (ventral, retrolateral, dorsal). D-F, T. clarkei , 6 palp (ventral,
retrolateral, dorsal). G H, T. harveyi, epigynum (ventral, dorsal).

Tangana clarkei sp. nov.

(Figs 5 D-F, 6B)

ETYMOLOGY. In honour of Arthur Clarke, speliologist.

MATERIAL. HOLOTYPE: <J, Exit Cave, 43°29’S,
146°51’E, 7 Mar. 1997, A.Clarke (TM J3259). OTHER
MATERIAL: juvenile, same data as holotype (TM J33 1 6).

DIAGNOSIS. Large cave spider with very long
legs. MAbifiircate distal ly.Retrodorsal branch of
RTA bluntly pointed.

DESCRIPTION. Male. CL 8.7, CW 6. 1 , AL 7.9,
AW 5.5. Coloration similar to other species but
without pattern on abdomen. Posterior eye row
straight, anterior row slightly recurved. Ratio of

NEW SPIDERS FROM SOUTHEASTERN AUSTRALIA

259

AME: ALE: PME: PLE is 8: 10: 10: 11. Labium
longer than wide, 1: 0.8. Sternum longer than
wide, 1 : 0.8. Legs 4 123. 1, 30.9; II, 29.3; III, 28.6;
IV, 38.3. Notation of spines. Femora: I, Dill,
P011, R021; II, Dill, PI 02, R011; III, Dill,
Pill. Rill; IV, Dill, Pill, Rill. Patellae: III,
D00 1 ; I V, D00 1 . Tibiae: I, Pill, V222, R0 1 1 ; II,
Pill. V222, Rill; III, Pill, V222, Rill; IV,
Dill, P011, V222, Rill. Metatarsi: I, D010,
Pill, V221, R1 12; II, D1 10, PI 1 1, V222, Rill;

III, D110, PI 12. V22 1 , R 1 1 2; IV, D100, PI 12,
V221, R112. Tarsi: III, POOL R001; IV, P001,
V003, R001.

6 palp (Fig. 5D-F). Without paracymbium.
MA with large dorsal and small prolateral
branches. RTA with short rounded retrolateral
and pointed retrodorsal apophyses.

Female unknown.

DISTRIBUTION. (Fig. 6B). Cave spider from
SW Tasmania.

Tangana harveyi sp. nov.

(Figs 5G,H, 6A)

ETYMOLOGY. In honour of Mark Harvey, one of the
collectors of this spider from Victoria.

MATERIAL. HOLOTYPE. 9, The Beeches, Victoria,
37°28’S, 145°49’E, 25 Mar. 1991, Mark S. Harvey, M.E.
Blosfelds (WAM 98/2046). OTHER MATERIAL. 2
juveniles, same data as holotype (WAM 98/2047-8).

DIAGNOSIS. Epigynal spermathecae more
widely separated than in other species.

DESCRIPTION. Female. CL 7.2, CW 4.8, AL
6.0, AW 4.0 (abdomen shrunken). Red-brown
carapace; dark cardial area and five chevrons
with accompanying pale spots on abdomen; pale
venter. From above, posterior eye row straight,
anterior row slightly recurved. Ratio of AME:
ALE: PME: PLE is 8: 11: 9: 12. Labium slightly
longer than wide 1: 0.9. Sternum longer than
wide 1: 0.8. Legs 4123. 1,21.2; II, 19.7; III, 17.7;

IV, 23.2. Notation of spines. Femora: I, Dill,
P002, R011: II, Dill, Pill, R100; III, Dill,
P101, R01 1; IV, Dill, P101, R001. Tibiae: I,
P011, V222; II, Pill, V222; III, Dill, Pill,
V212, Rill; IV, Dill, Pill, V212, Rill.
Metatarsi: I, PI 01, V221, R001; II, P01 1, V221,
R011; HI, D110, PI 12, V221, R112; IV, D110,
PI 12, V221, R112. Tarsi: III, POM, R001; IV,
POOL R001.

Epigynum (Fig. 5G,H). Spermathecae more
widely separated than in other species.

Male unknown.

DISTRIBUTION (Fig. 6A). One female and two
juveniles from southern Victoria.This is the first
record of a tasmarubriine from mainland
Australia.

ACKNOWLEDGEMENTS

1 thank the curators and collection managers of
the Australian Museum, the Tasmanian Museum
and Art Gallery, the Queen Victoria Museum and
Art Gallery, the South Australian Museum, and
the West Australian Museum for loans of
material for this study. 1 am grateful to the
Council of the Australian Biological Resources
Study for financial support of illustrators,
Catherine Harvey and Christine Lambkin; thanks
also to Catherine for her skill in computer
graphics. I thank colleague Robert Raven for
scanning micrographs and construction ol the
maps showing distribution.

LITERATURE CITED

DAVIES, V. TODD 2002. Tasmahrochus , a new spider
from Tasmania, Australia (Araneae.
Amphinectidae: Tasmarubriinae). Journal of
Arachnology 30: 219-226.

PLATNICK, N.I. & SHADAB, M.U. 1975. A revision
of the spider genus Gnaphosa (Araneae:
Gnaphosidae) in America. Bulletin of the
American Museum ofNatural History 155. 1-16.
SUHM, M., THALER, K, & ALBERTI, G 1996
Glands in the male palpal organ and the origin of
the mating plug in Amautvbius species (Araneae.
Amaurobiidae) Zoologischer Anzeiger.
234:191-199.

MIOCENE PALYNOFLORAS FROM SUBSURFACE SEDIMENTS IN THE
BUNDABERG DISTRICT

MARY E. DETTMANN AND H.TREVOR CLIFFORD

Dettmann, M.E. & Clifford, H.T. 2003 06 30: Miocene palynofloras from subsurface
sediments in the Bundaberg district. Memoirs of the Queensland Museum 49(1): 261-267.
Brisbane. ISSN 0079-8835.

Well-preserved palynofloras extracted from fluvial sediments of the Cainozoic Elliott
Formation in the Bundaberg Trough, central-eastern Queensland enable dating the
formation, the age of which hitherto has been poorly constrained. Representation oi the
pollen taxon, Canthiumidites be 1 1 us (Stover & Partridge) Mildenhall & Pocknall, in
association with spores referable to Polypodiidites usmensis (van der Hammen) Khan &
Martin, both of which are restricted to Early Miocene and younger sediments in the
Capricorn Basin, confirm an age no older than the Early Miocene, and enable correlation or
the Bundaberg sequence with Early Miocene fluvio-deltaic sediments penetrated by LibQ
Sandy Cape 1-3R on Fraser Island. Records oi Polypodiidites usmensis in the Fairymead
beds, which underlie the Elliott Formation, indicate that the Fairymead beds arc also no older
than Early Miocene. □ Elliott Formation, palynofloras, Miocene, Capricorn Basin.

Mary E. Dettmann, Department of Botany, University of Queensland, St Lucia 4072; H.
Trevor Clifford, Queensland Museum, PO Box 3300, South Brisbane 4101, Australia,
April 2002.

Subsurface sediments in the Bundaberg area,
southeastern Queensland (Fig. 1), contain an
important aquifer, which is tapped as a source of
groundwater, principally for sugar-cane
irrigation. Fossil fruit-stones including the
recently reported Spondylostrobus symthii
F.Muell. (Dettmann & Clifford, 2002), having
been recovered during the course of water-bore
drilling by the Department of Natural Resources
and Mines. The fruits came from a sequence of
unconsolidated coarse to fine sands, silts, and
interbedded clays capped by a thin sequence of
Quaternary sands. The sand/silt/clay sequence is
lithologically similar to the Elliott Formation, the
type section of which occurs in the region
(Robertson, 1979), This formation, based on its
stratigraphic relationships with weathering and
erosional surfaces of underlying and possibly
penecontemporanous sediments, has been
reported to comprise Eocene and Oligocene to
Miocene sediments (Grimes, 1 988). With the aim
of resolving the subsurface stratigraphy and age
of the important aquifer a continuously cored
section, DNR 1360 0253 (Fig. 2), was drilled
south of Bundaberg. Sediments sampled from
this section have been palynologically analysed
for age assessment and the results are detailed
herein.

GEOLOGICAL SETTING

The Cainozoic history of southeastern Queens-
land has been one of basic volcanism associated

with Paleocene break-up of the Australian plate
followed by intermittent sedimentation and
periods of deep weathering (Grimes, 1988). In
the Bundaberg region subsurface sections
penetrated basalts believed to be of Paleocene
age and overlying fluvial sediments referred to
the Fairymead beds that were tentatively dated as
Eocene (Ellis, 1968; Foster, 1978; Robertson,
1979; Grimes, 1988) (Table 1). Outcrops of
fluvial sediments believed to be younger and
mapped as Elliott Formation occur in an area
between the Kolan and Elliott Rivers within the
Bundaberg Trough (Robertson, 1979; Day et
al.,1983); the type area on the banks of the Elliott
River southeast of Bundaberg has a flat to gently

TABLE 1. Stratigraphy of Upper Mesozoic and
Cainozoic sediments in the Bundaberg Trough as
currently recognised (Robertson, 1 979; Gnmes, 1 988).

Age

Formation

Lithology

Pleistocene and
Holocene

Estuarine, lagoonal,
and fluvial sands

Early Pleistocene

Hummock Basalt

Olivine basalts

‘.'Oligocene-Miocene

‘Elliott Formation’
(younger)

Sandstone, claystone

? Eocene

Elliott Formation
(older)

Sandstone, claystone

? Eocene

Fairymead beds

Sandstone, claystone

?Paleocene

Pemberton Grange
Basalt

Olivine basalts

Albian

Burrum Coal
Measures

Coals, shales and
silts

262

MEMOIRS OF THE QUEENSLAND MUSEUM

sloping surface. To the west of the type area, the
surface buries deeply weathered Mesozoic and
older rocks, but it is uncertain whether or not the
surfaces are contemporaneous. In places a
younger, lithologically similar unit is thought to
be represented (Grimes, 1988) and this later
Elliott Formation’ may be of Oligocene to Miocene
age (Grimes, 1988, p.69). Overlying Pleistocene
and Holocene sands are restricted to alluvial
terraces and coastal plains (Grimes, 1988).

MATERIAL AND METHODS

The sequence encountered in DNR 1 360 0253,
situated adjacent to Foleys Road, 20km south of
Bundaberg, comprises coarse to fine sands, silts,
and clays beneath Quaternary sands (Fig. 2).
Several horizons were sampled and prepared tor
palynoloeical analysis using standard procedures
(Phipps & Playford, 1984). All samples yielded
abundant fungal remains and small-to-medium-
sized wood fragments were recovered from the
coarse sands. Two samples (24.6 m and 56.8 m)
provided well preserved assemblages of palyno-
morphs including diverse spores and pollen
grains (Table 2). Strew slides were examined in
transmitted light and photographed at
magnifications up to X 1250.

Illustrated specimens are housed in the
palaeontological collection of the Queensland
Museum. Registered numbers (QMF) refer to the
catalogue of that repository and are listed in Table
3 together with slide numbers and co-ordinates
derived from a standard ‘England Finder’ slide.

FIG. 2. Lithologic log of section cored by DNR 1360
0253 showing sampling horizons.

BIOSTRATIGRAPHY AND AGE

The diverse, well-preserved palynomorph
assemblages recovered from DNR 1 360 0253 are
dominated by angiosperm pollen with significant
contributions from ferns and lesser frequencies
of gymnosperm pollen (Table 2). A Tertiary age is
clearly indicated by the association of Halora-
gicidites harrisii , Nothofagidites emarcidus , N.
asperus , Malvacipollis spp., and Myrtaceidites
spp. Qualitative/quantitative attributes of the
assemblages are in accord with those reported
(Wood, 1986) from an Early Miocene section
penetrated by GSQ Sandy Cape 1-3R located on
Fraser Island and some 1 00km E of Bundaberg
(Fig. 1). Specifically, the Bundaberg palyno-
floras and those from between 421 .65-433. 5m in
Sandy Cape 1-3R share high frequencies of fern
spores, including significant representation
(>10%) of Cyathidites paleospora , together with
Polypodiidites spp. and Crassoretitriletes
vanraadshooveni. The associated angiosperm
pollen florules include common Haloragicidites
harrisii , Rhoipites , and Nothofagidites coupled
with infrequent Clavastephanocolporites
meleosus and Canthiumidites bellus (Wood,

MIOCENE PALYNOFLORAS FROM QUEENSLAND

263

1986) (Table 2). This Sandy Cape 1-3R section
includes fluvio-deltaic sediments containing
rotaliid and miliolid foraminifers at 425m;
overlying calcareous sandstones and calcarinites
contain foraminiferal associations of Early
Miocene age (Palmieri, 1984). The fluvio-deltaic
beds are believed to express deposition early in a
transgressive episode initiated during the Early
Miocene; correlative sediments occur in several
sections to the north in the Capricorn Basin,
offshore from Rockhampton (Palmieri, 1984).
Two of the Capricorn Basin sections, AGO
Aquarius 1 and AGO Capricorn 1, yielded
diverse palynofloras (Hekel, 1972) that share
many of the features of those from the Sandy
Cape and Bundaberg sections. They contain
moderate frequencies of Haloragicidites Harris ii,
Nothofagidites , Rhoipites , and fern spores.
Amongst the fern taxa are rare Crassoretitriletes
vanraadshooveni and common Polypodiidites ,
including P usmensis , which has first appear-
ances in Miocene sediments (Hekel, 1972).

The Sandy Cape 1 -3R fluvio-deltaic sediments
overly two basalt flows, the lower of which is
underlain by marine sediments containing
foraminiferids of late Oligocene age (Grimes,
1982; Palmieri, 1984). Between the two basalt
flows are deltaic sediments; palynofloras from
these sediments (537-539.4 m) differ from those
at 42 1 .65-433.5 m above the basalts in containing
high frequencies of Haloragicidites harrisii , low
frequencies of fern spores, and Malvacearum-
pollis mannanensis (Wood, 1 986).

In summation, evidence from marine sections
in the Capricorn Basin reveals that Poly-
podiidites usmensis and Canthiumidites bellus
are restricted to Early Miocene and younger
sediments. The association of these two taxa in
terrestrial sediments between 24.67 and 56.83 m
in DNR 1 360 0253 near Bundaberg argues for an
age no older than Early Miocene.

The only other Cainozoic sediments in the
region from which palynological data have been
published are from the Fairymead beds in DME
FairymeadNS 1 (Robertson, 1979) (Fig. 1, Table
1). A sample from 61.7m in this well provided
common Haloragicidites harrisii, infrequent
Nothofagidites and proteaccous grains in
association with Polypodiidites usmensis (Foster,
1979). Occurrence of the last mentioned taxon
implies an age no older than Early Miocene. Such
a dating raises the question of whether or not the
Elliott Formation and Fairymead beds were
deposited contemporaneously. Foster (1979) had

argued for a late Early Eocene age of the Fairy-
mead palynoflora based on its broad similarity to
palynofloras of the Proteacidites asperopolus
Zone of southeastern Australia as defined by
Stover & Partridge (1973). He also considered
that palynofloras from the upper fluvio-deltaic
sediments in GSQ Sandy Cape 1-3R were
dominated by taxa restricted to Eocene sediments
in southeastern Australia, but suggested these
may have been reworked (Foster, 1978).

From present evidence, first appearances of P
usmensis and C. bellus in the Capricorn Basin
may be earlier than in the more southerly basins
of Australia. C. bellus , the nominate species for
the C. bellus Zone of the Gippsland Basin and the
C. bellus equivalent Zone in the Murray Basin
has documented first appearances in those regions
as late Early Miocene and last appearances in the
M galeatus Zone of Pliocene age (Stover &
Partridge, 1973; Macphail, 1996). This contrasts
to the appearances of the pollen taxon in
foraminiferal Zone N4 (of Palmeiri, 1986) of
earliest Miocene age in the Sandy Cape section.
P. usmensis , a spore taxon with affinities with the
fern Stenochlaena is known from the Murray
Basin where it is restricted to sediments ot the C.
bellus Zone equivalent dated as late Early -
Middle Miocene age (Macphail, 1996); in the
Capricorn Basin the species ranges from earliest
Miocene (foraminiferal Zone N4) to Pliocene age
(Hekel, 1972). Wood (1986) advanced evidence
for the disparate stratigraphic ranges of several
palynomorph taxa in Tertiary sediments of
Queensland and those of more southerly regions
of Australia; further evidence has been
subsequently detailed by Beeston (1994) and
Dettmann & Clifford (2000).

ACKNOWLEDGEMENTS

Mr Peter Baker, formerly of the Department of
Natural Resources and Mines, Bundaberg is
thanked for providing sediment samples and logs
of DNR 1360 0253. We are especially grateful to
Dr Vince Palmieri for helpful discussions and
reviewing an early draft of the manuscript. We
thank Dr^Mike Macphail and Dr John McKellar
for their helpful reviews of the manuscript. MED
thanks the Director, Queensland Museum for
providing facilities and acknowledges support
from the Australian Research Council during
tenure of a Senior Research Fellowship.

264

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. A-G, spores, x 750 and H-P, pollen, x 1000 from DNR 1360 0253 borehole. A, Gleicheniidites sp.; B,
Dictyoph yllidites arcuatus Pocknall & Mildenhall; C, D, Polypodiidites usmensis (van der Hammen) Khan &
Martin, optical section and high focus respectively; E, Polypodiidites speciosus (Harris) Khan & Martin; F,
Microfoveolatosporites sp.; G, Peromonolites sp.; H, Dacrycarpites australiensis Cookson & Pike; I \Arecipites
sp.; J, K, Tricolpites sp. 1 polar views of two specimes; L, M, Tricolpites sp. 2, specimens in polar and lateral
aspect respectively; N, Tricolpites sp. 3; 0,P, Rhoipites sphaerica (Cookson) Pocknall & Crosbie, lateral aspect,
optical section and high focus respectively.

LITERATURE CITED

BEESTON, 1994, J.W. Tertiary palynology in the
Mount Coolon and Riverside areas. Queensland
Geology 6: 127-179.

DAY, R.W., WHITAKER, W.G, MURRAY, C.G,
WILSON, I.H. & GRIMES, K.G. 1983.
Queensland Geology. Geological Survey of
Queensland Publication 383.

MIOCENE PALYNOFLORAS FROM QUEENSLAND

265

FIG. 4. Pollen, x 1000. A, Proteacidites ivanhoensis Martin; B, C, Propylipollis pseudomoides (Stover)
Dettmann & Jarzen, high focus and optical section respectively; D, Propylipollis pseudomoides (Stover)
Dettmann & Jarzen; E, Lewalanipollis sp.; F, G, Canthiumidites belhvs (Stover & Partridge) Mildenhall &
Pocknall, optical section and high focus respectively; H, Polyoriflcites oblatus Martin; I, Haloragacidites
suttorensis Beeston; J, Cupanieidites reticularis Cookson & Pike; K, Malvacipollis sp.; L,
Clavastephanocolporites meleosus Martin, Macphail & Partridge; M, Tetracolporites sp.; N, Ericipites sp.; O,
Nothofagidites asperus (Cookson) Stover & Evans; P, Nothofagidites emarcidus (Cookson) Stover & Evans; Q,
Haloragacidites harrisii (Couper) Harris.

DETTMANN, M.E. & CLIFFORD, H.T. 2000.
Monocotyledonous fruits and seeds, and an
associated palynoflora from Eocene-Oligocenc
sediments of coastal central Queensland. Review
of Palaeobotany and Palynology 110; 141-173.

2002. Spondylostrobus F.Mueller; operculate fruits
of an extinct dicotyledon from the mid-Tertiary
of Australia. Review of Palaeobotany and
Palynology 122:219-237.

ELLIS, P.L.1968. Geology of the Maryborough 1:250
000 Sheet area. Geological Survey of Queensland,
Report 28.

FOSTER, C.B. 1978. Report on Tertiary spore-pollen
assemblages from GSQ Sandy Cape 1-3R.
Geological Survey of Queensland, Record
( 1 981/40 unpublished).

1979. Early Tertiary (Eocene) miospores from
Fairymead NS 1 . Appendix 2 in Robertson, A.D.
1 979. Revision of the Cainozoic geology between

266

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Palynomorph distribution in DNR 1630-0253, 20kin S of Bundaberg. Percentages derived from
counts of 200 palynomorphs per sample; + denotes present in sample, but not in count.

Palynomorph Taxa

Affinity

Depth

1 24.67- 7Sm 1 8. 81 m

Ferns:

|j Azolla Lam., microsporangia

Salviniaceac. Azolla

+

1 Crassiretitriletes vanraadshoovenii Germeraad et al.

Schizaeaceae, Lygodium
niicrophvllum-tvpe

+

Cyathidites paleospora (Martin) Aliev & Broadbridge

Cyatheaceae. Cvathea

12.5

16.5

ij Dictvoph yllidites arcuatus Pocknall & Mildcnhall

_! ?Gleicheniaccae

3.5

2

Foveosporites sp. of Wood (1986)

?OphiogIossaceae

Gleicheniidites sp.

Gleichenianceae

+

Ischvospo riles sp.

Dicksoniaceae

L a e\ igafospo riles ovatus Wilson & Webster

Filicales

10 5

1 M icrofoveolatosporites sp. (-Polypodiisporites sp. of Wood,

1986. fig. 3.6)

Filicales

+

Peromonolites spp.

Filicales

Polypodiidites speciosus (Harris) Khan & Martin

PolvDodiaceae

+

4

P. usmensis (van der Hammen) Khan & Martin

Blechnaccae. Stenochlaena

1

Gvmnosperms:

Araucariacites australis Cookson

.. Araucariaceae

Dacrycarpites austra liens is Cookson & Pike

Podocarpaccac. Dacrvearpus

2

2

Lvgistepollenites florinii (Cookson & Pike) Stover & Evans

Podocarpaceae. Darrydium

1

1

Podocarpidites elliptuus Cookson

3

Trichotomosulcites subgranulatus Couper

1 Podocarpaceae

1 s

Monocotyledons:

Arecipites sp.

?Arecaceae? Liliaceae

4.5

]

Cyperacaepollis sp.

Cyperaceae

+

_Canthiumidites bellus (Stover & Partridge) Mildcnhall & Pocknall
Clavastephanocolporites meleosus Martin, Macphail & Partridge

Rubiaccae. Randia

+

+

Alangiaceae, Alangium
villosum- type

+

Cupanieiditcs reticularis Cookson & Pike

^ Sapindaceae, Cuoanieae

+

Ericipites sp.

?Eoacridaceae

Haloragacidites cainozoicus Cookson & Pike

Casuarinaccae

H. harrisii (Couper) Harris

Casuarinaceae

36 5

H. suttorensis Beeston

Unidentified angiosperm

15 5

1 Z.J

7 ^

Polyorijiates oblatus Martin

+

Ilexipollenites anguloclavatus McIntyre

2

i Lewalanipollis sp.

Protcaceae

Malvacipollis subti/is Stover & Partridge

5

3

Malvacipollis spp.

Euphorbiaceae

2

Mvrtaceidites parvus-mesonesus (Cookson & Pike) Stover &

Evaas

Myrtaceae

1

1

| Nothofagidites asperus (Cookson) Stover & Evans

Nothofagaceae, Nothofagus
Lophozonia

5.5

N. emarcidus (Cookson) Stover & Evans

Nothofagaceae, Nothofagus
Brassospora

2

17.5

Periporopollenites sp.

+

! Propvlipollis pseudanwides (Stover) Dettmann & Jarzen

Protcaceae

6

Proteacidites ivanhoensis Martin

j

+

Proteacidites spp.

Protcaceae

2

Rhoipitex sphaerica (Cookson) Pocknall & Crosbie

2

4

5

Rhoipites microreticulatus (Harris) Macphail et al

Unidentified angiosperm

2

Rhoipites spp.

Unidentified angiosperm

Tetracolporites sp.

+

+

•f

+

+

+

Tricolpites sp. 1

Tricolpites sp. 2

Tricolpites sp. 3

Unidentified angiosperm
Unidentified angiosperm
- Unidentified angiosperm

MIOCENE PALYNOFLORAS FROM QUEENSLAND

267

TABLE 3. Register of illustrated specimens.

Taxon

Figure

Prep./Slide

England Finder
Co-ordinates

Catalogue no.

Arecipites sp.

31

M 1 04/C 1

S54

QMF51095

Canthiumidites bellus

4F,G

M104/C1

V40

QMF5I096

Clavastephanocolpori tes meleosus

4L

M101/F1

K36/4

QMF51094

Cupanieidiles reticularis

4J

M101/F2

T41

QMF51097

Dacrycarpites australiensis

3H

M101/F1

P53

QMF51098

Dictyophyllidites arcuatus

3B

M101/F1

D37

QMF51099

Ericipites sp.

4N

M101/C2

K19

QMF51100

Gleicheniidites sp.

3A

M104/F2

X30/2

QMF51101

Haloragicidites harrisii

4Q

M104/F2

U33/1

QMF51102

H. suttorensis

41

M104/F1

S41

QMF51103

Lewalanipollis sp.

4E

M104/C2

U23/1

QMF51105

Malvacipollis sp.

4K

M101/F2

P50/4

QMF51106

Microfoveolatosporites sp.

3F

M 101 /FI

F18

QMF51 107

Nothofagidites asperus

40

M101/F1

051

QMF51 108

N. emarcidus

4P

M101/F1

M54/3

QMF51109

Peromonolites sp.

3G

M101/F1

Ml 9/2

QMF511I0

Polyorificites oblatus

4H

M104/C1

023

QMF51104

Polypodiidites speciosus

3E

M101/F1

Q54

0MF51111

P. usmensis

3C,D

M104/F2

L49/3

OMF51112

Propylipollis pseudomoides

4B,C

M104/C1

N50/4

QMF51114

4D

M104/C2

G36/2

QMF51115

Proteacidites i vanhoensis

4A

M101/F1

F28/3

QMF51113

Rhoipites sphaerica

30

M101/C2

D37

QMF51116

3P

M 104/Cl

T36

OMF51117

Tetracolporites sp.

4M

M101/C1

041/3

QMF51118

Tricolpites sp. 1

3J

Ml 04/Cl

049/1

QMF51119

3K

M104/C2

P44/4

OMF51120

Tricolpites sp. 2

3L

M101/F2

L50/3

OMF51121

3M

M101/C2

L14

QMF51122

Tricolpites sp. 3

3N

M104/C2

J49/3-4

OMF51123

the Kolan and Elliott Rivers. Queensland
Government Mining Journal 80: 363.

GRIMES, K.G 1982. Stratigraphic drilling report
GSQ Sandy Cape 1-3R. Queensland Government
Mining Journal 83:224-233.

1988. Cainozoic geology, south-east Queensland.
Pp. 53-81. In Hamilton, L.H. (ed.) Field
Excursions Handbook for the Ninth Australian
Geological Convention. Geological Society of
Australia, Queensland Division, Brisbane.

HEKEL, H. 1972. Pollen and spore assemblages from
Queensland Tertiary sediments. Geological
Surv ey of Queensland, Publication 355: 1-29.

MACPHAIL, M.K. 1996. Paly nostratigraphy of the
Murray Basin, inland southeastern Australia.
Records of the Australian Geological Survey
Organisation 1996/57: 1-38.

PALMIERI, V. 1984. Neogene Foraminiferida from
GSQ Sandy Cape 1-3R bore, Queensland: a

biostratigraphic appraisal. Palaeogeography,
Palaeoclimatology, Palaeoecology 46: 165-183.
PHIPPS, D. & PLAYFORD, G. 1984. Laboratory
techniques for extraction of palynomorphs from
sediments. Papers of the Department of Geology,
University of Queensland 1 1 : 1-23.
ROBERTSON, A.D. 1979. Revision of the Cainozoic
geology between the Kolan and Elliott Rivers.
Queensland Government Mining Journal 80:
350-363.

STOVER, L.E. & PARTRIDGE, A.D. 1973. Tertiary
and Late Cretaceous spores and pollen from the
Gippsland Basin, southeastern Australia.
Proceedings of the Royal Society of Victoria 85:
237-286.

WOOD, GR. 1986. Late Oligocene to Early Miocene
palynomorphs from GSQ Sandy Cape 1-3R.
Geological Survey of Queensland, Publication
387: 1-27.

A NEW GENUS OF HENICOPID CENTIPEDE (CHILOPODA: LITHOBIOMORPHA)

FROM NEW CALEDONIA

GREGORY D. EDGECOMBE

Edgecombe, G.D. 2003 06 30: A new genus of henicopid centipede (Chilopoda:
Lithobiomorpha) from New Caledonia. Memoirs of the Queensland Museum 49(1):
269-284. Brisbane. ISSN 0079-8835.

Two species from New Caledonia, Easonobius tridentatus gen. et sp. nov. and Paralamyctes
humilis Ribaut, 1923, together represent a new genus of Henicopini. South African species
formerly classified together with Easonobius humilus in Analamyctes Chamberlin, 1 955, are
distantly allied, members of Paralamyctes (Paralamyctes) Pocock, 1901. Morphological
characters of Easonobius , including electron microscopic study of the head, indicate
membership in the Lamyctes-Henicops Group. Parsimony analysis tavours a closest
relationship between Easonobius and the Australasian genus Henicops Newport, 1 844. The
type species of Lamyctes ( Eumyctes ) Chamberlin, 1951, shares apomorphic characters with
the Cape genus Lamyctopristus Attems, 1 928. to which Eumyctes is transferred. □ C hilopoda ,
Lithobiomorpha , Henicopidae , Easonobius, New Caledonia , taxonomy, phytogeny.

Gregory D. Edgecombe, Australian Museum, 6 College Street, Sydney 2010, Australia
(e-mail: greged@austmus.gov.au); 22 March 2003.

Paralamyctes humilis Ribaut, 1923, was
named based on a single, small male from
Oubatche in northeastern New Caledonia.
Subsequently, Chamberlin (1955) reassigned P.
humilis to the genus Analamyctes Chamberlin,
1955, and, more precisely, to an invalidly erected
subgenus Capolamyctes . Analamyctes
(Analamyctes) was conceived as occurring in
Argentina (the type species A. tucumanus
Chamberlin, 1955, from Tucuman Province, and
Paralamyctes andinus Silvestri, 1903, from
Mendoza Province). Analamyctes (Capola-
myctes) received species from the Cape region of
South Africa (Paralamyctes asperulus Silvestri,
1903; P. levigatus Attems, 1928; P tabulinus
Attems, 1 928) together with the New Caledonian
P. humilis.

In a revision of Paralamyctes Pocock, 1901,
Edgecombe (2001) dismissed Chamberlin’s
(1955) reassignment of South African species to
Analamyctes , these taxa being accommodated
within a monophyletic Paralamyctes
(Paralamyctes) Pocock, 1901. This conclusion is
supported by both morphological (Edgecombe,
2003a) and molecular data (Edgecombe &
Giribet, 2003a). A reconsideration of Anala-
myctes was made possible by a restudy of its type
species, A. tucumanus. Argentinian species of
Analamyctes are members of a Lamyctes-
Henicops Group, only distantly allied to P.
(Paralamyctes) (Edgecombe, 2003b).

The present study reconsiders Paralamyctes
humilis , which until now has defied phylogenetic

placement. New collections made by G. B.
Monteith (Queensland Museum) in New
Caledonia include additional specimens of both
sexes of this species, as well as specimens that
represent another, closely related species.
Morphology of these species is documented by
electron microscopy, including mouthparts that
have proven useful in henicopid systematics, and
they are are coded for their morphological
characters in a dataset for henicopid phylogeny.

Institutional abbreviations cited in this work
are: AM - Australian Museum, Sydney; MNHN
Museum National d’Histoire Naturclle, Paris;
QM - Queensland Museum, Brisbane; ZMB -
Museum fur Naturkunde, Berlin. Morphological
terminology is as explained by Edgecombe
(2001: 203). Drawings were prepared with a
camera lucida attachment to a Leica MZ1-.
Scanning electron microscopy used a Leo 4o5V I
with a Robinson backscatter detector, and digital
images assembled into plates with Photoshop.

SYSTEMATICS

Easonobius gen. nov.

partim Analamyctes (Capolamyctes) Chamberlin, 1955
nomen nudum.

TYPE SPECIES. Easonobius tridentatus gen. et sp. nov.

ETYMOLOGY In honour of Dr Edward H. Eason
(1915-1999), for his contributions to lithobiomorph
systematics, with the standard suffix, -obius.

DIAGNOSIS. Member of Lamyctes-Henicops
Group lacking pseudoporodont (shared with

270

MEMOIRS OF THE QUEENSLAND MUSEUM

Henicops and Analamyctes ); posterior angles of
tergites 7,9, 11 and 1 3 produced (shared with
Henicops and Lamyctopristus); dental margin of
maxillipede coxostemite with 2+2 or 3+3 teeth;
tarsi of all legs bipartite (shared with
Analamyctes ); coxal process of first maxilla with
laciniate or plumose setae near dorsal margin
(shared with Henicops ); mandible with single
row of exclusively bipinnulate aciculae; last
distal spinose projection on tibia of leg 13; first
genital stemite of 6 undivided (plesiomorphies
excluding membership in Henicops). Antenna
with 26-33 articles; tergite of intermediate
segment with strongly concave posterior margin.

ASSIGNED SPECIES. Paralamyctes humilis
Ribaut, 1923.

DISCUSSION. Chamberlin distinguished two
subgenera of Analamyctes in a key, using a single
character. Analamyctes (Analamyctes), grouping
the Argentinian species A. tucumanus and A.
andinus , was defined based on 'posterior angles
of none of the dorsal plates produced’.
Analamyctes ( Capo lamyctes) grouped^, humilis
with three nominal South African species of
Paralamyctes (Edgecombe, 2001, 2003a).
Analamyctes (Capolamyctes) was distinguished
based on ‘posterior angles of tergites 9, 1 1 and 1 3
or 7, 9, 1 1 and 1 3 produced’. Though Chamberlin
was explicit about the membership and diagnosis
of Capolamyctes , he neglected to designate a type
species. The name fails Article 13.3 of the ICZN
Code, and is dismissed as a nomen nudum. As
argued in detail below, the original concept of
Capolamyctes delimits a polyphyletic group. Of
Chamberlin’s assigned species, only Parala-
myctes humilis is at all closely related to the type
species of Analamyctes.

Easonobius resembles Analamyctes in having
bipartite tarsi on all legs, a state restricted to these
taxa within the Lamyctes-Hen icops Group (but
possibly plesiomorphic by comparison to
Paralamyctes and Zygethobiini). Easonobius
most obviously differs from Analamyctes in its
tergal shapes, as used in Chamberlin’s (1955)
subgeneric scheme, with projections on TT7, 9,
11 and 13 versus nearly transverse margins in
Analamyctes. This does not in itself obviate a
close relationship, since tergite projections
occasionally vary within some well defined
henicopine clades, such as Paralamyctes
( Haasiella ), e.g., present in P. (H.) subicolus and
P. (//.) trailli; absent in P. ( H .) cammooensis and
P. (H.) ginini.

Easonobius is resolved as most closely related
to Henicops Newport, 1 844, in the best supported
phylogenetic analyses, described below.
However, several apomorphic characters unite
Australian and New Zealand members of
Henicops in the traditional sense (e.g., Attems,
1914, 1928; Chamberlin, 1920; Archey, 1937) as
a clade that excludes Easonobius. These
characters (numbered as in Table 1) include a
subdivision of the basitarsus indicated by paired
larger setae (40: 1 ), the first genital stemite of the
6 being divided longitudinally into two sclerites
(43:1 ), the distitarsus of leg 1 5 being divided into
tarsomeres (53: 1 ), and more setose gonopods in
both sexes. Henicops as traditionally delimited is
strongly supported (jackknife frequency 99%;
Fig. 8), and it is not expanded in scope and
rediagnosed to incorporate the species here
recognised as Easonobius.

Comparable in several respects is a group of
mostly Southern African species referred to
Lamyctes ( Eumyctes ) Chamberlin, 1951 [type
Henicops sinuatus Porat, 1893] and L.

( Neomyctes ) Chamberlin, 1951 [type Lamyctes
( Neomyctes ) ergus Chamberlin, 1951]. These are
distinguished from typical Lamyctes Meinert,
1868, by their projections on tergites 9, 1 1 and 13.
The style of tergite projections in species such as
L. (Eumyctes) sinuatus is as in Easonobius , and
these species also share a strongly concave
posterior margin to the tergite of the intermediate
segment (Attems, 1909, fig. 53). Chamberlin’s
(1951 versus 1955) distinction between
Eumyctes! Neomyctes and Analamyctes
(including E. humilus) placed fundamental
weight on the absence or presence of a tarsal
articulation. A closer relationship can be
proposed for the type of Eumyctes , L. (E.)
sinuatus , and the Cape genus Lamyctopristus
Attems, 1928, than the former shares with
Easonobius. Lamyctes (Eumyctes) sinuatus has
dense, strongly developed tuberculation on the
tergites, to a degree observed only in Lamycto-
pristus amongst all known Henicopidae. In both
L. (E.) sinuatus and L. granulosus (=L. validusl ),
tuberculation is more pronounced on the male
than on the female (Lawrence, 1955: 23), e.g.,
being well developed on the head shield of the
male. These species are also similar, and
resemble Henicops Newport, 1844, in having
tergite projections and distal spinose projections
on the tibia of leg 14. They share other peculiar
characters in addition to their tergal tubercul-
ation. In Lamyctopristus validus as well as
Lamyctes (Eumyctes) sinuatus , the tarsi of

A NEW CENTIPEDE GENUS

271

FIG. 1. A-F, Lamyctopristus ( Eumyctes ) sinuatus (Porat, 1893). ZMB 4962, 9 , Kamaggas, Northern Cape
Province, South Africa. A-E, mandible. A, medial view ofgnathal edge, scale 1 00pm; B, ventral part ot gnatna
edge, scale 50pm; C, dorsalmost tooth and furry pad, scale 20pm; D-E, aciculae, scales 20pm. 1 0pm. r , anterior
view of pretarsus, scale 30pm. G, Henicops maculatus Newport, 1844. Anterior view of pretarsus ot leg 14,
scale 15pm. H, Parcilamyctes ( Paralamyctes ) asperulus Silvestri, 1903. SAM-ENW-C53 14. Anterior view of
pretarsus of leg 14, scale 15pm.

272

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 2. A, Lamyctopristus (Eumyctes) sinuatus (Porat,
1893). ZMB 4962, 9, Kamaggas, Northern Cape
Province, South Africa, terminal segments and
gonopods. scale 200pm. B-F, Easonobius tridentatus
gen et sp. nov. QM S6029 1 , 9, Pic d'Amoa. Scale m
B applies to C-E. B. leg 1 2, scale 200pm; C, leg 1 3; D,
leg 14; E, leg 15; F, distal part of tarsus and pretarsus
of leg 15, scale 100pm.

anterior legs have a distinct curvature within the
distitarsal portion, this occurring despite the lack
of articulations. Mandibular characters (Fig. 1 )
are also consistent with a close relationship
between L. (Eumyctes) sinuatus , Lamyctopristus
and Henicops. The former species has a large
number of aciculae on the mandible, with the
density and arrangement of the aciculae two-deep
(Fig. IB, D), rather than a single row, being
otherwise observed only in Lamyctopristus
validus (Edgecombe, 2003b, fig. 34D) and in all
species of Henicops (Edgecombe et al., 2002, fig.
5C). Also as in Henicops and Easonobius , but not
Lamyctes, the accessory denticles on the dorsal-
most tooth of the mandible are simple, angular
elements (Fig. 1C), rather than multifurcating
scales (Edgecombe et al., 2002, tig. 7B, for
Lamyctes emarginatus). As well, the novel
expansion of basal article of the female gonopod
in Lamyctopristus may have a precursor in the
relative breadth of this article in L. ( E .) sinuatus
(Fig. 2A). Phylogenetic analysis including these
characters (see below) recognises L. (Eumyctes)
sinuatus as more closely related to Lamyctopristus
than to Lamyctes. To incorporate this relationship
into the classification, Eumyctes is reassigned to
Lamyctopristus.

FIG 3. Easonobius tridentatus gen. et sp. nov. A,
MNHN P244, holotype 9, Pic d’Amoa, dorsal
habitus, scale 1 mm; B, QM S6029 1 , 9 , Pic d’ Amoa,
terminal segments and gonopods, scale 100pm. C,
QM S60292, d, Aoupinie, terminal segments and
gonopods, scale 100pm.

Easonobius tridentatus sp. nov.

(Figs 3-6)

DIAGNOSIS. Easonobius with width of head
shield up to 1.5mm; penultimate and preceding
few antennal articles as wide as or wider than
long; dental margin of maxillipedc coxostemite
moderately wide, gently convex, with 3+3 teeth;
Tomosvary organ small; a few lacinate setae on
coxal process of first maxilla; posteromedian
embay ment in margin of T7 transverse or faintly
convex; short, spinule-like setae on tergites and
along tergal margins.

ETYMOLOGY. For the three teeth on the dental margin of
the maxillipede coxostemite.

MATERIAL. HOLOTYPE. MNHN P244, 9 (Fig. 3A),
Pic d’Amoa, N slopes, Province Nord, New Caledonia,
20°58’S 165°17’E, 500m, GB. Monteith, 24 November
2001-31 January 2002. PARATYPES. QM S60291, 9

A NEW CENTIPEDE GENUS

273

FIG. 4. Easonobius tridentatus gen. et sp. nov. SEMs, scales 100pm except G, JO pm. QM S6(L91, 2, Pic
d’Amoa. A, dorsal view of anterior part of head, proximal part of antennae; B,D,E, dorsal side ot antenna, U,
ventral view of clypeus and sclerotised bridge between antennae; F-G, cephalic pleurite and detail ol omosvary
organ.

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MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Easonobius tridentatus gen. et sp. nov. SEMs, scales 100pm except F, 50pm, G-H, 20pm, 1, 2pm. A-F, I,
QM S60291, 9, Pic d’Amoa. A, ventral view of maxillipede; B, detail of dental margin of coxostemite; C,
maxillipede telopodite; D, dorsal view of coxostemite; E, second maxilla; F, tarsus and claw of second maxilla;
I, distal part of lacinate seta on coxal process of first maxilla. G-H, AM KS 81365, 9, Pic d’Amoa, dorsal and
posterior views of claw, accessory claws and sensory spur of leg 14.

A NEW CENTIPEDE GENUS

275

(Figs 2B-F, 3B, 4, 5A-F, I, 6), AM KS 81365, 9 (Fig.
5G-H), from type locality, GB. Monteith. 3 1 Januaiy 2002.
OTHER MATERIAL. New Caledonia, leg. G.B.
Monteith. PROVINCE NORD: QM S60292, 6 (Fig. 3C),
21° II S 165°18’E, Aoupinie, top camp, 850m, 2-4
November 2001. PROVINCE SUD: QM S60293. d,
21°45’S 166°00’E, Mt Do summit 1000m, 21 November
2000.

DESCRIPTION. Length (head shield to end of
telson, slightly extended specimen) up to 16mm;
width ot head shield up to 1 .5mm. Colour (based
on specimens in absolute ethanol): head shield
orange with purple mottling, including region
surrounding ocellus; antenna pale orange,
sometimes with pale lavender tint along most of
length; tergites lavender with deep purple
longitudinal median band and mottling beside/on
borders; stemites pale lavender except for orange
stemites 14 and 15; legs pale yellow except for
pale yellowish-orange tarsi.

Head shield. Wider than T1-T5, equally wide as
T7, with shallow median notch, lacking
longitudinal median furrow (Fig. 4A); maximal
posterior extent of transverse suture at about 26%
length ol head shield; border as wide medially as
posterolaterally. Ocellus large, moderately
domed (Fig. 4A). Tdmosvary organ relatively
small (Fig. 4G), with outer margin near edge of
cephalic pleurite, beneath ocellus (Fig. 4F).

Clypeus with cluster of about six apical setae
(anterior pair and transverse band of three or
four), several smaller setae scattered postero-
lateral to these ( Fig. 4C); usual band of four setae
just in front ot labrum. Labral margin gently
concave where cluster of bristles projects beyond
margin; numerous branches along length of each
bristle.

Antenna. 34-38% length of body, 3.7 times length
ot head shield in largest specimen (Fig. 3A);
3 1 -27 and 31-31 articles in $ 9 , 30-30 and 3 1 -30
in S 6 ; basal two articles much enlarged relative
to others (Fig. 3A); articles 3-4, 7-8, 10-11 and
one or two more distal pairs short. Terminal
article up to 2.2 times length of penultimate; most
of distal articles of similar shape, slightly wider
than long (Fig. 4D). Setal density similar from
third article, with mix of longer trichoid sensilla
and shorter curved sensilla (Fig. 4B, E); most
setae oriented normal to antennal surface or sloping
anteriorly, arranged in imprecisely defined whorls,
as many as seven whorls on longer articles.
Maxillipede. Dental margin gently convex, with
3+3 teeth; outer tooth more distant than inner pair
to each other (Fig. 5B), well inside anterolateral
corner of dental margin. Median notch

moderately deep, parabolic or semicircular. Setae
rather evenly scattered over anterior two-thirds of
coxostemite (Fig. 5A); irregular band of short
setae on anterior part of dorsal surface of coxo-
stemite (Fig. 5D). Pretarsal part of tarsungulum
about equal in length to tarsal part; long setae
distinctly denser on inner side of tarsungulum
than on outer (Fig. 5C); setation on tibia and
femur fairly even on inner, outer and ventral
sides.

Mandible. Four paired teeth (Fig. 6A). Eleven
aciculae, each with large, blunt pinnules approx-
imately symmetrical on anterior and posterior
margins (Fig. 6C-D). Fringe of branching bristles
skirts aciculae; ventral bristles with moderately
wide bases, with even, rather dense branchings
along entire length of each bristle (Fig. 6B); fairly
abrupt transition to three overlapping rows of
multifurcating scale-like bristles against second
tooth; scales branch near their bases to form
continuous fringe of slender, hair-like spines;
fringe narrowing dorsal ly. Grooved ridges bearing
row of blunt accessory denticles well developed
on teeth (Fig. 6A-B); most accessory denticles
small, triangular, even on dorsal tooth (Fig. 6E).
Proximal part of dentate lamina consists of a
narrow band of fused scales, strongly different-
iated from furry pad (Fig. 6E); furry pad
composed of simple and multifurcating bristles.
First maxilla . Coxal parts of coxosternum
meeting along most of their length medially (Fig.
6F), separated posteriorly by small, wedge-
shaped stemite. Coxal process with cluster of up
to 1 5 simple setae at tip, a few setae along inner
margin; four or five laciniate setae above simple
setae near dorsal edge of coxal process (Fig. 6H),
thicker than simple setae, branching into up to
five short spines at their distal tips (Fig. 51). Distal
article of telopodite with two rows of up to 14
plumose setae along inner margin (Fig. 6G);
plumose setal rows fringed along ventral side by
row of shorter simple setae, along dorsal side on
anterior half of article by row of slender spines,
these more densely spaced than simple or
plumose setae (Fig. 6G); ventral surface of distal
article with numerous, evenly scattered simple
setae.

Second maxilla. Sternite fused to coxa, margins
distinct. Irregular band of about eight setae across
anterior pail of coxa (Fig. 5E). Joint between
trochanter and prefemur defined as a notch along
inner margin of telopodite (Fig. 5E). Inner face of
tarsus with up to about 20 plumose setae, densely
branching along their distal halves (Fig. 5F).
Pretarsal claw small, composed of up to five

276

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 6. Easonobius tridentatus gen. et sp. nov. SEMs. QM S60291, 9, Pic d’Amoa. A-E, right mandible. A,
gnathal lobe, scale 50pm; B, fringe of branching bristles and teeth, scale 10pm; C-D, aciculae, scales 10pm; E,
furry pad, scale 10pm; F, first maxillae, scale 100pm; G, distal article of telopodite of first maxilla, scale 50pm;
H, laciniate setae on coxal process of first maxilla, scale 10pm.

A NEW CENTIPEDE GENUS

277

digits of varied length and thickness, median
digit the largest (Fig. 5F).

Tergites (Fig. 3A). Weakly wrinkled, gently
turned up against borders. T1 trapeziform,
anterior width slightly less than T3, 8 1 % width of
widest tergite (T1 0), posterior margin transverse
or faintly concave; posterior angles of TT1-5
rounded; lateral border subparallel in T3,
posterior margin faintly or weakly concave;
posterior margin of T5 distinctly concave, that of
T8 slightly more so; TT9, 1 1 and 13 with strong,
blunt projections and wide, transverse or convex
median sector; T7 with shorter projections,
median sector subtransverse or weakly convex;
TT2, 4 and 6 bordered laterally, thickened
posteromedially; TT10, 12 and especially 14
with concave posterior margins, blunt posterior
angles. Tergite of intermediate segment with
concave posterior margin in both sexes. Tergite
of first genital segment less sclerotised than
telson tergite. Short, spinule-like setae scattered
across anterior third and laterally on long tergites;
numerous short, spinule-like setae along lateral
margins of tergites.

Legs. Distal spinose projection on tibiae of legs
1-13 (Fig. 2B, C), absent on 14 (Fig. 2D) and 15
(Fig. 2E). Legs 12-15 with length ratios 1: 1.2 :
1.6 : 2.5. Tarsal joints marked by desclerotised
band and weak flexure on anterior legs,
articulation weakly continuous on dorsal side of
leg; distitarsus about 55% length of basitarsus on
leg 12 (Fig. 2B), 63% length of basitarsus on leg
1 5; leg 1 5 basitarsus nine times longer than wide,
distitarsus about 10 times longer than wide, tibia
5.5 times longer than wide (Fig. 2E). Prefemur
with numerous short setae on dorsal and ventral
sides, with one or a few longer setae near
midlength on ventral side of legs 1-13; more
evenly short setae on prefemur of legs 14-15;
setae relatively fewer on femur and tibia, of
similar size to most on prefemur, densest on
tarsus; short setae on distitarsus of legs 1-14
sloping distally; setae relatively sparse on tibia
and tarsus of leg 15. Anterior and posterior
pretarsal accessory claws nearly symmetrical on
all legs, about half length of main claw, weakly
diverging (Fig. 5G); minute sensory spine on
posterior side of claw base (Fig. 5H), lacking on
anterior side.

Coxal pores. All round, separated by less than
their diameter when abundant, inner pores
smaller; 4, 5, 5, 5/4, 5, 5, 5 in largest $,
3, 4,5, 5/3, 4,5, 5 (Fig. 3B) and 2,3, 4, 4/2, 3, 4, 4 in
progressively smaller $$; 3, 3, 4, 3/3, 3, 3, 3 in
largest cJ, 1,2, 2, 2/ 1,2, 2, 2 in smaller 6 (Fig. 3C);

pore row not set in a groove, separated from
anteroventral face of coxa by rounded surface.

Female (Fig. 3B). Stemite of segment 1 5 weakly
to gently convex posteromedially, fringed with
short setae along posterior margin. Setae fairly
evenly scattered on posterior two-thirds of
stemite of first genital segment, few or lacking on
anterior third; transverse band of setae slightly in
advance of posterior margin. Gonopod with two
relatively small, conical spurs, inner spur slightly
smaller; up to 20 setae on basal article of
gonopod, up to 8 on second article, two or three
on distal article; claw simple.

Male (Fig. 3C). Stemite of segment 15 weakly
convex posteromedially, most setae along mar-
gins. Stemite of first genital segment undivided, a
few setae in front of posterior margin. Articles of
gonopod with four, two, and one small setae
(proximally to distally).

DISCUSSION. Specimens assigned to Easonobius
tridentatus sp. nov. resemble E. humilis (Ribaut)
in having a similar number of antennal articles
(27-31 in the new species versus 26-33 in E .
humilis ), projections with the same shape on
tergites 9, 1 1 and 13, a bipartite tarsus on legs
1-12, and absence of a pseudoporodont. The
holotype of Easonobius humilis is a small male
(width of head 0.75mm; length of body 6.6mm)
lacking most legs, and no other material was
assigned to the species by Ribaut (1923). The
original illustrations are accurate except for the
position of the Tomosvary organ, which was
depicted as anterior to the ocellus (Ribaut. 1 923,
fig. 24) when in fact the organ lies beneath the
ocellus in the holotype and in new specimens
assigned to the species, as is also the case in E.
tridentatus. Ribaut noted 33 antennal articles in
the description based on the right antenna: the
unfigured left antenna has 30 articles.

All specimens of Easonobius tridentatus are
larger than specimens of E. humilis , and all have
3+3 teeth on the dental margin of the maxillipede
(Fig. 5A-D) versus 2+2 teeth on a narrower
margin in E. humilis (Fig. 7A-B). Although some
henicopids have an ontogenetic increase in
number of maxillipede teeth [Paralamyctes
validus : Archey, 1921: 182; Anopsobius
neozelanicus : Archey, 1937: 87; Paralamyctes
(Haasiella) cammooensis : Edgecombe, 2003b],
tooth numbers in the Lamyctes-Henicops Group
are fixed early in ontogeny when the adult
number is 2+2 or 3+3 teeth. For example, the
dental formula 3+3 is complete by 8-legged
larval stadium LII in Henicops from Victoria,

278

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 7. Easonobius humilis (Ribaut, 1923). AM KS 81366, 9 , Mt Koghis. A, ventral view of maxillipede, scale
100pm; B, detail of dental margin of maxillipede coxostemite, scale 50pm; C, ventral part of mandibular
gnathal edge, showing aciculae, scale 10pm; D, dorsal view of coxal processes of first maxillae and inner
margins of telopodites, scale 20pm; E, cephalic pleurite, showing Tomosvary organ, scale 50pm; F, distal
articles of antenna, scale 50pm; G, gonopods, scale 50pm.

A NEW CENTIPEDE GENUS

279

Australia (Museum Victoria NOH-1778, 1782,
1 786), and the coxostemite shape more closely
resembles the adult shape than is the case
between the much more similar sized specimens
of E. humilis and E. tridentatus. Lamyctes
fulvicornis (=L. emarginatus) likewise acquires
its adult dental formula (2+ pseudoporodont) in
LI I, and the LII dental margin is similar to that of
the mature stages (Andersson, 1984, fig. 6). The
substantial differences between the coxostemal
shape o fE. humilis and the smallest specimens of
E. tridentatus are thus unlikely to be attributable
to ontogenetic change in a single species.

A few other differences between small ( E .
humilis) and larger (E. tridentatus) specimens are
atypical for ontogenetic variation. The holotype
of E. humilis and two additional specimens
assigned to that species have a prominently
rounded (convex) median sector to the posterior
margin of T7, whereas this sector of the margin is
at most weakly convex in E. tridentatus.
Elaboration of tergal margins is typically
enhanced, rather than suppressed, in
lithobiomorph ontogeny (e.g., projections
become more prominent: Andersson, 1981) so
the modified margin of the small specimens
appears to have taxonomic significance. The
holotype of E. humilis and 9 QM S60637 have a
relatively longer antenna than do any specimens
of E. tridentatus , the elongation deriving from a
larger number of articles (maximum 33 in these
specimens) and an elongation of the distal
articles. The penultimate and adjacent articles in
E. humilis are longer than wide (Fig. 7F), the
reverse of the condition in E. tridentatus (Fig.
4D). The 6 gonopodofthe holotype of E. humilis
(Ribaut, 1923, fig. 26) is more setose than is that
of larger specimens of E. tridentatus (Fig. 3C).

Easonobius tridentatus (Fig. 4F) has a
substantially smaller Tomosvary organ than does
E. humilis (Fig. 7E). This difference may be
size-related because several small Henicopidae
have large Tomosvary organs; this is observed
repeatedly in blind lineages [Anopsobiinae;
Lamyctes coeculus\ Paralamyctes (Haasiella)
trailli ] but also in some small species that retain
ocelli [P. (. Haasiella ) cammooensis and P. (//.)
ginini].

Modified setae on the coxal process of the first
maxilla also serve to distinguish the species.
Easonobius tridentatus has several lacinate setae
near the dorsal margin of the coxal process,
above the main cluster of simple setae (Figs. 51),
with branching confined to a few spines at the

distal tip of the setae (Fig. 6H). In E. humilis a
single plumose seta (Fig. 7D) is instead present’in
addition to the simple setae.

Easonobius humilis (Ribaut 1923)

(Fig. 7)

Paralamyctes humilis: Ribaut, 1923: 23, figs 24-26.
Paralamyctes humilis : Wurmli, 1974: 526, fig. 2.
Analamyctes humilis : Chamberlin, 1955: 50.

Analamyctes humilis: Edgecombe, 2001: 206.

DIAGNOSIS. Wide head shield 0.75-0. 8mm;
penultimate and preceding few antennal articles
longer than wide; dental margin of maxillipede
coxostemite narrow, with 2+2 teeth, margin
strongly sloping posterolaterally distal to outer
tooth; Tomosvary organ large; single plumose
seta near dorsal edge of coxal process of first
maxilla; posteromedian embayment in margin of
T7 convex; tergites lacking spinule-like setae.

MATERIAL. HOLOTYPE. Naturhistorisches Museum
Basel, Zoologische Abteilung 303a, d, Oubatche,
Province Nord, New Caledonia. OTHER MATERIAL.
New Caledonia, leg. GB. Monteith. PROVINCE SUD-
AM KS 81366, 9 (Fig. 7A-G), Mt Koghis, 22° ITS
166°0rE, 750m, 29 November 2000; AM KS 82627, d,
Mt Koghis, 500m, 2-3 November 2002; QM S60637, 9,
S60651, 9, Mt Humboldt, source, 21°53’S 166°24’E,
1300 m, 5-8 November 2002, rainforest.

DISCUSSION. Specimens from Mt Koghis (Fig.
7) and Mt Humboldt are assigned to E. humilis
despite the substantial geographic distance from
the type locality. The specific diagnosis indicates
characters that these specimens share with each
other to the exclusion of the larger E. tridentatus.
The five specimens have a maximum number of
26, 28 (N=2) and 33 (N=2) antennal articles. The
strength of tarsal articulations on anterior legs
varies between specimens from the same locality
(faint on Mt Humboldt specimen QM S60637 as
in the holotype; well defined on Mt Humboldt
specimen QM S60652). Coxal pore counts are
1,1, 2,2/1, 1,2, 2 (holotype) and 1 ,2,2, 2/0,2, 2, 2
(AM KS 82627) in males and l, 2,2, 2/1, 2, 2, 2 in
females. The female gonopods (Fig. 7G) have a
pair of bullet-shaped spurs with their bases
adjacent to each other (see also Wurmli, 1974).

PHYLOGENETIC RELATIONSHIPS

Morphological characters of Easonobius
humilis and E. tridentatus are scored for the
character set of Edgecombe (2003b). Several
new characters (characters 52-57 in Table 1) bear
on the relationships of Henicops and
Lamyctopristus. Other than adding the two New
Caledonian species, taxonomic sampling is as in

280

MEMOIRS OF THE QUEENSLAND MUSEUM

100

69» —

89

99

83 L

58

52

50 1——

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65|

£

tridentatus
den tat us
maculatus
n. sp. QLD
— spcnceri
asperulus
weberi
harrisi
monteithi
tridens

nevcrneverensis

chilensis

cassisi

mesibovi

gray I

?grayi

hornorae

validus

ginini

cammooensis

tram

subicolus

Lithoblus obscurus
Lithobius variegatus
Australobius scabrior
Bothropolys multidentatus —
Shikokuoblus japonic us -|
Anopsoblus neozelanicus
Anopsobius n. sp. NSW
Anopsobius n. sp. TAS
Dichelobius flavens
Zygethobius ponds
Cermatobius Japonicus
Analamyctes tucumanus
Analamyctes andlnus
Lamyctcs africanus
Lamyctes coeculus
Lamyctes omarginatus
Eumyctes slnuatus
Lamyctopristus validus
humills

Lithobiidae

Anopsobiinae

^ Zygethobiini

]

Easonobius

Henicops

P. (Paralamyctes)

| P (Nothofagobius)
| P. (Thingathinga)

P (Haasiella)

*5

§

o'

*1

(/)

O

o

c

TJ

3

oT

o

FIG. 8. Consensus of cladcs present in more than 50% of jackknife
replicates based on morphological data in Table 2, showing jackknife
frequencies. Groups with jackknife frequencies in italics are
contradicted in some of the 4,233 minimal length cladograms.

Edgecombe (2003b) except for
the inclusion of Lamyctopristus
( Eumyctes ) sinuatus (Porat,

1893), as discussed above, and
Paralamyctes ( Paralamyctes )
asperulus Silvestri, 1903 ( =P
tabulinus Attems, 1928; see
Edgecombe, 2003a). This species
is included to test Chamberlin’s
(1955) concept of a New
Caledonian/South African clade
(his k Capolamyctes ' concept).

Data were analysed with
PA UP* version 4.0b 1 0 ( SwofTord,

2002). A heuristic search used
5,000 random stepwise addition
sequences that sampled five trees
per iteration, followed by TBR
(tree bisection-reconnection)
branch swapping on these trees.

Cladograms were rooted with
Lithobiidae as outgroup to
Henicopidae. Multistate characters
were coded as unordered. Node
support was evaluated via
parsimony jackknifing (Farris et
al., 1996). Jackknife frequencies
were computed with PAUP* with
1000 replicates having 33%
deletion. Each jackknife replicate
involved a heuristic search with
20 random stepwise addition rep-
licates and TBR branch swapping
on 20 trees per replicate.

With the above analytical
procedures, 4,233 shortest clado-
grams of 1 35 steps (Consistency
Index 0.54; Retention Index 0.82;

Rescaled Consistency Index
0.44) were found in all 5,000
replicates. Figure 8 depicts clades
resolved in more than half of the
jackknife replicates, indicating
which of these groups are collapsed in the strict
consensus. All clades with more than 50% jack-
knife support are present in all shortest cladograms.

A Lamyctes-Henicops Group is one of the most
strongly supported clades in Henicopidae based
on molecular data (Edgecombe et al., 2002;
Edgecombe & Giribet, 2003b). This group,
which includes the nominate genera together
with Analamyctes , Easonobius , Eumyctes and
Lamyctopristus , is supported in 78% of the
shortest morphological cladograms, with a

jackknife frequency of 58%. Apomorphies
indicating membership of Easonobius in the
Lamyctes-Henicops Group are the alternation of
groups of short and long antennal articles (Fig.
4E) and an abrupt transition in the structure of the
fringe of branching bristles on the mandible
(characters 4:1 and 25:1, respectively). Within
the group, only Henicops and a clade that unites
Eumyctes and Lamyctopristus are present in all
minimal length cladograms and have strong
jackknife support (both 99%); the inter-
relationships of species assigned to Lamyctes ,

A NEW CENTIPEDE GENUS

281

TABLE 1. Morphological characters used in phylogenetic analysis (see Edgecombe et al., 2002; Edgecombe,
2003b, for descriptions and discussion of characters 1-51).

1 . Ocelli: (0) cluster of ocelli; ( 1 ) single ocellus.

2. Convexity of ocellus: (0) bulging; (1) flattened.

3. Antennal segmentation: (0) 1 7 or more segments; (1)15 segments.

4. Change in lengths of antennomeres: (0) gradual change in length
along antenna; ( 1 ) markedly uneven in proximal part of antenna, with
short, paired antennomeres interspersed between groups of longer
ones.

5. Long, tubular antennomeres: (0) some antennomeres equally wide
and long, proximal two antennomeres much larger than succeeding
few; ID all antennomeres longer than wide, proximal two
antennomeres not substantially larger than succeeding few.

6. Tbmosvary organ: (0) on small sclerotisation antcrovcntral to ocelli;
(i) near margin of cephalic pleurite; (2) near midwidth of cephalic
pleurite.

7. Tomdsvary organ on margin of head: (0) organ on surface of
cephalic pleurite; (1) organ in membranous field on margin of head.

8. Median furrow on head shield: (0) terminates in front of transverse
suture; ( 1 ) deep and continuous to transverse suture.

9. Shoulder in labral margin: (0) absent; (1) present.

10. Pleurites of maxillipcdc segment connected ventrally, forming a
continuous band between maxillipcdc coxostemite and stemitc of first
pedigerous segment: (O') pleurites discontinuous; (1) pleurites
continuous.

11. Shape of maxillinede coxostemite: (0) subtriangular coxostemite
with narrow, curved dental margin; ( 1 ) subtrapezoidal coxostemite
with narrow, straight dental margin; (2) narrow dental margin,
markedly V-shaped, with deep median notch; (3) subsemicircular
coxostemite with wide, convex dental margin; (4) trapezoidal
coxostemite with narrow, curved dental margin; (5) wide,
subtransversc dental margin; (6) narrow, straight dental margin
projected forward; (7) trapezoidal coxostemite with moderately wide,
weakly V-shaped dental margin.

12. Paired cusps on teeth on maxillipcdc coxostemite: (0) absent
(unpaired, conical teeth); ( 1 ) present.

13. Porodont: (0) absent; (1) translucent, seta-like porodont; (2)
conical, tooth-like pseudoporodont.

14. Proportions of maxillipede tarsungulum: (0) pretarsal section of
approximately coual length to tarsal section; ( 1 ) pretarsal section much
longer than tarsal section.

15. Dense setation on inner part of maxillipede tibia and femur: (0)
absent: ( 1 ) present.

16 Body narrowed across anterior part of trunk: (0) T1 of similar
width to head and T3; (1 ) T1 markedly narrower than head and T3.

17. Angulation (projections) of posterolateral comers of tergites: (0)
some angular or toothed; ( 1 ) all rounded.

18. Posterior margin oftergite 7 embayed, with median sector straight
and thickened ventrally: (0) absent; ( 1) present.

19. Course of posterior margin oftergite 8: (0) concave; ( 1 ) transverse.

20. Spiracle on first pedigerous trunk segment: (0) absent; ( 1 ) present.

2 1 . Row of digitiform pinnules with pointed tips along dorsal edge of
aciculae: (0) absent; (l) present.

22. Entire acicula scries simple: (0) absent; (1) present.

23. Fringe of branching bristles on mandible: (0) extends along entire
gnathal margin, skirting aciculae; ( 1 ) terminates at aciculae.

24. Ventral bristles in fringe on mandible with a wide base: (0) absent;
(1) present.

25. Differentiation of branching bristles on mandible: (0) branching
structure of bristles grades evenly along fringe; (1) abrupt transition
between row's of scale-like bristles and single row of plumose bristles.

26. Width of fringe of branching bristles dorsally: (0) fringe narrowvd
dorsallv. not developed along all bristles of furry pad; ( 1 ) fringe wide,
dense, developed along whole length of furry pad.

27. Accessory denticles on mandible all triangular, continuous
between teeth, without grooved ridges on teeth: (0) absent; ( 1 ) present.

28. Furry pad intergradcs with accessory denticles: (0) absent; (l)
present

29. Shape of first maxillary stemitc: (0) small, wedge-shaped, with
median suture; (1) large, bell-shaped, coxae not merged anterior to
stemitc, suture between coxa and stemite confined to posterior edge of
maxilla.

30. Basal joint of telopodite of first maxilla fused on inner side to coxal
process: (0) telopodite distinctly demarcated; (1) telopodite fused to
adjacent part of coxa.

31 Setae on coxal process of first maxilla: (0) dense cluster of
differentiated setae; (l) simple setae: (2) laciniate setae or plumose
amidst simple setae.

32. Coxa of leg 1 5 with long, lobate process ending in a spine: (0)
absent; (1) present.

33. Prefemur of leg 15 w'ith spurs: (0) spurs absent; ( 1 ) single ventral
spur; (2) several spurs in a wnorl

34. Coxal pores: (0) on legs 1 4 and 1 5 only; ( 1 ) on legs 13-15 only; (2)
on legs 12-15 only; (3) on legs 11-15.

35. Coxal pores set in deep groove, largely concealed by anteroventral
face of coxa in ventral view: (0) absent; ( 1) present.

36. Distal spinose projections on tibiae of legs 1-11: (0) absent; (1)
present.

legs 1-13 only; (3) strong projection on legs 1-14 only
projection on legs 1-15.

piVjWVIIVII WU

38. Tarsus of legs 1-12: (0) divided into basitarsus and distitarsus; (1 )
undivided

39. Articulation between basitarsus and distitarsus on anterior pairs of
legs: (0) distinct on dorsal side of leg; (1) fused on dorsal side ot leg,
distinct ventrally.

40. Subdivision of basitarsus indicated by paired larger setae: (0)
absent: (1) present.

41. First tarsal segment of legs 1-12 bisegmented (tripartite tarsus). ( )
absent; (1) present.

42. Accessory apical claws: (0) anterior and posterior accessory claws;
(1) posterior accessory claw only. Supposed absence of an anterior
accessory claw in Lamyctopristus validus (Edgecombe. *.003b) i. n
error. A leg associated with the holotype has an anterior and posterior
accessory claw.

43. First genital stemitc of 6 divided longitudinally into two sclentes:

(0) undivided; ( 1 ) divided.

44. Segmentation of 6 gonopod: (0) four segments with a seta-like
terminal process; ( 1 ) stout gonopod with one or two segments.

45. Number of spurs on 9 gonopod: (0) two; (1) three; (2) five to
seven.

46. First article of 9 gonopod extended as a short process: (0) absent;

(1) present. . ,

47. Claw of 9 gonopod: (0) simple (unipartite); (1) tripartite, dorsal

and ventral accessory denticles present. .

48. T6m6sv£ry organ large, positioned posteriorly on pleurite. ( )
ibsent; (1) present.

dUjvJll« jM woviiit

49 Maxillinede teeth progressively decreasing in size and spacing
medially: « 5 ) absent; (I ) present.

U1CUIOIIJ. ouwim \ ■ / I ,

50. Aciculae differentiated into two (outer and inner) rows: (0) absent
(single row of aciculae); (1) present.

51 Accessory denticles on dorsal part of mandible: (0) simple
triangular accessory denticles; (1) flattened, multi furcating scales, (2)
tuberculate scales.

52. Tergal tuberculation: (0) absent or faint; (1) strong, more

pronounced in 8 6 than 9 9 . .

53. Distitarsus of leg 1 5 divided: (0) undivided (single tarsomerc), ( )
divided into two or more tarsomercs.

54 Curvature of distitarsal part of leg: (0) straight; ( 1 ) curved.

55. Insertion of anterior protarsal acccssory claw, (O) on dorsolatera
side of main claw; (1) on ventrolateral side ot main claw mosi
heniconids have the anterior and posterior accessory claws originating
dorsolateral I v on the main claw (Fig. 5H for Easonobius ’
Fig 1G fox' Henicops maculatus). Some species of ^ihfl/am.vcres
(Paralamvctes) (Fig. 1H) and Eumyctes sinuatus (Fig. IF) have the
anterior accessory claw originating near the ventral margin ot the main
claw

IUW. _ ,

6. Definition of scutes on pretarsal accessory claws: (0) absent or
cak; ( 1 ) strong (Edgecombe & Ginbet. 2003a: character 57).

7. Definition of scutes on proximodorsal part of main pretarsal claw:
)) distinct; ( 1 ) indistinct (Edgecombe & Giribet, 2003a: character 58).

282

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Codings for 57 morphological characters listed in Table 1 .

—

Lithobius obscurus

0-0000-000 0011000000 0000000000 002200-000 0101001000 0000-00

Lithobius variegatus rubriceps

0-0000-000 7011000000 0000000000 002200-000 0101000000 0000-00

Australobius scabrior

0-0000-000 7010000000 0000000000 002200-000 0101101000 0000-00

Bolhropolys multidentatus

0-0010-000 201 1000000 0000000000 002200-000 0101001000 0000-00

Shikokuobius japanicus

-00010001 0010001010 0010010000 11120101-0 0000000000 2000000

Dichelobius Jlavens

-00010101 2010001010 0010010000 11100101-0 0000010000 2000001

Anopsobius neozelanicus

-10010101 2110001000 0010010000 11100111-0 0000010000 2000001

Anopsobius sp. nov. NSW

-10010101 2110001010 0010010000 11100111-0 0000010000 2000001

Anopsobius sp. nov. TAS

-10010101 2010001010 0010010000 11100111-0 0000010000 2000001

Zygelhobius pontis

1000010001 3000000100 0010010101 1003113000 0000000000 0000000

Cermatobius japonicus

1100010001 3001000100 0000010111 1002112070 0000100000 0011000

Lamyctes emarginatus

1001010001 0020001001 0001000001 10020101-0 0000000000 1000000

Lamyctes africanus

1001010001 0020001001 0001000001 10020111-0 0010000000 1000000

Lamvctes coeculus

-00010001 0020001001 0001000001 10020101-0 0077000100 1000000

Lamyctopristus validus

1001010001 0021000001 0007700001 10020131-0 0000200001 710117?

Lamyctopristus (Eumyctes) sinuatus

1001010001 0001000001 0001000001 10020131-0 0000000001 0101100

Henicops maculatus

1001020001 1000000001 0001000001 2002013011 1010000001 0010000

|| Henicops dentatus

1000010001 1000000101 0001000001 2002013011 1010000001 0010000

Henicops sp. nov. QLD

1001020001 1000000001 0001000001 2002013011 0Q10000001 0010000

Analamyctes tucumanus

1001010001 0000001001 0001000001 1002012000 0000000000 0000000

Analamyctes andinus

1001010001 0020001001 0001000001 1002013000 0000000000 1000000

Paralamyctes ( Paralamvctes ) spenceri

1000010101 4001000001 1000000111 1002012000 0000000000 0000000

Paralamvctes f Paralamyctes ) asperulus

1000010101 4001000101 1000000111 1002012000 0000000000 0000100

Paralamvctes ( Paralamyctes ) weberi

1000010101 4001000001 1000000111 1002013000 0000000000 0000100

Paralamyctes (Paralamyctes) tridens

1000011101 3001000001 1000000111 1002012000 0000000000 0000000

Paralamyctes (Paralamyctes) monteithi

1000111111 3001100001 1000000111 1002013000 0000000000 0000100

Paralamyctes (Paralamyctes) harrisi

10001 11111 3001 100001 10000001 1 1 1002013000 0000000000 0000000

Paralamyctes (Paralamyctes) nevemeverensis

1000010101 4000000001 1000000111 1002014000 0000000000 0000000

Paralamyctes chilensis

1000010101 4000010001 0000000011 1002013000 0000010000 0000000

Paralamyctes (Notho/agobius) cassisi

1000010101 4000010001 0100001011 1002014000 0000110000 0000000

Paralamyctes (Notho/agobius) mesibovi

1000010101 4000010001 0100001011 1002014000 0000110000 0000000

Paralamyctes (Thingathinga) gravi

1100010111 5001000101 0100000011 1002113010 0000000000 0000011

Paralamyctes (Thingathinga) Ygrayi

1100010111 5001000101 0100000011 1002014010 0000000000 0000011

Paralamvctes (Thingathinga) homerae

1100010111 5001000101 0100000011 1072017010 0000000000 0000011

Paralamyctes (Thingathinga) validus

1000010111 5001000001 0100100011 1002114010 0000000000 0000011

Paralamyctes ( Haasiella ) trail li

-00010101 6001000011 0000000011 10020121-0 0000000110 0000000

Paralamyctes ( Haasiella ) subicolus

1100010101 5001000011 0000000011 10020131-0 0000000010 0000000

Paralamyctes (Haasiella) cammooensis

1100010101 4000001011 0000000011 10020111-0 0000000110 0000000

Paralamyctes (Haasiella) ginini

1100010101 4000001011 0000000011 10020111-0 0000000100 0000000

Easonobius humilis

1001010001 0000000101 0001000001 2072012000 0000000000 0070000

| Easonobius tridentatus

1001010001 1000000101 0001000001 2002012000 0000000000 0000000

Analamyctes and Easonobius are labile.
Amongst the minimal length cladograms, the two
species assigned to Easonobius have three
alternative resolutions: monophyletic sister
group of Henicops , monophyletic sister group of
Analamyctes tucumanus, or paraphyletic with E.

tridentatus being closer to Henicops than is E.
humilis.

Parsimony jackknifing favours a closer
relationship between Easonobius and Henicops
than with Analamyctes , the former grouping
having a jackknife frequency of 58%.

A NEW CENTIPEDE GENUS

283

Easonobius tridentatus in particular shares its
shape of the maxillipede coxostemite (character
11:1) with Henicops , closely resembling species
such as Henicops dentatus in having a gently
convex dental margin with the outermost (third)
tooth set well inward of the anterolateral comer
(Fig. 5 A, B). The coxostemal shape in E. humilis
(Fig. 7 A) is instead more similar to that o fAnala-
myctes , Lamyctopristus and Lamyctes (character
11:0), and in some shortest cladograms the
similarity between E. tridentatus and Henicops is
a synapomorphy. A cluster of lacinate setae
amidst the simple setae on the coxal process of
the first maxilla (character 3 1 :2) is also shared by
Easonobius (Fig. 6H) and all species of
Henicops. A similarly positioned group of three
laciniate setae was described by Ribaut (1923,
and Ribaut’s unpublished drawings of the
holotype) in another New Caledonian species,
Lamyctes brevilabiatus Ribaut, 1923. Molecular
sequence data suggest that 'Lamyctes' brevi-
labiatus is nested within Henicops as the latter is
traditionally delimited (Edgecombe & Giribet,
2003b). This species is excluded from Easonobius
based on its indistinctly jointed tarsi on legs 1-12,
unprojected tergites (e.g., transverse posterior
margins of TT11 and 13), and bipartite first
genital stemite in the male. It is further dis-
tinguished at the species level from both known
members of Easonobius by its more segmented
antenna (38-47 articles in QM S60636, S60651,
AM KS 82580, KS 82626) and extreme
proximity of the inner two teeth on the
maxillipede coxostemite (distance between outer
and middle teeth about 2.5 times that between
middle and inner tooth).

Alternative relationships for Henicops are
favoured in some of the shortest cladograms. In
particular, Henicops and Lamyctopristus
(including Eumyctes) sometimes unite to the
exclusion of Easonobius , with this resolution
being supported by a distal spinosc projection on
the tibia of leg 14 (character 37:3) and man-
dibular aciculae differentiated into two rows
(character 50:1).

Turning to Chamberlin’s (1955) hypothesis
that Easonobius humilis is especially closely
related to South African species, an exemplar of
the latter group, Paralamyctes asperulus* nests in
a monophyletic Paralamyctes and, in most minimal
length cladograms, in P. (Paralamyctes). That
clade is defined by two unique mandibular
characters (characters 21:1 and 28:1) as well as
molecular synapomorphies (Edgecombe et al.,
2002; Edgecombe & Giribet, 2003a). The

classification of this and allied South African
Paralamyctes with Easonobius humilis based on
a single, highly homoplastic antennal character
(Chamberlin, 1955), is emphatically rejected.
The classification of Ribaut (1923), in which E.
humilis was assigned to Paralamyctes based on
its bipartite tarsi on all legs (character 38:0). is
opposed by the absence of a complete median
furrow on the head shield (character 8:0; Fig. 4A)
and a reduced, rather than bell-shaped, first
maxillary stemite (character 29:0; Fig, 6F). The
apomorphic homologues unite Paralamyctes
(Edgecombe, 2001).

ACKNOWLEDGEMENTS

I thank Suzanne Bullock, Sue Lindsay and
Yongyi Zhen for their expert assistance with
illustrations, electron microscopy, and image
editing, respectively. Geoff Monteith (Queens-
land Museum) provided collections from New
Caledonia that made this study possible. Antoni
Serra (Universitat de Barcelona) kindly reviewed
the manuscript. Eduard Stoeckli loaned types
from the Naturhistorisches Museum, Basel, and
Jason Dunlop (Museum fiir Naturkunde) hosted
study in Berlin and arranged loans.

LITERATURE CITED

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1937. Revision of the Chilopoda of New Zealand.
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L. (ed.) Zoologische und anthropologische
Ergebnisse einer Forschungsreise im westhchen
und zentralen Sudafrika ausgeluhrt in den Jahren
1903-1905. Denkschriften der Medizinisch-
naturwissenschaftlichen Gesellschaft zu Zena 14
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1914. Die indo-australischen Mynapoden. Archiv
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1 95 1 . On Chilopoda collected in North-East Angola
by Dr. A. de Barros Machado. Museo do Dundo,

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Subsidios para o Estudo da Biologia na Lunda.
(Companhia de Diamantes de Angola, Servicos
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EDGECOMBE, GD. 2001. Revision of Paralamyctes
(Chilopoda: Lithobiomorpha), with six new
species from eastern Australia. Records of the
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2003a. Paralamyctes (Chilopoda: Lithobiomorpha:
Henicopidae) from the Cape region. South
Africa, with a new species from Table Mountain.
African Entomology 1 1 : 97-1 15.

2003b. The henicopid centipede Haasiella
(Chilopoda: Lithobiomorpha): new species from
Australia, with a morphology-based phylogeny
of Henicopidae. Journal of Natural History 37.
2003c. A new species of the Gondwanan centipede
Anopsobius (Chilopoda: Lithobiomorpha) from
New South Wales, Australia. Zootaxa 204: I - 1 5.
EDGECOMBE, G.D. & GIRIBET, G. 2003a.
Relationships of Henicopidae (Chilopoda:
Lithobiomorpha): new molecular data, class-
ification and biogeography. In Hamer, M. (ed.)
Proceedings of the 12th International Congress of
Myriapodology. African Invertebrates.

2003b. A new blind Lamyctes (Chilopoda:
Lithobiomorpha) from Tasmania with an
analysis of molecular sequence data for the
Lamyctes-Henicops Group. Zootaxa 152: 1-23.

EDGECOMBE, GD., GIRIBET, G & WHEELER,
W.C. 2002. Phylogeny of Henicopidae
(Chilopoda: Lithobiomorpha): a combined
analysis of morphology and five molecular loci.
Systematic Entomology 27: 31-64.

FARRIS, J.S., ALBERT, V.A., KALLERSJO, M.,
LIPSCOMB. D. & KLUGE, A.G. 1996.
Parsimony jackknifing outperforms neighbor-
joining. Cladistics 12: 99-124.

POCOCK, R.I. 1901 . Some new Genera and Species of
Lithobiomorphous Chilopoda. Annals and
Magazine of Natural History 7: 448-451.

PORAT. C.O. von. 1893. Myriapoder fr&n Vest- och
Syd-Afrika. Bihang till Svenska Vetenskaps-
akademiens Handlingar 18: 1-15.

RIBAUT, H. 1923. Chilopodes de la NouvelJe-
Caledonie et des lies Loyalty. Pp. 1 -79. In Sarasin,
F. & Roux, J. (eds) Nova Caledonia. Recherches
scientifique en Nouvelle-Caledonie et aux lies
Loyalty. A. Zoology 3(1) (C.W. KreideFs Verlag:
Berlin).

SILVESTRI. F. 1903. Contribuzione alia conoscenza
dei Chilopodi. II. Nuove specie di Paralamvctes.
Rcdia 1: 256-257.

SWOFFORD, D. L. 2002. PAUP*. Phylogenetic Analysis
Using Parsimony (*and Other Methods). Version
4.0b 10. (Sinauer Associates: Sunderland, MA).

WURMLI, M. 1974. Ergebnisse der Osterreichen
Neukaledonien-Expedition 1965. Chilopoden.
Annalen des Naturhistorischens Museums in
Wein 78: 523-533.

NOTE ADDED IN PROOF

Anopsobius n. sp. NSW (Fig. 8, Table 2) has been formalised as Anopsobius wrighti
Edgecombe, 2003c.

GREAT ARTESIAN BASIN SPRINGS IN SOUTHERN QUEENSLAND 191 1-2000

RUSSELL J. FAIRFAX AND RODERICK J. FENSHAM

Fairfax, R.J. & Fensham, R.J. 2003 06 30: Great Artesian Basin springs in Southern
Queensland 1911-2000. Memoirs of the Queensland Museum 49(1): 285-293. Brisbane.
ISSN 0079-8835.

This study presents descriptions of Great Artesian Basin springs in south-central Queensland
from 191 1-1912. It compares these records with a second survey conducted in 1 999-2000. Of
58 springs documented in the initial survey, 42 were inactive and 13 active in the year -000.
Three were not re-located. Observations concerning fluctuations in spring flows, causes of
spring inactivity and wetland vegetation are provided. O Great Artesian Basin, bores,
springs, Queensland, history.

Russell J. Fairfax & Roderick J. Fensham, Queensland Herbarium, Brisbane Botanic
Gardens, Mt Coot-thu Road, Toowong 4066, Australia; (e-mail: russell.fairfax
@epa.q ld.gov.au); 29 January 2003.

The Great Artesian Basin (GAB) lies beneath
much of the semi-arid land in northeastern
Australia (Fig. 1) and is the continent’s most
voluminous aquifer. The Basin is charged by
rainfall that enters porous sedimentary sand-
stones outcropping along the Great Dividing
Range (Habermehl, 1982; 2001 ). Water naturally
exits the aquifer within these recharge areas as
‘recharge-rejection springs’ and as ‘discharge
springs’ on the lower, inland margins of the basin
where the age of the water may exceed 800,000
years (Radke et al., 2000). GAB springs are
regionally clustered and referred to as spring
Super-groups (Fig. 1). For the purposes of
presentation the term Spring-group is used to
represent multiple vents/springs where no
adjacent pair of springs is more than about 1km
distant. Spring-groups are often referred to by the
place name ‘Springs’ e.g. Baroona Springs. 545
Spring-groups have been documented from
Queensland (Fensham & Fairfax, in press).

Individual GAB springs have been recorded as
discharging a few to several million litres of
water per day (Habermehl, 1998; Fensham &
Fairfax, in press), corresponding to muddy areas
of a few square metres to permanently flowing
streams. Many of the wetlands contain flora and
fauna endemic to GAB springs, notably fish,
plants, snails and other invertebrates. Some
species such as the Elizabeth springs Goby
(Chlamydogobius micropterus) have only been
recorded from one isolated Spring-group and
numerous other endemic species are limited
locally or regionally (e.g. Ponder & Clark, 1990;
Ponder, 1986). Some springs are conspicuously
mounded (to 6m high) due to the accumulation of
discharged-mineral matter. Such mud springs

generally do not support vegetated wetlands and
fn Queensland are most common in the Eulo
Super-group.

Artesian bores first tapped the GAB in the late
1 870’s. Since then over 5,000 flowing bores have
been sunk: the pressure within the Basin has
dropped and currently about half of the bores still
flow. Directly related to this reduction in artesian
pressure (referred to as ‘draw-down’) about
two-thirds of Queensland springs in the discharge
areas no longer flow (Fensham & Fairfax, in
press).

The Queensland portions of the Eulo and
Bourke spring Super-groups form the current
study area (Fig. 1 ). The region can be considered
as semi-arid, with an average mean annual
rainfall at Eulo in the order of332mm/yr (Clewett
et al., 1994). These Super-groups lie within the
Warrego GAB Hydrologic Zone in which the
average pressure head has decreased by 39m
(ramje 0-1 20m) since development of the GAB
(GABCC 1 998). This zone has the greatest loss
of pressure within the GAB. Springs from the
Eulo Super-group currently discharge about a
third of their early 20th Century output of
1.6ML/day, 62% of 116 Spring-groups have
become completely inactive and a further 14/o
partially inactive (Fensham & Fairfax, in press).
That study found no inactive springs from the
Queensland portion ol the Bourke Super-group,
although Pickard (1992) reports that 15 of 45
springs were extinct from the NSW portion.

Three of the four GAB spring endemic plants in
Queensland (Myriop hy llum art esi u m ,
Sporobolus pamelae and Eragrostis sp. RJ
Fensham 3705) were recorded from the Eulo
Super-group in 1999/2000. The endangered salt

286

MEMOIRS OF THE QUEENSLAND MUSEUM

pipewort Eriocaulon carsonii was recorded from
this area (Queensland Herbarium, unpubl. data)
as well as from one spring in the NSW portion of
the Bourke Super-group (Pickard, 1992). Several
species of GAB endemic snails and isopods have
also been recorded from Eulo springs ( W. Ponder,
pers. comm.).

The dramatic decline in spring flows and
threatened GAB spring-dependent fauna and
flora confer great conservation value upon
remaining springs. The biggest threat to the water
source, i.e. draw-down of the aquifer, is being
addressed by a program to cap remaining flowing
bores and improve the efficiency of water use
(e.g. GABCC, 1998). In some areas where this
has happened there are signs that the output of
both springs (Fensham & Fairfax, in press) and
bores (GABCC, 1998) has increased. Opportun-
ities to rehabilitate springs (e.g. revegetation)
may present themselves in the future. However,
our knowledge of spring wetlands prior to their
demise is poor and unrecorded. One written
historical source is the reports of artesian bore
inspectors in the late 19th and early 20th
Centuries. Although focusing on the condition
and output of bores these surveyors also briefly
described the physical characteristics of springs
(e.g. size and flows) and modifications (e.g. wells
and bores) made to them (Fairfax & Fensham,
2002). Such descriptions enable a comparison
between past and present, and provide a bench-
mark for which spring recovery can be gauged.

This article summarises various aspects of
springs described within a survey of artesian
bores and springs conducted in 1911-1912.
Descriptions from that survey are compared with
another field survey conducted by the authors in
February 1999 and August 2000, in which a
spring was considered active if it was wet at the
ground surface or wobbled when jumped upon. A
spring was also considered active in the case of
wells when water was visible regardless of depth
below ground surface. Thus our definition of
‘active’ does not imply that the spring maintains a
natural wetland. Limitations and insights of the
original survey information and changes in
spring activity over the last century are discussed.
The current activity of these springs has been
collated within the broader study of Fensham &
Fairfax (in press).

THE 1911-12 SURVEY

The first survey was conducted by C. Ogilvie
and E. Edwards between October 1911 and April
1912, and documented within the handwritten,

FIG 1 . Map of the GAB based upon Habermehl & Lau
(1997). The shaded pattern represents the recharge
area; arrows represent modeled flow lines after Welsh
(2000). Dashed lines represent spring Super-groups
adapted from Habermehl (1982).

unpublished Artesian Reports Volumes VIII, IX
and XVI held by the Queensland State Archives
(reference numbers RSI 3037-1-8, 9 and 16).
Within these reports the locations of most springs
are either described or mapped with sufficient
detail to enable their relocation.

Ogilvie & Edwards’ 1911-12 survey recorded
56 separate springs (at least 48 Spring-groups)
within the Eulo Super-group (Fig. 2; Table 1 ). Six
of these were not surveyed as they were
considered ‘unimportant’. It can be assumed that
these springs were dry or had particularly
negligible flows. The descriptions of three
groups suggest they were completely dry and
may have been inactive prior to the first artesian
bores in the district ( 1 890’s). Flows (gallons per
day) were estimated for 22 springs. The survey
was not comprehensive, as a further 68 Spring-
groups were located for that district from other
historical sources and field surveys during 1 999
and 2000 (Fensham & Fairfax, in press). Ogilvie
and Edwards also described two Spring-groups
within the Bourke Super-group, approximately
200km east of the Eulo springs (Fig. 2; Table 1 ).

The descriptions were concise and largely
restricted to the nature of modifications made to
the springs to facilitate either greater flows or

GREAT ARTESIAN BASIN SPRINGS

287

FIG 2. Map of the Eulo district showing springs active in 1912 and 2000 (T); active in 1912 and mact.ve 2000
(A); other inactive Spring-groups 2000 (O); other active Spring-groups 2000 (•), and Spring-groups sin 2000
containing both active and inactive springs (□). The latter 3 categories comprise springs not surveyed during
1911-12. Numbers correspond to springs listed in Table 1 . Approximate current property boundaries are shown.
The inset portrays springs within the Bourke Super-group. All springs represent Spring-groups except - and 3,
10 and 16, 19 and 20, and 53 and 54.

access to clean water. Indeed, it was mainly those
springs with improvements that were visited and
described; springs that could not be made to yield
useful amounts of water were considered un-
important and generally not visited (Table 1 ). No
mention was made of evidence of use of springs
by Aboriginals (Robins, 1998) or native fauna
associated with the springs. However, incidental
notes included observations on fluctuations in
spring flows, causes of some spring inactivity
and broad botanical references (Table 1 ).

DISCUSSION

The work of Ogilvie and Edwards provides the
earliest known benchmark from which GAB
springs of southern Queensland can be assessed.

At least three of the 56 springs documented from
the Eulo Super-group were inactive when this
survey was conducted around 1912. Following
re-survey in 1999-2000, 42 (75%) had become
inactive and 1 1 remained active to some extent.
Three springs were unlocated in 1999- 2000
because their locations were not described in the
original survey. Although not all springs within
the Eulo Super-group were documented in
191 1-12, the level of spring extinction revealed
by this study is similar to the more comprehen-
sive survey of that Super-group by Fensham &
Fairfax (in press): 24% of Spring-groups active,
62% inactive, and 14% with both active and
inactive springs. The two Spring-groups

288

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1. Summary of spring descriptions, use, flows, modifications made to them by 1912 and water taste. Bore
flows were sourced from Dunstan (1913). gpd=gallons per day (1 gallon = 4.536L). A=active, X=inactive.

Spring(S)

Name

No.

Abridged Spring Description

Human Use, Quality
to Taste (191 1-1912)

Flow (Gpd)

Developments

1999-2000

Status

Eulo Super

troup

Barb

1

Several acres containing a number of dry mounds

X

Baroona ( 1 )

2

One large flowing and several smaller non-flowing
vegetated peaty springs; clumps of tall rushes,
miranda (Typha spp.)

‘Rankin [the lessee] says the flow seems to slacken
during summer, but this is rather doubtful evidence
as the total overflow is only about 500 gpd and this
could readily be evaporated from the damp ground
surrounding the well and consequently not reach the
claypan about 30 yards distant.’

‘The springs are stated to be independent of local
rainfall for several years at least.’

4 . . .this spring watered about 4000 cattle during a
drought in the early days of Tinnenburra Run but this
is certainly excessive as its maximum flow at the
bottom of the well is only 4 1 1 0 gp.d . . . '

Cooks Galley,

Shearer’s Hut; fair

4610

Well, pump,
troughing

X

Baroona (2)

3

A few overflowing and about a dozen smaller
vegetated springs; whole surface thickly coated with
duckweed, mosses, rushes and other vegetation

1000

Pump

X

Bingara
(old HS)

4

A single mud spring

Travelling public;

.good

X

Bingara (HS)

5

Four small improved springs and two large wet
non-flowing mounds

Homestead; good

500

Wells

A

Bingara

Springs

6

30-40 acres of dry mounds and wet mud springs

Small

Bore (40,000
gpd).

unspecified

improvements

X

Bitherty

7

1 dry and 1 wet mud spring with no evidence of
recent overflow. An Acacia bush locally regarded
as a sign of subterranean water present

Exception-
ally large

Well

X

Bokeen

8

Unspecified, ‘unimportant*

Outstation, boundary
riders; good

Well, pump,
troughing

X

Boomerang

9

Unspecified, ‘unimportant 4

A

Unnamed

(Boorara)

10

A group of mounds up to 18 foot high practically
all drv

X

Boorara

11

Scattered group of dry mounds 4-6 feet high

small

?

Boorara

Woolshed

12

Single wet mud spring

Woolshed

Well, bore
117,000 gpd)

X

Bullenbilla

(1)

13

About 20 mud mounds all wet at the surface, 6
overflow slightly and form small patches of
green feed

500

Bores

X

Bullenbilla

(2)

14

5 very large mounds ( 1 wet at surface) and 2
small carpet springs

5

Excavated

X

Burtanya

15

Unspecified, ‘unimportant 4

X

Bush

16

2 main improved springs and several smaller,
one of which supplies a pool 10 foot diameter by

6 inches deep; very dense clump of reeds at
troughs

Two wells,
troughing, bore
that yielded
9,000 gpd in

1 90 1 ; dry by
1913

X

Caiwarro

Mud

17

‘When visited none of the springs were actually
flowing, although 7 mounds were wet and boggy.
There are altogether about 13 mounds, the big-
gest of which is about 8 ft high and not running. I
am led to believe that the springs occasionally
overflow, but there is no regularity in the periods
of discharge’.

X

Caiwarro

Bore

18

Small mud spring

Bore (25,000
gpd)

X

Colanya bore

19

Small spring

Bore (75,000
gpd in 1913;
90,000 in 1901)

X

GREAT ARTESIAN BASIN SPRINGS

289

TABLE 1 ( Cont .)

Colanya

springs

20

1 'A miles of mounds to 1 5 feet high, some are wet
and overflow, a few permanent holes; wells over-
grown inside and out with reeds

Good

1600

3 wells,
troughing

X

Corina

21

150-200 mostly dry mounds (extending about 2
miles) 5-20 feet high

200

Reservoir,

troughing

X

Curracunya

22

A few acres of boggy flat, springs do not over-
flow to any extent

Homestead, pub

Well

X

Currawinya

23

A few mounds scattered within 1 mile diameter, 1
overflows slightly

Tank

X

Currawinya

Station

24

Unspecified; possibly containing a small bamboo
(Phragmites australis)

Homestead; excellent

600

Well

X

Dewalla

25

2 large (the main spring discharges into a grassy
bog 1 chain wide), some smaller mud springs free
of growth & several drv mounds

Travelling public;
excellent

500

Unspecified
improvement
plus bore

A

Fish

26

Group of dry mounds plus several small active;
like Umatcha Springs

Fair

100

Excavated

X

Gooncrah

27

Isolated mud spring

Outstation; good

Unspecified

improvement

X

Gooning

28

1 large, wet patches/pools (to 1 2 ft diameter) &
several minor in vicinity. Free of odour and taste
despite a dense growth of reeds around the spring

Station (dry season);
excellent

1500

Well, pump,

tanks,

reservoir

X

Gourminya

29

Unspecified, ‘unimportant’

X

Gowerah

30

Unspecified, ‘unimportant'

?

Horseshoe/

Twomanee

31

‘These springs (7 main) form practically one group
round the granite outcrops. On the first \isit to
Twomanee which was in the middle of a summer's day
(14/12/1 1 ) no well-defined springs could be found. On
this visit however at 8am distinct flows were noticed in
places that were merely wet patches on the previous
occasion. It appears from these observations that tlte
flow is not constant throughout the day though the fall-
ing off would be more noticeable in mid summer when
the evaporation is a maxim'

‘In most of iIk* springs the evaporative effect is very
marked, and especially at the main Horseshoe Spring.

I lerc tiie water Issuing from the bog surrounding the
spring (during a week's inspection ) appeared to advance
and recede with comparative regularity according to die
time of day, reaching approximately 1 chain further at

8am than at 5pm (when it began to advance). The effect
was more marked on tlie very' hot days so that observa-
tions taken at different periods might be expected to

15000

A

KapongeeA

32

Unspecified

X

Kaponyee

33

Unspecified, 3rd best spring in district

3rd big

X

Kungie

34

Boggy ground that never discharges any water,
not mud-spring like

X

McNichols

35

A few scattered mounds, ‘unimportant*

A

Nowanee

38

Unspecified, 'unimportant' (largest group in dis-

X

Mingcburra

36

Two small groups of non over-flowing springs
‘Since the bore w'as put down, he [J.G.Cooney,
driller and selector] says that several of the
springs have dried up, and it is now' safe to ride

Good

Excavated,

bore

X

Myrton

37

Mounds 2-7 ft high, a few discharge small quantity of
water. Surplus from the bore forms a small swamp
overgrown with reeds peculiar to the spring

Bore

X

Ooliman
(at bore)

39

Grassy bog 20ft wide plus smaller diy springs ‘This in-
cludes a flowing spring, which yields about 1 5<M) gal-
lons of good quality water per day. This spring is west
of the bore and cool. The other members of the group
are now' dry but formerly yielded water before the in-
terference referred to in Ooliman bore notes coincident
with the sinking of that bore’.

‘A few dry springs belonging to this group lie about 3
chains north of the bore and in the crater of one of
these Edwards found a sample of fairly rich copper
ore, which had evidently been thrown up during an
‘explosion’ of the spring/

Good

1500

Bore

(28,000 gpd)

A

290

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1 ( Cont .)

Ooliman
(at hut)

40

Two main springs and a few non-flowing im-
proved mounds ‘It was this group that was
reported to have failed on the completion of
Ooliman bore. It is situated about 1 mile south
west of the bore...’

Outstation; 1st Class

112

Wells plus
troughing

□

Talaroo bore

41

Unspecified

Bore (75,000

Ml _

X

Tareen

42

A small flowing spring

Homestead; good

Well,
troughing,
pump, tank

X

Tarko

43

Largest group of mounds in district (extending

3 miles). 3 main flowing springs, a few scat-
tered wet patches of sand, some wet and over-
flowing mounds, main spring no longer used
and overgrown with reeds

Travelling public;
excellent

3000

Troughing,

excavation

X

The Springs

44

A small group of mud springs not overflowing

Tank

?

Thorlinda

45

Group of unspecified springs

3 wells and
bore

X

Tilberry

46

2 flow ing springs and a few dry mounds

Outstation

1110

Well,
troughing,
pump, tank

A

Tunca

47

Group of springs of unspecified character

150

Well

X

Tunga

48

Unspecified '...since sinking Tunga bore the
flow has stopped and the ground is almost dry’.

Fairly large

Bore (20,000

wAl

X

Tungata

49

About 50 large dry mounds and 2 overflowing
springs; growth of tall reeds

Good

1100

Well, excava-
tion, troughing

A

Umatcha

50

A group of dry mounds to 20ft high & 100ft
diameter over several acres

X

Wanko

51

Unspecified, ‘unimportant 1

A

Wiggera

52

Best in district, one large (flows for 3 miles) and
several carpet springs. A few reeds flourish on the
edge of the pool & good feed in the watercourse
‘The supply is now very small but a boundary
rider who was on the run before the adjacent bore
was sunk states that the spring gave out when the
bore first struck water. Originally the spring
maintained the same length of drain as the bore
now does. fWaterl Quality; excellent'.

Good/excellent

42100

Troughing,

excavation,

bore

X

Wirrarah

(Werrerah)

53

A group with one flowing spring and 12 wet
mounds are Imile southeast ‘The flow was
originally, judging from the description of one
of the station hands, about 10000 gpd, but now
it docs not exceed 200 gpd (difficult to esti-
mate exactly)’.

Good

10000

Well, bore

X

lm S of
Werrerah

54

12 mounds wet at the surface but non- flowing

X

Wooregym

55

A group of springs in an area about 220 by 1 1 0
yards. Other springs not improved give off
lartie auantities of gas (as does Wiggera)

Homestead; good

6 wells, bore
(48,000 gpd)
in near vicinity

A

Youlcen

(Youlain)

56

2nd best in district, 2 main and several smaller
springs, 5 acres of swamps, growth of green grass,
water temperature 103°F, cold spring unlocated

Good

30000-40000

X

Bourke Su

pergrou

- - - - .

Tego

57

A small group of mud springs ‘Robertson
(Tego) states that the discharge apparently docs
not fluctuate during the various seasons, but of
course the effect of ev aporation in summer
causes the general appearance to alter’.

200

Well, tank,
dam

A

| Towry

58

Boggy areas, bogs and pools plus V: mile2 of
smalf springs. Patches of water grass, bog of
miranda (Typha spp.) 'The discharge docs not
vary with the seasons and the springs arc a
fairly regular watering place*.

Usual

700

Excavated,

bores

A

GREAT ARTESIAN BASIN SPRINGS

291

surveyed in the Bourke Super-group in 1911
contained active springs in 1 999.

Some of the currently inactive springs are
mounded mud springs that previously flowed
relatively small quantities of water. However,
larger springs have also stopped flowing,
including the largest three surveyed: Wiggera,
Youlain and Kaponyee Springs (Table 1). Other
springs had ceased to flow altogether by 1912
(e.g. Mingeburra. Ooliman, Tunga), and Ogilvie
& Edwards documented that some springs, such
as Werrerah Springs had greater flows prior to
bores being sunk (Tabic 1). The original spring
flow estimates and descriptions should therefore
be treated as underestimates of the pre-artesian
bore era.

Many bores were sunk adjacent to springs. For
example, from Bingara and Dynevor Downs runs
Ogilvie and Edwards recorded that ‘All the bores
on the runs, 6 running and 3 abandoned, were put
down within a few yards of mud springs ...’
Dunstan (1913) noted that some bores on runs
that contained springs, including Dynevor
Downs had reduced Hows within a few years
after they were sunk (Table 1). Several of the
bores in the district initially flowed more than
lOML/day, and recent bore-flow measurements
are 10% of the original estimates (Table 2). The
large flows from bores and their proximity to
springs almost certainly accounts for the
observed reduction in spring-flows. None of the
springs active in 1999-2000 appear to have
increased in flow since 1911-12.

It is important to consider natural fluctuations
in spring flows when assessing longer-term
changes in flow rates (e.g. Williams & Holmes,
1978). Ogilvie & Edwards report that Towry
Springs had a constant flow rate, the flow of
Horseshoc/Twomanee fluctuated with daily
evaporation and the flow from Tego Springs did
not apparently fluctuate between seasons
although a summer evaporation effect was
evident (Table 1). However, in most cases such
changes would have been inconsequential com-
pared to estimated flow rates and spring activity.
Irregular discharges from mud springs were
recorded for Caiwarro and an occasional
‘explosion' was reported for Ooliman Springs
(Table 1 ). This is consistent with anecdotes from
landholders regarding mud spring activity
collated during the 1999-2000 survey. Some
large springs have only become inactive in recent
decades. The habitat notes on a Queensland
Herbarium specimen of Cyperus laevigatus

indicate that Kaponyee spring was active in 1971
and a local resident has confirmed that Wiggera
spring was active in the 1960s (I. Pike, pers.
comm.).

Two-thirds of all springs recorded by Ogilvie &
Edwards had been modified in some way prior to
1912, mostly by constructing wells. Wells were
often boxed (lined) to a level above the natural
outlet of the vent thus eliminating any wetland
associated with the spring.

At least 10 of the 14 Eulo Super-group springs
whose descriptions included references to
vegetation have become inactive. Of the
remaining four, two wetlands are supplied by
abandoned bores (Ooliman & Dewalla), one has
been decimated by stock (Tungata) and
Horseshoe has been excavated to form a small
dam. None of the 14 contained GAB endemic
plants during the recent survey. Descriptions
such as bogs, swamps, peat and carpet springs
were applied to seven springs (Table 1). Such
descriptions are compatible with situations in
which GAB endemic plants have been observed,
both in the Eulo district and elsewhere in the
GAB (Queensland Herbarium unpub. data) and
may represent former habitat. Wiggera Springs
was the largest spring in the district and described
as comprising carpet springs. A local resident
recalls the endangered Eriocaulon carsonii from
that spring when it was active (I. Pike, pers.
comm.).

This study provides further evidence of the
impact of bores on springs. The positive news is
that examples exist from within the study area of
increased artesian pressure since the bore-
capping program (GABCC, 1998). I hus
completion of the bore rehabilitation program
and judicious prioritising of GAB water use may
enhance flows for remaining springs and could
even reactivate springs.

ACKNOWLEDGEMENTS

This project was partly funded by the
Endangered Species Program (Environment
Australia). Ian Pike from Eulo is thanked for
sharing his enthusiasm, time and tine date wine.
The landholders of all relevant properties are
thanked for their kind assistance, without which
the surveys would not have been possible. The
Department of Natural Resources and Mines
provided information on bores in the region and
historical maps which were valuable in locating
springs. Steve Eastaugh’s companionship and
assistance in the field is also greatly appreciated.

292

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Bores from the Eulo 1 :250 000 map sheet for which pre- 1913 and recent flow rates are available. Flow
rates provided are the maximum when uncontrolled. Initial flow rates of named bores are sourced from Dunstan
(1913). Bore information is courtesy the Department of Natural Resources and Mineswith some initial flow
rates sourced from Dunstan (1913). ‘Controlled' applies to bores that have been capped.

Bore Name/ number

Pre- 191 3 flow
(L/day)

Recent year of
inspection

Bore status

Recent flow (L/day)

% pre-1913 flow |

4560

285,120

1992

Controlled

144,288

50.6

Mulgar(9) 4553

998,01 1

1992

Un-controlled

340,416

34.1

1491

31,104

1956

Controlled

8,640

27.8

Kungie 4550

1,168,655

1972

Un-controlled

304,992

26.1

403

16,498.080

1987

Controlled

4,192,128

25.4

Tunka 1617

13,608

1997

Un-controlled

3,456

25.4

Taleroo 1619

340,200

1937

Controlled

76,896

22.5

1831

307,584

1992

Controlled

63,936

20.8

Wooregym 1488

217,728

1974

Un-controlled

43,200

19.8

Woolshed(2) 1616

90,720

1997

Un-controlled

17,280

19.0

4547

2.445,984

1993

Un-controllcd

315.360

12.9

2272

5,069.088

1987

Un-controlled

595,296

11.7

4551

12,274,848

1992

Controlled

1,294,272

10.5

1821

5,908,896

1992

Controlled

610,848

10.3

4552

1,249,344

1982

Un-controlled

67,392

5.4

2276

6,590,592

1985

Un-controlled

330,912

5.0

Colanya 1613

408.240

1979

Un-controlled

19,872

4.9

Bingara 1486

181,440

1997

Un-controlled

6,912

3.8

4561

5,227,200

1982

Un-controlled

145,152

2.8

402

11,643.264

1987

Un-controlled

203,904

1.8

Tunca 1487

90,720

1997

Ceased

0

0

Tarko 1490

45,360

1911

Ceased

0

0

Woolshed 1614

77,112

1969

Ceased

0

0

Caiwarro 1820

113,400

1950

Ceased

0

0

1822

12,960

1911

Ceased

0

0

2275

432,000

1928

Controlled

0

0

Ooliman 2427

127,008

1960

Ceased

0

0

2429

104,544

1937

Ceased

0

0

Mulgarfl I ) 4554

4,091,904

1943

Ceased

0

0

Boortra 4556

12.273,120

1959

Ceased

0

0

Tinnenburra( 1 ) 4558

4,089,312

1915

Ceased

0

0

Mulgar(lO) 4559

136,080

1974

Ceased

0

0

Total

92,543,226

8,785,152

9.5

LITERATURE CITED

CLEWETT, J.F., CLARKSON, N.M., OWENS, D.T. &
ARBRECHT, D.G 1994. Australian RAINMAN.
Version 2.2. (Department of Primary Industries:
Brisbane).

DUNSTAN, B. 1913. Queensland mineral index and
guide. Queensland Geological Survey Pub. No.
241.

FAIRFAX, R.J. & FENSHAM, R.J. 2002. In the
footsteps of J. Alfred Griffiths: a cataclysmic
history of Great Artesian Basin springs in

Queensland. Australian Geographic Studies 40:
210-230.

FENSHAM, R.J. & FAIRFAX, R.J. in press. Spring
wetlands of the Great Artesian Basin,
Queensland, Australia. Wetlands Ecology and
Management.

GREAT ARTESIAN BASIN CONSULTATIVE
COUNCIL (GABCC). 1998. Great Artesian
Basin Resource Study (November 1998). (Great
Artesian Basin Consultative Council: Canberra).

HABERMEHL, M.A. 1982. Springs in the Great
Artesian Basin, Australia - their origin and nature.

GREAT ARTESIAN BASIN SPRINGS

293

Bureau of Mineral Resources Geology and
Geophysics, Australia Report 235: 1-50.

1 998. Section 1 .7 Hydrogeology. Pp. 44-60. In Cox,
R. & Barron, A. (eds) Great Artesian Basin
Resource Study, November 1998. (Great
Artesian Basin Consultative Committee:
Brisbane).

200 1 . Hydrogeology and environmental geology of
the Great Artesian Basin, Australia. Pp. 127-143.
In Gostin, V.A. (ed.) Gondwana to Greenhouse -
Australian Environmental Geoscience.
Geological Society of Australia Special
Publication 21.

HABERMEHL, M.A. & LAU, J.E. 1997. Hydro-
geology of the Great Artesian Basin (Map at 1:2
500 000). (Australian Geological Survey
Organisation: Canberra).

PICKARD, J. 1992. Artesian springs in the Western
Division ofNew South Wales. Graduate School of
the Environment, Working paper 9202.
(Macquarie University: Sydney).

PONDER, W.F. 1986. Mound springs of the Great
Artesian Basin. Pp. 403-420. In Dc Dekker, P. &

Williams, W. (eds) Limnology in Australia.
(CSIRO: Melbourne).

PONDER, W.F. & CLARK, GA. 1990. A radiation of
hydrobiid snails in threatened artesian springs in
western Queensland. Records of the Australian
Museum 42: 301-363.

RADKE, B.M., FERGUSON, J., CRESWELL, R.G.,
RANSLEY, T.R. & HABERMEHL. M.A. 2000.
Hydrochemistry and implied hydrodynamics of
the Cadna-owie-Hooray aquifer. Great Artesian
Basin. (Bureau of Rural Sciences: Canberra).

ROBINS, R. 1998. Archaeological investigations at
Youlain Springs, southwest Queensland.
Memoirs of the Queensland Museum, Cultural
Heritage Series 1 : 57-74.

WELSH, W. 2000. GABFLOW: A steady state
ground-water flow model of the Great Artesian
Basin. (Bureau of Rural Sciences: Canberra).

WILLIAMS, A.F. & HOLMES, J.W. 1978. A novel
method of estimating the discharge of water from
mound springs of the Great Artesian Basin,
central Australia. Journal of Hydrology 38:
263-272.

1

AN OBSERVATION OF CALLING NORTHERN TINKER FROGS ( TAUDACTYLUS
RHEOPHILUS) ON MOUNT BELLENDEN KER

ALASTAIR B. FREEMAN

Freeman, A.B. 2003 06 30: An observation of calling northern tinker frogs on Mount
Bcllenden Ker. Memoirs of the Queensland Museum 49(1): 295-297. Brisbane. ISSN
0079-8835.

Since its ‘rediscovery’ in 1 996 the northern tinker frog has only been recorded spasmodically
on Mount Bcllenden Ker with calling males last recorded in 1996. In December 2000 calling
northern tinker frogs were recorded on seven occasions during two trips. Subsequent trips by
QPWS staff and James Cook University researchers have failed to detect the species at the
site. It is concluded that this is a critically endangered species and some basic
recommendations are detailed for future monitoring. □ Northern tinker frog, calling,
management, endangered, Mount Bellenden Ker.

Alastair Freeman, Ecosystem Management Unit, Queensland Parks and Wildlife Sendee,
PO Box 834, Atherton 4883 (e-mail: Alastair.Freeman@epa.qld.gov.au); 25 September
2002 .

Taudactylus rheophilus is an endangered
species (Queensland Nature Conservation Act
1994, Environmental Protection and
Biodiversity Conservation Act 1999) known
from Carbine Tableland, Lamb Range, Thornton
Peak and Mt Bellenden Ker in the Wet Tropics
(McDonald, 1992). Between 1991 and 1996 this
species was not recorded in the wild and it was
assumed that it had declined or even disappeared.
In 1996 they were ‘rediscovered’ on Mt Lewis
and Mt Bellenden Ker (Marshall, 1998). Since
this time reports of this species at both these sites
have been few and spasmodic (Hero et.al., 2002;
D. James, pers. com.; E. Bolitho, pers. com.;
Keith McDonald & A.B. Freeman, unpubl. data).
No other populations have been located despite a
significant search effort, particularly on the
Carbine Tableland (K. McDonald & A.B.
Freeman, unpubl. data).

The records off rheophilus on Bellenden Ker,
between 1996 and November 2000 include five
individuals recorded along 150m of stream in
1996 (Marshall, 1998), one juvenile captured in
1998 (Hero ct. al., 2000) and an unconfirmed
observation of a Taudactylus tadpole in 2000 (E.
Bolitho, pers. com.). The latter was made by a
researcher with 4 years experience with Taudac-
tylus tadpoles from mid-east Queensland and was
based on the type of pool that it was observed in
and the behaviour of the tadpole. Unfortunately
the animal was not captured so a confirmed
identification could not be made.

This paper details observations of T. rheophilus
made by QPWS station the 12th, 13th, 18th, 19th
and 20th of December 2000 on Mt Bellenden Ker

and three subsequent trips on the 3rd September,
2001 and 23rd and 24th of January, 2002 and the
26th April 2002.

STUDY AREA

Mt Bellenden Ker is the second highest peak in
Queensland at 1582m and part of the Bellenden
Ker Range. The summit vegetation is low growing
simple microphyll vine-fern forest (Tracey,
1982). On Mt Bellenden Ker T. rheophilus is
known from a perennial stream in the upper
catchment of the Mulgrave River, 600m SE of the
summit (17°15’56’\ 145°5r22”).

METHODS

The first T. rheophilus sighting made by the
QPWS team occurred on the 12th December
2000 during an opportunistic survey associated
with an ongoing frog monitoring program.
Subsequent searches concentrated on the area
where the initial observation had been made.
Active searching (turning rocks) was abandoned
when it become apparent that this was likely to
cause damage to the habitat. Subsequent
sampling involved passively recording calling
individuals.

When a T rheophilus was seen or heard for the
first time air temperature and humidity along
with general weather information were recorded
for the site (the one exception for this was the
morning of the 13th).

For the two individuals that were captured
snout vent length (SVL) was measured to the
nearest 1mm using 200mm dial calipers and
weight was taken to the nearest 0.1 gm using a

296

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1 . Sightings of T. rheophilus on Bellenden Ker. * Tissue samples were taken for analysis by disease
researchers at James Cook University, Townsville from two individuals. These samples subsequently came
back as negative for chytrid fungus.

Date

Time

Weather

No. of
frogs

Site

' i

Measurements

12 Dec

20:00

Overcast, Light showers, Air
temp 14.5BC, Humdity 94%

1*

Calling between rocks in crevice 52cm
above stream. Captured by hand.

SVL: 24.9mm Wght: l.65g

Sex: 6

13 Dec

08:10

Overcast. Light showers. Air
temp cool.

1

Calling amongst rocks and streamside
vegetation. Ceased calling 08:20.

18 Dec

15:00

Overcast, Light rain. Air temp
18.5BC, Humidity 95%

2*

Found under rock on edge of water.
One captured by hand.

SVL: 20.5mm Wght: 0.85gm
Sex: ?

18 Dec

15:30

Overcast, Heavy rain. Air temp
!8.0Bc, Humidity 100%

2

Calling amongst rocks and streamside
vegetation.

19 Dec

08:00

Overcast, Light rain, Air temp
16.0BC, Humidity 100%

2

Calling amongst rocks and streamside
vegetation. Still calling at 09:00.

19 Dec

15:45

Overcast, Heavy rain. Air temp
I6.0BC, Humidity 100%

2

Calling amongst rocks and streamside
vegetation. Calling intermittent before
petering out anmnd 16:05.

20 Dec

08:10

Overcast, Light showers. Air
templ6BC, Humidity 95%

1

Calling amongst rocks and streamside
vegetation. Calling continued at
irregular intervals until 09:10.

lOgm Pesola™ spring balance. Sex of the male
was ascertained by the presence of nuptial pads.

RESULTS

The initial observations, on the 12th of
December 2000, were made by A.B. Freeman, L.
Dwyer, C. Schmidt. During this trip two calling T.
rheophilus were heard over two days. One of
these was captured to confirm identification. This
was the first time calling T. rheophilus had been
recorded on Bellenden Ker since 1996. A further
trip was made for two days during the following
week in an attempt to clarify the status of T.
rheophilus on Mt Bellenden Ker. This was after a
trip to the Carbine Tableland to establish if T.
rheophilus was active at this site.

Approximately 18 hours was spent at the
Bellenden Ker site over the two trips, with
searching taking place during the day and the
night. Most of this time was spent at the site
where frogs had previously been recorded.

T. rheophilus was observed or heard seven
times during the two trips to Bellenden Ker
(12- 13th December and 18-20th December)
(Table 1). Observations included calling individ-
uals, visual sightings and individuals in hand.
Two individuals were captured by hand, one
individual was observed but not captured and
another 2-3 individuals were heard.

Over 400m of stream was searched above, and
below, the site while the frogs were calling. Most
of the search effort concentrated on downstream
habitat, as it appeared to be more suitable for T.

rheophilus although historically the whole
stream had T. rheophilus (K. McDonald, pers.
com.). All the observations of T. rheophilus were
made in an area of habitat less than 9m 2 in size.

On the 21st December, 2001 a trip was also
made to the Carbine Tableland in the hope that
the activity on top of Bellenden Ker would be
replicated at the other site where they were
known to be present. No activity was detected on
this trip.

Subsequent trips were made in September
2001, January 2002 and April 2002 to the
Bellenden Ker site. The September and April
trips were made to ascertain if breeding had taken
place by checking the site for tadpoles. None
were detected. The visit in January took place
over two days with the primary aim of surveying
the stream day and night, no activity was
detected.

DISCUSSION

The number of individual frogs within this area
appears to be small. A conservative estimate of
the population was made at 3-5 males. While it
cannot be said for certain that frogs do not occur
outside this area, searches both upstream and
downstream of the site have not located other I
rheophilus. This includes four previous searches
by QPWS staff of the stream during earlier trips
to the site over a period of two years. These
searches were carried out at similar times of the
year in similar conditions. Experienced frog
researchers from James Cook University also

OBSERVATIONS OF CALLING NORTHERN TINKER FROGS

297

made ten trips to the summit of Bellenden Ker
between the 12th January and the 2nd March
2003. On each of these trips the stream where T.
rheophilus was recorded was checked for frog
activity with none being recorded (E. Bolitho,
pers. com.; Richard Retallick, pers. com.).

There is little doubt that T. rheophilus is a
critically endangered species surviving in very
low numbers on Mt. Bellenden Ker and possibly
the Carbine Tableland. The Carbine Tableland
population has not been detected since January
1999 (K. McDonald & A.B. Freeman, unpubl.
data). I am concerned by the lack of T. rheophilus
activity recorded on Bellenden Ker since
December 2000. Three trips between January
2001 and April 2002 have failed to find any sign
of the species.

T. rheophilus is not an easy species to work
with on Bellenden Ker. The specific habitat of the
frogs is notable for the dense ground layer of
Helmholtzia acorifolia on granite boulders.
Calling individuals are difficult to locate visually
in the densely vegetated rocky terrain. It was
noticeable that the actions of just one person
searching for calling frogs amongst this habitat
resulted in significant trampling of the vegetation
on part of the site. Future monitoring should
concentrate on censusing calling frogs during the
day and night or nighttime searches for
non-calling individuals out on vegetation or
rocks. Active searching by turning and
dismantling habitat should be kept to a minimum.
Visits to the specific site of T. rheophilus other
than conducting monitoring should not be
permitted.

MANAGEMENT RECOMMENDATIONS

The population should be monitored more
frequently than is currently the case particularly
over the early wet season months (Oct-Dec).

Visits to the specific site by people other than
those directly involved in the monitoring
program should not be permitted.

Future monitoring should concentrate on the
census of calling animals and night time searches
for non-calling individuals out on vegetation and
rocks. Active searching involving disturbing
habitat should be kept to a minimum.

ACKNOWLEDGEMENTS

My thanks to Carol Schmidt and Lisa Dwyer
who provided invaluable field assistance and to
Amanda Freeman and Keith McDonald who
commented on drafts of this paper. Keith
McDonald, Harry Hines, Chris Marshall and
Emily Bolitho provided comments and
information that was much appreciated.

LITERATURE CITED

HERO, J-M, HINES, H.B.. MEYER, E., MORRISON,
C. & STREATFEILD, C. 2002. New records of
“declining" frogs in Queensland, Australia (-April
1999). In Nattrass, R. (ed.) Frogs in the
community. Proceedings of the Brisbane
Symposium 13-14 February 1999. (The
Queensland Frog Society: Brisbane).
MCDONALD, K.R. 1992. Distribution patterns and
conservation status of north Queensland
rainforest frogs. Queensland Department of
Environment and Heritage Conservation
Technical Report 1 : 1-51.

MARSHALL, C. 1998. The reappearance of
Taudactylus (Anura: Myobatrachidae) in north
Queensland streams. Pacific Conservation
Biology 4: 39-41.

TRACEY J 1 982. The vegetation of the humid tropical
region of Queensland. (CSIRO Publishing:
Melbourne).

1

FAUNAL SURVEY OF NEW ENGLAND. V. THE LIZARDS AND SNAKES

HAROLD HEATWOLE, JOHN de BAVAY AND PAUL WEBBER

Heatwole, H., de Bavay, J. & Webber, P. 2003 06 30: Faunal survey of New England. V. The
lizards and snakes. Memoirs of the Queensland Museum 49(1): 299-325. Brisbane. ISSN
0079-8835.

A 34-year study (1966-2000) of the lizards and snakes of the New England region of New
South Wales yielded 1 4 species of geckos, 4 of pygopodids, 8 of dragons, 3 of goannas, 47 of
skinks, and 30 of snakes, for a total of 106 species. Four additional species reported by Swan
(1990), and an additional one from the Atlas of NSW Wildlife brings the total known
squamatan fauna of New England to 1 1 1 species. Most terrestrial families of squamates in
New' England represent 10-15% of the total Australian species. The exceptions arc the
families Elapidae (26%) and Colubridae (18%). Generic representation is higher and more
variable. A fifth of the species are widespread. Other species are predominantly separated
into eastern and western faunal groups. In some species the Great Dividing Range coincides
with east-west distributional boundaries. In others, the borders of the distributional range lie
one side or the other of this topographical feature. In both western and eastern groups, there
are species associated mainly with either the northern or southern quadrants; otherwise
distributions of only a few species are oriented w ith respect to the north-south axis. There
were no strictly central species. Some species are represented by only one or a few records
and the full extent of their distributions may not be appreciated, or they may be sparsely
distributed, rare, or cryptic. A number of species were distributed randomly with regard to
particular environmental attributes; these were unlikely to be affecting distribution within
New England. Where correlations did exist, causal effects could only be hypothesised, but
optimal habitats could be defined by over-representation in localities with particular levels of
elevation, rainfall, or temperature, or characterised by certain types of vegetation, rock or
soil. □ New England, snakes, lizards, faunal survey, distribution, habitat.

Harold Heatwole, Department of Zoology', North Carolina State University, Raleigh NC
27695-7617, USA; John de Bavay, School of Biological Sciences, University of New
England, Armidale 2351, Australia; Paul Webber, Webber and Associates, 125 Taylor
Street, Armidale 2350, Australia; 15 September, 2001.

An initial paper of the faunal survey of New
England described the general geography of the
area (Heatwole & Simpson, 1986); this was
followed by accounts of the gastropods (Simpson
& Stanisic, 1986), birds (Ford & McFarland,
1991) and frogs (Heatwole et al., 1995). The
present paper reports on the lizards and snakes.

There have been several interpretations of the
boundaries of New England (see Lea et al., 1977
and electoral maps). We recognise the boundaries
as defined in the previous papers of this series
(Fig. 1).

Treatment of the fauna is at the spec tic level;
subspecific designations are not included.

METHODS

In the interests of conservation, we collected
only a few voucher specimens of each species
from any given area. Nevertheless, over the 34
years of the study (1966-2000) we collected and
identified more than 6,000 specimens of snakes
and lizards and deposited them in the Australian
Museum.

This material was augmented by other
herpetological collections in the Australian
Museum (up to 2000), and by the holdings of the
Museum of Victoria, and the Queensland
Museum (both up to 1996). Initially, we eliminated
records referring to specimens that had been
discarded or lost, or which had deteriorated
beyond accurate identification, corrected some
misidentifications, and verified or corrected data
for unusual specimens such as geographical
outliers. After this editing, we relied heavily on
the registers of those museums. Especially
important has been the progressive upgrading by
the Australian Museum of their herpetological
collections, whereby incompletely unidentified
specimens were identified, previously identified
specimens checked for accuracy, nomenclatural
changes incorporated, and geographical co-
ordinates and other data verified or corrected.

Taxonomic nomenclature follows that of
Cogger (2000) and Cogger et al. (1983), but
incorporates recent name changes and generic
allocations by King (1983), Wells & Wellington

300

MEMOIRS OF THE QUEENSLAND MUSEUM

( 1 985), Ingram & Covacevich ( 1 988), Hutchison
(1990), Hutchison et al. (1990) Sadleir (1990),
Rawlinson (1991), Greer (1992, 1997), Hutch-
ison & Donnellan (1992), Couper et al. (1993,
1994, 1997), Sadleir et al. (1993) and Shea
(1995a, 1995b).

A number of surveys and environmental
consultancies for specific purposes amassed a
corpus of data relating to the New England area.
Specimens arising from these studies were deposited
in the Australian Museum and automatically
became part of our database. In addition, a large
number of the records were not based on voucher
specimens, but rather upon field identifications
of varying reliability. We have not included any
sight records in our database but do discuss them
below where relevant.

Our final, edited dataset, consisting of our
collections plus the other acquisitions of the three
museums, is based on more than 12,800
specimens that were collected from the late 19th
century to the end of the twentieth century. Using
the statistical package, StatView, locality data
were formatted for the Arc View Mapping System
to produce distribution maps of each species. In
this system, multiple records of a given species
for a particular locality were grouped as a single
entity and latitudes and longitudes from museum
records were converted to a decimal degree.

The final edited dataset totalled 4,045 records
and comprised 3,175 localities within the
boundaries of New England (Fig. 1), and 888
records from sites immediately extralimital (Figs
2, 3-108) that were included for mapping
purposes. This has been placed in each of the
three museums both as hard copies and CD ROM.

The New England survey of frogs (Heatwole et
al., 1995) related species distributions to four
environmental parameters: mean daily minimum
temperature for July, mean annual rainfall,
elevation, and vegetation. Because many
squamates behaviourally thermoregulate, are
cryptic, or are substrate-dependent in various
ways, an enlarged set of parameters was
employed, including soil type and rock type.
Heatwole & Simpson (1986) presented a map of
the lithology of New England based on ‘An Atlas
of New England’ (Lea et al., 1 977). For the present
paper an improved map, derived from the New
South Wales Department of Mineral Resources
1/25000 Geological map, was used.

The soil map derives from Lea et al., 1 977. This
was digitised, and the map prepared using the
Environmental Research mapping Program

FIG 1. Collecting grid and study area within New
South Wales indicating the main towns, connecting
roads, and the Great Dividing Range (GDR). Inset
shows the location of the study area in New South
Wales.

(ERMS) in the Armidale office of the National
Parks and Wildlife Service of New South Wales
(Ferrier,1989).

The vegetation map is based on a modification
of the classification of vegetation systems for
northeastern New South Wales (NSW National
Parks and Wildlife Service, 1994). The nine
categories of that classification were lumped into
three categories: ‘disturbed’ (including pastoral
and agricultural land), ‘dry forest and woodland’
and ‘wet forest’ (including rainforest).

New, more detailed maps of elevation, rainfall
and temperature were prepared. The
Environmental Research Mapping System
(ERMS) in the Armidale office of the National
Parks and Wildlife Service of New South Wales
(Ferrier, 1989) was used to prepare maps of the
distribution of soils, and the distribution of
various variables.

Long-term monthly means of minimum
temperature, maximum temperature, and
precipitation were derived by linking a gridded
digital elevation model with climate surface
models developed by Hutchinson (1989). These

NEW ENGLAND LIZARDS AND SNAKES

301

FIG. 2. Location of all collecting sites for squamates in
the present study, both within the study area and
immediately extralimitally.

monthly estimates were then used to derive, for
each grid cell in the study area, the mean
minimum temperature of the coldest month, the
mean maximum temperature of the hottest
month, and the mean annual rainfall. Values for
the squares of the grid were grouped into a series
of categories. The five categories for mean
minimum temperature (°C) of the coldest month
were: -4° to -2°; -1°; 0°; 1°; 2°-6°. Those for the
mean maximum temperature (°C) of the hottest
month were: 2 1 °-23°; 24°; 25°; 26°-28°; 29°-32°.
Six categories for mean annual rainfall (mm)
were: 700; 700-800; 801-900; 901-1000; 1001-
1100; >1100. Mean elevations (m) of squares
were grouped as follows: 0-700; 701-900; 90 1 -
1 000; 1 00 1 - 1 1 00; 1 1 0 1 - 1 600. Finally squares of
the grid were tallied as to the type of vegetation,
soil and parent rock they contained and the data
grouped accordingly. Vegetation: disturbed;
woodland and dry forest; wet forest. Soils: RP
(red podzolic); YS/YP (yellow sol/yellow pod);
YP/GP (yellow pod/gley pod); CP (chocolate
prairie); YP (yellow podzolic); BE (black earth
prairie); all others (the remaining, less common
soil types combined). Type of parent rock:
sedimentary rocks; granite; basic igneous rocks;
acid volcanics; all others (the remaining, less
common types of rocks).

For statistical treatment, the proportion of grids
falling into these various categories was
calculated separately for each environmental
attribute. Then from these proportions, the
expected number of grids in each category
expected to be occupied by a given species was
calculated and the observed values compared to
expected values and tested for statistical sig-
nificance by chi-squared tests, using a rejection
level of 5%. These tests were conducted in 1996
on the database available at that time and not
recalculated to include the few later acquisitions;
only localities located within the boundaries of
New England (Fig. 1) were included in the
statistical analyses.

For valid statistical testing, no category should
have an expected value less than live. Some
categories of vegetation, soil type and rock type
were sparsely represented in the region and did
not meet this requirement. To overcome this, the
smaller categories were lumped to raise their
collective expected value into the appropriate
range. Some environmental parameters had more
disparate frequencies of categories than did
others. For example, in vegetation systems,
rainforest and other wet forest occupy only a
small proportion of the total area of New
England, and there were relatively tew collecting
localities located within it. Nevertheless, it was
unrealistic to lump these important habitats with
those of a distinctly different character merely in
order to allow the mechanics of statistical testing
to proceed. For this reason, in order to reach the
criterion of an expected value of five for
rainforest/wet forest, only species of squamates
with a high number of occurrences (N) in New
England could be tested. Other parameters, with
more uniform frequencies of environmental
categories, could be tested for species with a
lower N. The minimum values of N for the
different environmental parameters after
lumping of categories were: Elevation * 28; Rock
Type = 34 for four categories (an N of 95 allowed
testing of five categories); Maximum
Temperature = 36; Rainfall = 46; Soils - 48 for
five categories and 80 for six; Minimum
Temperature = 49; Vegetation System = 74.

Two examples (Table 1 ) illustrate the technique.
( 1 ) Eulamprus quoyii in relation to elevation: this
species occurred in 64 grid squares in the study
area. Since 26.39% of the grid squares had a mean
elevation of 0-700m, if this species were
distributed at random with respect to elevation,
then it would be expected to occur in 16.89 grid
squares at this elevation (26.39% of 64); the

302

MEMOIRS OF THE QUEENSLAND MUSEUM

observed value was 1 1 . Similarly expected values
were 1 1 .96 for 70 1 -900, 1 0.42 for 901-1 000, 1 2.2
for 1001-1100, and 12.44 for 1101-1600. The
corresponding observed values were: 8, 16, 13
and 16. The chi-square value of the comparison
of observed to expected values was 7.414
(0.25>P>0.10). Since P>0.05, E. quoyii was con-
sidered to be distributed randomly with respect to
elevation, within the range of elevations
occurring within the study area. (2) Ctenotus
robustus in relation to rainfall. The expected
(observed) values for the 46 grid squares in which
this species occurred was: 4.50 for <700mm
rainfall, 14.86 for 700-800mm, 9.17 for 801-
900mm, 5.82 for 90 1-1 000mm, 5.43 for 1001-
1100, and 6.17 for >11 00mm. The respective
observed values were 8, 23, 8, 5, 2 and 0. The
chi-square value was 15.197 and 0.01>P>0.005
(highly significant). Thus, this species was not
randomly distributed with respect to rainfall.
Note that the observed values were greater than
expected by randomness in the first two
categories (<700mm and 700-800mm) and less
than expected in all the rest. The optimal habitat
for C. robustus therefore seems to be the drier
parts of the area.

For some species, over-representation occurred
only in one category, in other species over-
representation occurred over a broader range of
values and encompassed several categories.

Copies of the detailed statistical tests, including
all observed and expected values, chi-squares
and P values, as well as histograms of frequency
distributions of environmental parameters, have
been placed on file in the Australian Museum and
in the Queensland Museum.

Although nearby extralimital localities were
plotted on species’ distribution maps, only the
localities within the boundaries of the New
England region were included in the statistical
analyses.

A transparency of the State of New South
Wales, with the area of New England delineated
as defined in the present study, was produced at
the same scale as Swan’s (1990) maps. By
overlapping this transparency onto Swan’s maps
and comparing them, the species and localities
included in his study were detected and compared
with ours.

COVERAGE

The long duration of this survey allowed
opportunity for a comprehensive geographical
coverage of the area (Fig. 2). Collections were

made throughout the area with no large gaps,
although the collecting localities were slightly
less densely distributed in the west than
elsewhere.

Even with a relatively complete geographic
coverage, a distorted picture could he obtained if
important habitats were poorly represented. For
example, not collecting along streamsides in
New England would have failed to reveal the
presence of several species. We attempted to
cover all habitats. In order to assess the extent to
which that attempt was successful, the areas covered
by each category of the various environmental
parameters was calculated from the computer-
generated environmental maps, and the number
of collecting sites represented in each category
ascertained. For each parameter, the number of
collecting sites in a given category was roughly
proportional to the representation of that
category in the region. Thus, it appears that all
recognised habitats were adequately covered.

The seasonal coverage was not as good. Many
localities were visited only once. Consequently,
if certain species were active only outside that
period, they would have been missed.

OTHER RECORDS FOR NEW ENGLAND

Swan’s (1990) maps were based partly on
records not available to us and they add four
species (Hypsilurus spinipes , Tympanocryptis
diemenensis , Lerista punctatovittata , Crypto-
blepharus carnabyi) to the fauna of New
England. He reported Phyllurus cornutus from
the area, but Couper et al. (1993) restricted this
species to northeastern Queensland and allocated
the New England population to Saltuarius
swaini. Swan also reported Ramphotyphlops
broorni from New England but Shea (1995)
demonstrated the New England species to be R.
wiedii. 1'hus, we have excluded Phyllurus cornutus
and Ramphotyphlops broomi from the list of
species from New England.

The Sarah River Survey (Webber & Heatwole,
1991), an action plan for threatened Australian
reptiles (Cogger et al., 1993) and the Eastlink
Survey of 1 995- 1 997, conducted along a transect
through northern New England and southern
Queensland (Debus, unpubl. data) yielded no
species for the New England squamatan fauna
beyond those listed in Table 1 from other sources.
The Atlas of NSW Wildlife of (NSW National
Parks and Wildlife Service, database of the year
2000) summarised all records from New South
Wales, including New England, arising from a

NEW ENGLAND LIZARDS AND SNAKES

303

variety of sources and of varying reliability.
Almost all the records in the Atlas that were
based on voucher specimens had already been
included automatically in our database by virtue
of the specimens having been deposited in the
Australian Museum.

However, two new species records, Ctenotus
eurydice and Denisonia devisi, were based on
voucher specimens. The former is now regarded
as an invalid species (Sadleir, pers. comm.), but
the latter can be added to the known fauna of New
England.

The following species were recorded from New
England only on the basis of field identifications:
Phyllurus platurus , Egernia striata , Eulamprus
tympanum , Hemisphaeriodon gerarrdii , all with
a reliability index of 4 or 5 (with 1 being most
reliable, 6 least reliable). The last is known
extralimitally just east of New England and
further investigation may well extend its range
into the region. However, the first three species
almost certainly represent erroneous identifi-
cations and we do not consider them part of the
New England fauna; their known ranges are far
removed from New England (Cogger, 2000).

One of the species listed in our database,
Denisonia macula ta y may be in error. The record
is based on a single specimen from the Australian
Museum (AM R4765) that was later donated to
the M useum of Comparative Zoology at Harvard.
J. Rosado (pers. comm.) indicated that because of
renovations this specimen would be unavailable
for an identity check for at least another year.
Cogger (2000) showed the range of D. maculata
as coastal central Queensland, well beyond the
limits of New England. D. maculata is closely
related to D. devisi and the latter was once
considered a subspecies of the former (Cogger et
al.,1983). D. devisi is one of the species reliably
recorded from New England by the Atlas of NSW
Wildlife (Table 1 and see above) and it is likely
that the purported specimen of D. maculata is, in
fact, D. devisi.

Ctenotus eurydice : This species is now regarded
as an invalid species(R. Sadleir, pers. comm.),
since it cannot be reliably distinguished from C.
taeniolatus , and is therefore excluded from our
data base. The New England specimens of C.
eurydice were SPXEIOl 3 (NSW Wildlife Survey,
1988), SPXEI006 (Atlas of NSW Wildlife)
Neither of them can now be located.

Gehyra australis : There are three records of this
species from closely adjacent sites in the Moonbi
Ranges near Tamworth, NSW, (AM R29700,

AM R75990, VM D55544). They form a cluster
within a predominantly G. dubia population. The
Australian Museum specimens could not be
located, but that from the Victorian Museum was
initially identified as G. australis by the Acting
Curator of Herpetology, (D.J. Bray), and by one
of the authors (J.de B). The specimen was then
referred to the Australian Museum where it was
identified as G. dubia because of the presence of
an intemasal scale, a characteristic of that species
(R.Sadlier, pers. com.). Consequently the three
records of G australis were transferred to G.
dubia.

The reliably known squamatan fauna of New
England therefore consists of (a) our database of
106 species (14 species of geckos, 4 of pygo-
podids, 8 of dragons, 3 of goannas, 47 of skinks,
and 30 of snakes); (b) four additional species (2
dragons and 2 skinks) reported by Swan (1990);
and (c) an additional species based on voucher
specimens from the Atlas of NSW Wildlife
database, for a total of 1 1 1 species.

Several more are immediately extralimital and
further collecting may record their presence in
New England. Faunaf surveys involving parts of
New England are in progress (e.g., Nandewar
Ranges in western New England; Demon Nature
Reserve on the Timbarra Plateau in eastern New
England). The preliminary lists are subject to
revision and reassessment before they are
permissible for quotation; accordingly, it was not
possible to include information from them in the
present study.

THE REGIONAL FAUNA IN RELATION TO

THE HERPETOFAUNA OF AUSTRALIA

The majority of Australian squamates live in
tropical or arid habitats and are thus excluded
from New England (Cogger, 2000). Overall, 1 4%
of Australian species of squamates and 47% of
the genera occur in New England. At the specific
level, all Australian families of lizards are similar
in their proportional representation in New
England (12-15% of the total species from
Australia) (Table 2). By contrast, the different
ophidian families, have markedly divergent
proportional representations in New England at
the species level. The Typhlopidae and Boidae
have about the same proportional representation
(10-13%) as the saurian families, whereas the
colubrids and the elapids are unusually well
represented (Table 2).

Generic representation is higher than specific
representation. All Australian monitors belong to
Varanus and consequently New England has a

304

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1 . List of the lizards and snakes known from the New England Region and a summary of their types of
distributions. * recorded for the New England region by Swan (1990) but not present in our database; **
recorded in Atlas of NSW Wildlife database, based on a voucher specimen, but not in our database.

Key to symbols indicating distributional pattern within the New England region: U(e)=widspread and evenly
distributed; U(p)=widespread but patchily distributed; U(s)=widespread but sparsely distributed; W= mainly
distributed in the western part of the area; W(sw)=mainly in the southwestern sector; W(nw)=mainly in the
northwestern sector; E= distributed mainly in the eastern part of the area; E(ne)=mainly in the northeastern
sector; E(se)=mainly in the southeastern sector; S = mainly distributed in the southern part of the area; R=
recorded from less than ten localities in our database.

LIZARDS

FAMILY AGAMIDAE (Dragons)

Amphibolunis burnsi (Wells & Wellington, 1985): W(nw)
Amphibolurus muricatus (White, ex Shaw, 1790): U(e)
Amphibolunis nobbi Witten, 1972: U(e)

*Hypsilurus spinipes (Dumeril & Dumeril, 1851): E, R
Physignathus lesueurii (Gray, 1831): U(p)

Pogona barbata (Cuvier, 1829): W
Pogona vitticeps (Ahl, 1926): W, R
* Tympanocryptis diemenensis(Gray, 1841): S, R
Tympanocryptis lineata Peters, 1863: W, R
Tympanocryptis tetraporophora Lucas & Frost, 1895: W, R

FAMILY VARANIDAE (Monitors or Goannas)

Varanus gouldii (Gray, 1 838): U(s)

Varanus tristis (Schlegel, 1839): W(nw), R
Varanus varius (White, ex Shaw, 1 790): W, R

FAMILY GEKKONIDAE (Geckos)

Diplodactylus intermedius Ogilby, 1 892: W, R
Diplodactylus vittatus Gray, 1 832: W
Diplodactylus williamsi Kluge, 1963: W, R
Gehyra dubia (Macleay, 1 877): W
Gehyra variegata (Dumeril & Bibron, 1 836): W, R
Heteronotia binoei (Gray, 1845): W
Oedura lesueurii (Dumeril & Bibron, 1 836): W
Oedura monilis De Vis, 1 888: W, R
Oedura robusta Boulenger 1885: W
Oedura tryoni De Vis, 1884: W
Saltuarius swaini (Wells & Wellington, 1985): E
Saltuarius wyberba Cooper, Schneider & Covacevich, 1997:
E(ne), R

Underwoodisaurus milii (Bory de Saint- Vincent, 1 825): W
(Jnderwoodisaurus sphyrurus (Olgilby, 1 892); W

FAMILY PYGOPODIDAE (Snake-lizards)

Delma plebeia De Vis, 1888: W, R
Delma tincta Dc Vis 1888; W
Lialis burtonis Gray, 1835: U(p)

Pygopus lepidopodus (Lacepede, 1 804): U(s), R

FAMILY SCINCIDAE (Skinks)

Anomalopus leuckartii (Weinland, 1862): W
Anomalopus mackayi Greer & Cogger, 1985: W(nw), R
Anomalopus verreawdi (Dumeril & Dumeril, 1851): E, R
Bassiana platynota (Peters, 1 88 1 ): E
Calyptotis ruficauda Greer, 1983: E

Calyptotis scutirostnim (Peters, 1 873): E(nc)

Carlia tetradactyla (O’Shaughnessy, 1879): W
Carlia vivax (De Vis, 1884): U(p), R
Cautulazia (Ingram & Ehmann. 1981): E, R
Coeranoscincus reticulatus (GOnther, 1 873): E(ne), R
* Cryptoblepharus camabyi Storr 1976: R
Cryptoblephanis virgatus (Garman, 1901): U(p)

** Ctenotus eurydice Czechura and Wombcy, 1982: E(ne)
Ctenotus robustus Storr, 1970: U(e)

Ctenotus taeniolatus (White, ex Shaw, 1790): U(e)
Cyclodomorphus michaeli Wells & Wellington, 1984: R
Egemia cunninghami (Gray, 1832): U(e)

Egernia jrerei Gunther 1 897: E(nc), R

Egemia major (Gray, 1845): U(s), R

Egemia mepheei Wells & Wellington, 1984: E

Egemia modesta Storr, 1968; W

Egernia saxatilis Cogger, 1960: W, R

Egemia striolata (Peters, 1870): W

Egemia whitii (Lacepede, 1 804): E

Eremiascincus richardsonii (Gray, 1845): U(s), R

Eulampms heatwolei Wells & Wellington, 1984: E(se)

Eulamprus kosciuskoi (Kinghom, 1932): E(se)

Eulampms martini (Wells & Wellington, 1985): E(ne)
Eulampms murrayi (Boulenger, 1887): E
Eulampms quoyii (Dumeril & Bibron, 1 839): U(e)
Eulampms tenuis (Gray. 1831): U(s), R
Hemiergis decresiensis (Cuvier, 1829): U(p)

Lumpmpholis amicula Ingram & Rawlinson, 1981: E
Lampropholis caligula Ingram & Rawlinson, 1981: E(se), R
iMmpropholis delicata (De Vis, 1 888 ): E
Lampropholis elongata Greer, 1997: E(se), R
Lampropholis guichenoti (Dumeril & Bibron, 1 839): E
Lerista bougainvillii (Gray, 1 839): W
Lerista muelleri (Fischer, 1881): W(nw)

* Lerista punctatovittata (Gunther, 1867):W(sw), R
Lygisaums foliomm De Vis, 1884: W
Menetia greyii Gray, 1845: E(se), R
Morethia boulengeri (Ogilby, 1890): W
Nanoscincus maccoyi (Lucas & Frost. 1894): R
Ophioscincus tmneatus (Peters, 1876): E, R
Pseudemoia pagenstecheri (Lindholm, 1901): S
Saiphos equalis (Gray, 1 825): E
Sapmsc incus mustelinus ( O'Shaughnessy, 1874): E
Saproscincus rosei Wells & Wellington, 1985: E
Tiliqua scincoides (White, ex Shaw, 1 790): W

generic representation of 1 00%. There also is a gekkonids, pygopodids, agamids, boids and
high representation of Australian typhlopid colubrids (22-39%) (Table 2). Being inland, New
(50%), scincid (63%) and elapid (85%) genera in England lacks the predominantly marine families
New England but lesser generic representation by of snakes.

NEW ENGLAND LIZARDS AND SNAKES

305

TABLE 1 (Cont.).

SNAKES

FAMILY TYPHLOPIDAE (Blind Snakes or Worm Snakes)
Ramphotyphlops bituberculatus (Peters, 1863): W(sw), R
Ramphotyphlops nigrescens (Gray, 1845): U(e)
Ramphotyphlops proximus (Waite, 1893): W(sw), R
Ramphotyphlops wiedii (Peters, 1867): W

FAMILY BOIDAE (Pythons)

Antaresia maculosa (Peters, 1873): W, R
Morelia spilota (Lacepede, 1804): U(p)

FAMILY COLUBR1DAE

Boiga irregularis (Merrem, 1802): E, R

Dendrelaphis punctulata (Gray, 1 827): E(ne), R

FAMILY ELAP1DAE

Acanthophis antarcticus (Shaw & Nodder, 1 802): E, R
Austrelaps ramsayi (Krefft, 1864): E(se)

Cacophis harriettae Krefft, 1 869: E, R
Cacophis krefftii Gunther, 1 863): E, R

Cacophis squamulosus (Dumeril, Bibron & Dumeril 1 854): E
Demansia psammophis (Schlegel, 1 837): W
**Denisonia devisi Waite & Longman, 1920: W
Drysdalia coronoides (Gunther, 1 858): U(p)

Furina diadema (Schlegel, 1837): W
Hemiaspis signata (Jan, 1 859): E
Hoplocephalus bitorquatus (Jan. 1859): U(s)

Hoplocephalus stephensii Krefft, 1869: E(ne), R
Notechis scutatus (Peters, 1861): E
Pseudechis guttatus Dc Vis, 1905: W
Pseudechis porphyriacus (Shaw. 1794): E
Pseudonaja textilis (Dumeril, Bibron & Dumeril, 1854): W
Rhinoplocephalus nigrescens (Gunther, 1 862): E
Simoselaps australis (Krefft. 1 864): W
Sut a dwyeri (Worrell. 1956): U(e)

Sutasuta (Peters, 1863): W, R
Tropidechis carinatus (Krefft, 1863): E, R
Vermicella annulata (Gray, 1841): U(s)

DISTRIBUTIONAL PATTERNS

The distributions of all species (Figs 3-108)
can be grouped into several general categories.
The following interpretations have relied on our
database and maps, supplemented by the
additional localities on the maps of Swan (1990)
and from specimen-based entries in the database
of the Atlas of NSW Wildlife (NSW National
Parks and Wildlife Service, 2000).

TABLE 2. Representation in New England of the total
Australian Squamata. Total genera and species
calculated from Cogger (2000); number of genera
and species from New England from the present
database plus valid additions from Swan (1990).

Species & genera in New England (New England taxa as % of
Australian total)

Taxon

Genera

Species

Genera

Species

Lizards

Gckkonidac

18

111

6(33)

14(13)

Pygopodidae

8

34

3(38)

4(12) 1

Agamidae

13

65

5(39)

10(15)

Varanidae

1

26

1 (100)

3(12)

Scincidae

38

379

24 (63)

50(13)

Snakes

Typhlopidae

2

40

1(50)

4(10)

Boidac

6

15

2 (33)

2(13)

Acrochordidae

1

2

0(0)

0(0)

Colubridae

9

11

2(22)

2(18)

Elapidae

20

87

17(85)

22(26)

Hydrophiidae

12

31

0(0)

0i2)

Laticaudidae

1

2

0(0)

0(0)

Total

129

803

61 (47)

111 (14)

WIDESPREAD DISTRIBUTIONS. Widespread
species usually have broad tolerances and habitat
preferences. However, this is not always the case;
sometimes widespread distributions may be
patchy, reflecting discontinuous habitat within a
large area. For example, although Egernia
cunninghami is widespread (Fig. 46), it is largely
restricted to exfoliating granite outcrops. The
occurrences of Physignathus lesueurii and
Pseudechis porphyriacus are locally restricted by
the distribution of the rivers and other bodies of
water with which they are closely associated. In
other cases, especially in fossorial or secretive
species, patchiness may be more apparent than
real; a species like Pygopus lepidopodus may be
recorded only sparsely from an area merely
because it is difficult to find.

Twenty-two species (20% of the total squamatan
fauna) were widespread in New England. Of
these about equal numbers were evenly wide-
spread in the area (8 species), and widespread but
patchily (7 species) or sparsely (7 species)
distributed over the region (Table 1).

EASTERN VERSUS WESTERN DISTRIB-
UTIONS. There are gradients in climate and
natural vegetation in New England from east to
west; accordingly, 74% of the species have either
a predominantly western distribution, or a
predominantly eastern one. The Great Dividing
Range coincides with the boundary of the
geographic range of only a few species (e.g.,
Gehyra dubia , Eulamprus kosciuskoi) and in
some others, distributions have penetrated
slightly to the opposite side only in a few small
areas (e.g., Oedura lesueurii , Oedura tryoni ,

306

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 3. Amphibolurus bumsi

FIG 6. Physignathus lesueurii

FIG 9. Tympanocryptis lineata

FIG 4. Amphibolurus muricatus

• V

• J

• • J

Vv *1

V*.

• N

• V

/ •

•

• 1^^

1

FIG 7. Pogona barbata

FIG 10. Tympanocryptis
tetraporophora

FIG 5. Amphibolurus nobbi

y ,A \

FIG 8. Pogona vitticeps

• L

Ji

•

• > ^

-1

i

r

FIG 1 1 . Varanus gouldii

NEW ENGLAND LIZARDS AND SNAKES

307

FIG 18. Gehyra variegata

FIG 19. Heteronotia binoei

FIG 20. Oedura lesueurii

308

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 21 . Oedura monilis

FIG 22. Oedura robust a

FIG 23. Oedura try>oni

FIG 24. Saltuarius sxvaini

FIG 25. Saltuarius wyberba

FIG. 26. Undenx’oodisaurus rnilii

FIG 27. Underwoodisaurus sphyrurus

FIG 28. Delma plebeia

FIG 29. Delma tincta

NEW ENGLAND LIZARDS AND SNAKES

309

FIG 33. Anomalopus mackayi

310

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 39. Carlia vivax

FIG 40. Cautula zia

FIG 41. Coeranoscineus reticulatus

FIG 42. Cryptoblepharus virgatus FIG 43. Ctenotus robustus

FIG 44. Ctenotus taeniolatus

FIG 45. Cyclodomorphus michaeli FIG 46. Egernia cunninghami

FIG 47. Egernia frerei

NEW ENGLAND LIZARDS AND SNAKES

311

— - -1 j • t.

j

r

.

i •

«•

1 —

~r~^| — 1-
* S — 7

r \

» K

*• .•

y

^ i

- J

y b

•

•

• — ■

*

..-1— .jft

-j*

\.l |

1 \

1 \

■ f.

— v — \ #

• ^

l

J —

\ J

•

•i\ I
\?\

-

FIG 48. Egemia major FIG 49. Egernia mcpheei FIG 50. Egernia modesta

FIG. 5 1 . Egernia saxatilis

FIG 52. Egernia striolata

FIG 53. Egernia whitii

FIG 54. Eremiascincus richardsoni

FIG 55. Eulamprus heaiwolei

FIG. 56. Eulamprus kosciuskoi

312

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 57. Eulamprus martini

FIG 58. Eulamprus murrayi

FIG. 59. Eulamprus quoyii

FIG 60. Eulamprus tenuis

FIG 61 . Hemiergis decresiensis

FIG. 62. Lampropholis amicula

FIG 63. Lampropholis caligula

FIG 64. Lampropholis delicata

FIG 65. Lampropholis elongata

NEW ENGLAND LIZARDS AND SNAKES

313

FIG 66. Lampropholis guichenoti

FIG 67. Lerista bougainvillii

FIG 68. Lerista muelleri

FIG 69. Lygisaurus foliorum

FIG 70. Menetia greyii

FIG 7 1 . Morethia boulengeri

FIG 73. Ophioscincus truncatus

FIG 74. Pseudemoia pagenstecheri

FIG 72. Nannoscincus maccoyi

314

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 78. Tiliqua scincoides

FIG 79. Ramphotyphlops
bituberculatus

FIG. 80. Ramphotyphlops nigrescens

FIG 81. Ramphotyphlops proximus

Ji

£

, , ,

\ •

-j

• L

=

•

•

• 1 /

FIG 82. Ramphotyphlops wiedii FIG 83. Antaresia maculosa

NEW ENGLAND LIZARDS AND SNAKES

315

FIG. 84. Morelia spilota

FIG 87. Acanthophis antarcticus

FIG. 85. Boiga irregularis

\ J

X

,A i

f •

l

• 1

\ 1 •

-

•

FIG 88. Austrelaps ramsayi

FIG 86. Dendrelaphis punctulata

FIG 89. Cacophis harriettae

FIG. 90. Cacophis krefftii

FIG 91. Cacophis squamulosus FIG 92. Demansia psammophis

316

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 93. Denisonia devisi

FIG 94. Drysdalia coronoides

FIG 95. Furina diademu

FIG 96. Hemiaspis signata

FIG 97. Hoplocephalus bitorquatus FIG 98. Hoplocephalus stephensii

FIG 99. Notechis scutatus

FIG 100. Pseudechis guttatus

FIG. 101. Pseudechis porphyriacus

NEW ENGLAND LIZARDS AND SNAKES

317

FIG 102 . Pseudonaja textilis

FIG 103. Rhinoplocephalus nigrescens FIG 104. Simoselaps australis

FIG 105. Sut a dwyeri

FIG. 106. Suta suta

FIG. 107. Tropidechis carinatus

FIG. 108. Vermicella annulata

318

MEMOIRS OF THE QUEENSLAND MUSEUM

Egernia modest a, Hemiaspis signata). However,
the topography is not severe on the continental
divide and it is unlikely to constitute a physical
barrier to dispersal for many species. For a number
of species, the boundary of the geographic range
lies considerably to one side or the other of the
continental divide (e.g., Anomalopus leuckartii ,
Bassiana platynota , Lampropholis delicata , Lam-
propholis guichenoti , Pseudechis porphvriacus).
In such cases, it is likely that distribution is
limited by one or more climatic parameters with
an east-west gradient (see below), rather than by
the Great Dividing Range itself.

Some of the eastern species certainly can
disperse beyond the Great Dividing Range as
there are outlier populations on the western edge
of the study area at Mt Kaputar, where elevation,
rainfall and vegetation parallel those on the eastern
side, but differ from those of the western areas
surrounding the mountain. Such outliers are:
Bassiana platynota , Ctenotus taeniolatus ,
Egemia cunninghami , Egernia whitii, Eulamprus
heatwolei , Hemiergis decresiensis , Lampropholis
delicata , Pseudemoia pagenstecheri and
Rhinoplocephalus nigrescens. Conversely, a few
predominantly western species (e.g, Heteronotia
hinoei) are found in a few localities in the east.

A number of the species that are predominantly
western within the confines of New England are
absent or uncommon in eastern New England
because of their exclusion from the wetter forests
of that region; beyond that gap their ranges do
extend eastward, often to the coast. Examples are
Oedura robusta. Lygisaurus folio rum, Tiliqua
scincoides , Ramphotyphlops wiedii , Demansia
psammophis , Furina diadema , Pseudonaja
textilis and Simoselaps australis.

There are slightly more species with a pre-
dominantly western distribution (40%) than a
predominantly eastern one (30%). The difference
may be even more disparate than these figures
suggest, as there are a number of western species,
e.g., Trachydosaunis rugosus , Pseudechis australis ,
that have been found near, but not inside, the
western boundary of New England. With further
collecting, or upon completion of some of the
surveys still in progress, the ranges of some of
these will likely be found to extend into New
England.

It should also be noted that gorges and river
valleys permit the penetration of coastal species
into New England e.g. Boiga irregularis (Webber
& Heatole, 1991).

Different taxa vary in regard to east-west
distributional patterns. The geckos (13 species
western, 2 species eastern), pygopodids (2 w, 0
e), varanids (2 w, 0 e), agamids (5 w, 1 e),
typhlophids (3 w, 0 e) and boids (1 w, 0 e) are
mainly western rather than eastern, whereas the
reverse is true of the skinks (24 species eastern,
12 species western), elapids (11 e, 7 w) and
colubrids (2 e, 0 w).

NORTHERN VERSUS SOUTHERN
DISTRIBUTIONS. The north-south axis does
not feature prominently in the distribution of
squamates in New England. The northerly
distributed species occur on only one side or
other of the Great Dividing Range and hence
represent subsets of the eastern (8 species) or
western (4 species) groups, but whose southern
limits of distribution are reached in New
England. There are no species restricted to the
north that range widely from east to west in that
region.

There are just two species in New England that
span New England from east to west but
primarily in the south; these are Pseudemoia
pagenstecheri (Fig. 74) and Tympanocryptis
diemensis (Swan, 1990). There are six species
from only the southeastern quadrant and four
from only the southwestern one; these were
considered subgroups of the eastern and western
groups.

‘CENTRAL’ DISTRIBUTION. There was no
species w ith a strictly central distribution. How-
ever, Hemiergis decresiensis was predominantly
central in that it was represented by many
localities in the central region, along the con-
tinental divide and immediately adjacent to it;
beyond that area it was widespread (and was
listed as such for statistical purposes), but only at
scattered localities.

The only other species known only from
central locations were represented by so few
records that their distribution within New
England could not be interpreted.

INFREQUENTLY ENCOUNTERED. A salient
aspect of the New England reptilian fauna is the
large number of species that were recorded from
only a small number of localities (‘R’ in Table 1 ).
In many cases, supplementation by the additional
localities mapped by Swan (1990) allowed
allocation of these species to one of the
distributional categories mentioned above. The
remaining (4 species) were represented by only
one locality in New England and supplementary

NEW ENGLAND LIZARDS AND SNAKES

319

information was not available to aid in
interpretation of their local distributional pattern.
For example, Nannoscincus maccoyi (Fig. 72)
was found at only one locality, southeast of the
center of New England. This is a considerable
range extension as formerly it was not known
north of Sydney (Swan, 1 990). It is likely that the
ranges of such species will be extended with
further information, or these localities shown to
be outliers.

Some of these R species may be rare; in other
cases they may be more common than suggested
by their frequency in collections because they are
secretive and therefore less subject to being
found. Gibbons et al. (1997) noted that over a
long period of collecting, the known extent of
distributions tend to increase remarkably and
species’ ranges often are much broader than
indicated by less intensive collecting.

RELATION OF DISTRIBUTION TO

ENVIRONMENTAL CONDITIONS

In the following account, the range of
conditions under which the various species are
found within New England is given. Outside of
New England some of these species may extend
into areas with environments beyond the limits
encountered within the study area. Where
correlations are made between distributional
patterns and particular levels of environmental
features, the relationship is referable only to the
New England area, not to the geographic range of
a species as a whole.

ELEVATION. The lowest elevations (below
400m) are located at the edges of the region in the
northwest, southwest and southeast. These slope
upward to the New England plateau of 800- 1 200m
in the eastern half of the region with isolated
areas up to more than 1400 m, especially near the
southern boundary and from the east-central
region to the eastern border (Fig. 109).

The frequency of collecting localities had a
unimodal distribution skewed toward the middle
elevations. The greatest frequency of occurrence
was at 1 00- 1 1 00m and the second greatest at
900- 1000m. From these values there was a
progressive decrease toward the extremes of the
range of elevations.

All 26 species for which there were sufficient
data for statistical testing of distributions against
category of elevation were found at a wide
variety of elevations. Of these, seven were
distributed randomly with respect to elevation,
seven were over-represented at the lower

FIG. 109. Elevations (m) of the study area. Data from
the original Digital Elevation Model (DEM) supplied
by NSW Land and Property Information (NSW LPI),
1997.

elevations, two at the middle elevations, three at
middle to high elevations and seven at the higher
elevations (Table 3).

CLIMATE. Rainfall. A map of New England
with a rough approximation ot the isohyets was
presented in the first paper of this series (Heat-
wole & Simpson, 1986). A more detailed map
showing the distribution ot raintall was used tor
the present analyses (Fig. 1 10). The western and
northwestern parts of the region have relatively
low mean annual raintall (mostly 800mm or less)
increasing toward the east. The eastern border is
especially wet with rainfall > 1700mm per annum
occurring just to the east and southeast ot the
designated boundary.

The collecting sites were distributed um-
modally with respect to rainfall categories, but
skewed toward the lower levels of precipitation.
Almost a third of the sites were in areas
characterised by 700-800 mm of rain per annum,
and the next highest frequency was in the 800-
900mm category. All other categories individually
contained <13% of the collecting sites.

Of the 14 species for which there were
sufficient data for statistical testing, all occurred
over most of the spectrum of rainfall values and
two species were randomly distributed with

320

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3. Distribution of species or reptiles in relation to various environmental parameters. R=random; —
indicates that expectation was lower than 5 in the smallest category and therefore not testable statistically. *
‘Other’ refers to minor soil or rock types lumped into one category for statistical testing; CP=Chocolate Prairie;
RP=Red Podzolic; YS/YP=Yellow Sol/Yellow Pod; YP/GP=Yellow Pod/Glcy Pod.

Species

Elevation

Rainfall

Coldest

Month

Minimum

Temp

Hottest

Month

Maximum

Temp

Vegetation

Soil Type

Rock Type

Heteronotia binoei

Lower

—

—

26°-32°

—

—

R

Oedura lesueurii

Middle

Low to
Medium

-4° to 1°

25°-28°

—

YS/YP;YP.G

P

Granite

! Oedura tryoni

Lower

Low to
Medium

-4° to 1°

26°-32°

—

YS/YP;YP/G

P

Granite;

Other*

Amphibolurus nohbi

Lower

Low to
Medium

R

26°-32°

—

YP/GP;

Other*

R

Amphiboturus muricatus

R

R

-4° to -1°

R

—

R

R

Anomalopus leuckartii

Lower

Low

R

26°-32°

—

Other*

Granite,

Other*

Bassiana platvnota

Higher

—

—

—

—

—

Calvptotis scutirostrum

R

—

—

—

—

—

—

Ctenotus robustus

Lower

Low

—

26°-32°

—

R

Ctenotus taeniolatus

Higher

—

—

R

—

—

—

Egemia cunningham i

R

R

R

R

—

R

Granite

Egemia striolata

Lower

—

—

26°-32°

—

—

R

Egemia whim

Higher

—

—

—

—

—

Eulamprus murrayi

Middle

—

—

—

—

-

Eulamprus quoyii

R

Medium to
High

0°

R

—

—

R

Hemiergis decresiensis

Higher

Low

-4° to -1°

24°

—

RP; YS/YP;
Other*

Basic Igneus

Iximpropholis delicata

Middle to
High

Medium to
High

0° to 6°

21-25°

Woodland;
Dry Forest;
Wet Forest

RP

R

Lampropholis guichenoti

Middle to
High

Medium to
High

0°

21°-25°

Wet Forest

RP

R

Morethia boulengeri

Lower

Low to
Medium

—

R

—

—

R

Saiphos equal is

Middle to
High

Medium to
High

0°

21°-25°

Wet Forest

RP

Sedimentary

Saproscincus rosei

R

—

—

—

—

R

Ramphotyphlops nigrescens

Higher

—

—

24°-25°

—

—

Basic Igneus;
Granite;
Other*

: Austrelaps ramsavi

Higher

—

—

—

—

—

— .

A us t re laps superbus

—

—

—

—

—

Basic Igneus

Pseudechis porphyriacus

Higher

Low to
Medium

-1°

21°-24°

—

YS/YP; CP;
Other*

Sedimentary;
Basic Igneus

Pseudonaja textilis

R

—

—

R

—

—

R

Suta dwyeri

R

—

R

—

—

R

respect to rainfall (Table 3). However, three
species were over-represented in areas of low
rainfall, five species were found more often than
expected on the basis of chance in localities of
low to medium rainfall, and four species were
proportionately more common in places with
medium to high rainfall.

Temperature. Heatwole & Simpson (1986)
presented maps (based on Lea et al., 1977 ‘An

Atlas of New England’) of the isotherms of the
mean daily maxima and minima for the months of
July and January, and Heatwole et al. (1995)
reproduced one of them (mean daily minimum
for July). In general, the central part of the region
is the coldest with higher isotherms concentric-
ally arranged toward the periphery.

The present analysis was based on the mean
daily minimum of the coldest month (range -4 to

NEW ENGLAND LIZARDS AND SNAKES

321

FIG 1 10. Mean annual rainfall (mm) of the study area.
Original DEM supplied by NSW LPI. Mean potential
rainfall data derived from ESOCLIM program which
was supplied by CRES at National University , 1 997.

6°C) (Fig. Ill) and mean daily maximum of the
hottest month (range within the area of 12°C to
32°C) (Fig. 112).

The mean daily minima of the coldest month
were unimodally distributed with a peak at and
just below freezing, gradually descending toward
the higher and lower values. Intervals of mean
daily minima of 2°C or above accounted for only
slightly more than 9% of the collecting sites and
those of 2°C or above for only about 14%.

Of the twelve species for which statistical
analysis was feasible, three showed random
distribution with regard to levels of mean daily
minimum temperature of the coldest month
(Table 3). The remainder, while found over most
of the spectrum of minimum temperatures,
statistically favoured certain levels. Four species
tended to occur more often than expected in the
coldest localities, four at intermediate localities
(near zero), and one was enigmatic, being found
more often than expected by chance over a wide
range of the higher temperatures.

The mean daily maximum temperature of the
hottest month showed a bimodal frequency
distribution with one peak at 24°C (20.6% of the
collecting sites) and 25°C (2 1 .4%) and the other

at 28°C (12.1%); all other temperature intervals
individually accounted for <10% of the sites.

Of the 20 species for which statistical testing
could be conducted, seven were shown to be
distributed at random with respect to mean daily
maximum temperature for the hottest month
(Table 3). Four species were over-represented in
the thermal categories in the range of 2 1 °-24°C or
2 1 °-25°C, two in the ranges of 24°C or 24°-25°C,
and seven in various ranges of categories from
25° C and higher.

VEGETATION. The New England area is
devoted to the pastoral industry and consequently
is highly modified by human activity. Accord-
ingly. many of the collecting sites occurred on
cleared land (45%), with disturbed remnants
accounting for an additional 11% of the localities.
The forest types containing the greatest frequency
of occurrence of collecting sites were Dry Forest
Complex (19%) and Dry Open Forest (15%); all
other vegetation types individually contained
1 1% of the sites (Fig. 113).

Each of the previous papers in this series has
used a different classification of vegetation or
land-use. The present analysis is based on a
modification of the classification ot vegetation
systems for northeastern New South Wales (NSW
National Parks and Wildlife Service, 1994).

Only three species could be tested because of the
small areas occupied by wet forests and the statistical
limitations this imposed. Saiphos equal is and
Lampropholis guichenoti were found in wet
forest significantly more often than expected at
random, whereas Lampropholis delicata was
over-represented in woodlands and in both dry
and wet forests (Table 3).

SOIL TYPE. The soil types (Fig. 1 14) accounting
for the greatest number of collecting localities
were Yellow Pod/Grey Pod (24% of the sites).
Red Podzolic (21%) and Yellow Sol/Yellow Pod
(19%); the other 15 soil types each accounted for
<10% of the collecting localities and collectively
for 36%.

Eleven species were tested against the
distribution of soil types. Of these, two were
distributed randomly with respect to soil type
(Table 3), three were found oftener than expected
by chance on red podzolic soils, and live were
over-represented on more than one soil type, but
always involving yellow sol/yellow pod and/or
yellow pod/gley pod soils in combination with
others.

322

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 111. Mean daily minimum temperatures(°C) of
the coldest month (July) in the study area. Original
DEM supplied by NSW LPI. Mean potential temp-
erature data derived from ESOCLIM program which
was supplied by CRES at National University, 1 997.

ROCK TYPE. The two most abundant rock types
characterising collecting sites were sedimentary
rocks with low quartz (36% of the collecting
sites) and granite (23%); all other rock types
accounted for 13% or less of the collecting
localities (Fig. 115).

Twenty-three species allowed for statistical
testing of distribution against the various rock
types, of which 12 were randomly distributed,
five were over-represented on granite, four on
basic igneous, and two on sedimentary rocks
(Table 3).

Only three species could be tested against all
environmental characteristics (Table 3). Lampro-
pholis delicata showed random distribution only
with respect to parent rock type and consequently
all other characteristics may have affected its
distributional patterns. It is over-represented at
middle to high elevations, in woodlands and forests
on red podzolic soils where there is medium to
high rainfall and where mean minimum
temperature of the coldest month is zero°C or
above and mean maximum temperature of the
hottest month is in the range of 21°-25°C. L.
delicata does occur under other conditions, but is
under-represented there and thus it would appear

FIG 1 1 2. Mean daily temperatures (°C) of the warmest
month (January) in the study area. Original DEM
supplied by NSW LPI. Mean potential temperature
derived from ESOCLIM program at National
University, 1997.

that this suite of environmental conditions
defines the optimum habitat of this species.

Lampropholis guichenoti had an almost
identical distribution to L. delicata. It differed
only by being over-represented in a narrower
range of minimum temperatures of the coldest
month (0°C only, rather than the larger range
0°-6°C), in only wet forest rather than multiple
kinds of forest, and by being randomly dis-
tributed relative to soil type. Despite their great
similarity, and their great overlap in distribution,
these two species seem to have minor differences
in the biographical details.

Saiphos equalis is distributed very similarly to
L. guichenoti except that is is over-represented on
sedimentary rocks instead of being randomly
distributed with respect to rock type.

Eight species could be tested against all
environmental characteristics except vegetation
(the characteristic allowing for the testing of the
fewest number of species) (Table 3).

The most clear-cut result was for Egernia
cunninghami, which was distributed randomly
with respect to all environmental characteristics
measured except it was over-represented on granite.
This species inhabits crevices in exfoliating

NEW ENGLAND LIZARDS AND SNAKES

323

JB Rainforest
B3 Moist Oper
I I Dry Forest ..

■Motet Open Forest
| | Dry Open Forest/Woodland

H Plantation

Unmapped

g black earth -eucbrozem
3 black earth-prairie
] choc-black earth
1 chocolate-prairie
3 krasznozem-chocolate
3 non-calclc brown
| red brown earth
i] red podzolic

no data

solod sol yellow earth
solodlsed solonelz red brown earth
solonetzic red brown earth
yellow pod gley pod
yellow podzolic
yellow sol yellow pod

sol

FIG 113. Vegetation systems of the study area.
Derived from broad vegetation boundaries mapped
on 1 : 100k Landsat imagery. Mapping undertaken by
NPWS in 1991.

rocks throughout its range (Cogger, 2000);
granite is especially prone to such structure. It is
not surprising that the distribution of E.
cunnighami is strongly related to the occurrence
of this primary feature of its habitat.

Similarly, Amphibolurus muricatus that
showed random distribution with respect to all
environmental parameters tested except for mean
minimum of the coldest month; this species was
over-represented in the coldest part of the area
(Table 3). The remaining six species in this
category departed from randomness in respect to
a number of parameters and distributional
patterns may be influenced in more complex
ways than for A. muricatus and E. cunninghami.

For the remaining species tests there were three
or more environmental parameters for which
tests could not be conducted and consequently
optimal habitats are defined less completely.
Heteronotia binoei and Egernia striolata are
distributed randomly with respect to rock type,
Suta dwyeri and Pseudonaja textilis to elevation,
maximum temperatures and rock type,

FIG 114. Distribution of the various soil types in the
study area. From the Atlas of New England (Lea et al.,
1997), digitised and executed using Environmental
Research Mapping Scheme (Ferrier, 1989), NPWS,
NSW, Armidale office.

Saproscincus rosei to elevation and rock type,
Calyptotis scutirostrum to elevation, Ctenotus
taeniolatus to maximum temperatures, and
Morethia boulengeri to maximum temperatures
and rock type. Some of the species with a limited
number of available tests were over-represented
on some categories. Heteronotia binoei was
over-represented at lower elevations and high
maximum temperatures, Bassiana platynota at
high elevations, Ctenotus robustus at low rainfall
(see above), low elevation, and high maximum
temperatures, Ctenotus taeniolatus to high
elevations, Egernia striolata at low elevations,
Egernia whitii at high elevations, Eulamprus
murrayi at middle elevations, Morethia
boulengen at low elevations and low to medium
rainfall, Ramphotyphlops nigrescens at high
elevations and maximum temperatures in
midrange, and Aust relaps ramsayi at high
elevations (Table 3).

Various of the environmental attributes of the
region are correlated with each other (e.g.,
temperature, elevation, and rainfall) and hence

324

MEMOIRS OF THE QUEENSLAND MUSEUM

| j Sedimentary rocks<high % quartz) Quaternary sand and alluvium

1 Sedimentary rocks(low % quartz) H.;j Serpentine
B Acid volcamcs EZH Basic igneous rocks

■I Granitic rocks Hi Leuco granitic rocks

— '

FIG 115. Distribution of the various rock types in the
study area. Derived from the Department of Minerals,
geology 1 :250k Mapping. Digitised and classified by
NSW NPWS 1993.

the precise factor or combination of factors that
proximally affects distribution cannot be
identified individually from the present data.
However, assembling the suite of environmental
levels at which a species shows over-
representation, permits definition of its optimal
habitat, and allows construction of testable
hypotheses about proximal factors affecting
distributions. Detection of features with which a
species is randomly distributed is perhaps of even
jnore importance as it identifies environmental
parameters that are unlikely to be affecting a
species distribution, at least within the range of
values encountered in New England.

ACKNOWLEDGEMENTS

We are grateful to a host of research assistants,
students, colleagues, relatives and friends for
helping collect specimens; to the curatorial staff
of the Herpetology Divisions of the Australian
Museum, the Queensland Museum and the State
Museum of Victoria for permission to examine
specimens and for facilitating our work in
numerous ways; to Simon Ferrier, Harry Hines,
Alan Jones, Jamie Love, Craig Lawlor, Nick

Rollings, David Pauli, Bob Harden, Tracy
Maddocks, Stuart Cairns, Mike Roach, Russell
Hobbs, Maree Koch, Sandy Hamdorf and Louise
Percival variously for assistance with treatment
of data, literature searches, computer techniques,
preparation of maps, and typing.

The project was carried out under a succession
of permits from the National Parks and Wildlife
Service of New South Wales and with the
approval of the Animal Welfare Committee of the
University of New England. The research was
financed by the Internal Research Funds of the
University of New England and privately by the
authors.

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COGGER, H.G 2000. Reptiles and amphibians of
Australia. Sixth Ed. (Reed New Holland:
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COGGER, H.G, CAMERON, E.E. & COGGER, H.M.
1983. Amphibia and reptilia. Zoological
catalogue of Australia. Vol. 1. (Bureau of Flora
and Fauna: Canberra).

COUPER, P.J., COVACEVICH, J.A. & MORITZ, C.
1994. Designation of the type species of
Saltuarius , and other data on the genus. Memoirs
of the Queensland Museum 35: 26.

COUPER, P. J., SCHNEIDER, C.J. & COVACEVICH,
J.A. 1 997. A new species of Saltuarius (Lacertilia:
Gckkonidae) from granite based, open forests of
eastern Australia. Memoirs of the Queensland
Museum 42( 1 ): 91 -96.

FERRIER, S. 1989. Environmental Research Mapping
System (ERMS). User’s Manual for Version 2.
National Parks and Wildlife Service of New South
Wales,

FORD. H.A. & McFARLAND, D. 1991 . Faunal survey
of New England. III. Birds. Memoirs of the
Queensland Museum 30(3): 38 1-431.

GIBBONS, J.W., BURKE, V.H., LOVICH, J.E.,
SEMLITSCH, R.D.. TUBERVILLE, D., BODIE,
J.R., GREENE, J.L., NIEWIAROWSKI, P.H.,
WHITEMAN. H.H., SCOTT, D.E.,
PECHMANN, J.H.K., HARRISON, C.R.,
BENNETT, S.H., KRENZ, J.D., MILLS, M.S.,
BUHLMANN, K.A., LEE, J.R.. SEICiEL, R.A.,
TUCKER, A.D., MILLS, T.M., LAMB, T.,
DORCAS, M.E., CONGDON, J.D., SMITH,
M.H., NELSON, D.H., DIETSCH, M.B.,
HANLIN, H.G, OTT, J.A. & KARAPATAKIS,
D.J. 1997. Perceptions of species abundance,
distribution, and diversity: lessons from four
decades of sampling on a government-managed
reserve. Environmental Management 21:
259-268.

GREER, A.E. 1992. Revision of the species previously
associated with the Australian scincid lizard
Eulampris tenuis. Records of the Australian
Museum 44: 7-19.

NEW ENGLAND LIZARDS AND SNAKES

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1997. A new species of Lampropholis (Squamata :
Scincidae) with a restricted high altitude
distribution in eastern Australia. Australian
Zoologist 30(3): 360-368.

HEATWOLE, H. & SIMPSON, R.D. 1986. Faunal
survey of New England. I. Introduction and
general description of the area. Memoirs of the
Queensland Museum 22: 107-1 13.

HEATWOLE, H., de BAVAY, J M WEBBER, P. &
WEBB, G 1995. Faunal survey ofNew r England
IV. The Frogs. Memoirs of the Queensland
Museum 38: 229-249.

HUTCHINSON, M.F. 1989. A new objective method
for spatial interpolation of meteorological
variables from irregular networks applied to the
estimation of monthly mean solar radiation,
temperature, precipitation and windrun. CSIRO
Division of Water Resources Technical
Memorandum 5: 95-104

HUTCHISON. M.N. 1990. The generic classification
of the Australian elapid snakes. Memoirs of the
Queensland Museum 29: 397-405.

HUTCHISON, M.N. & DONNELLAN, S.C. 1992
Taxonomy and genetic variation in the Australian
lizards of the genus Pseudemoia (Scincidae:
Lygosominae). Journal of Natural History 26:
215-264.

HUTCHISON, M.N., DONNELLAN, S.C, BAVER-
STOCK, P.R., KRIEG, M., SIMMS, S. &
BURG IN, S. 1990. Immunological relationships
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535-554.

INGRAM, G & COVACEVICH, J. 1988. Revision of
the genus Lygisaurus dc Vis (Scincidae: Reptilia)
in Australia. Memoirs of the Queensland Museum
25(2): 333-354.

KING M. 1983. The Gehyra australis species complex
(Sauria: Geckkonidae). Amphibia-Rcptilia 4:
147-169.

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NSW NATIONAL PARKS AND WILDLIFE
SERVICE, 1994. Vegetation systems of

north-east NSW forests. North East Forests
Biodiversity Report No. 26: 1-19.

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print-out of database.

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of the Australian Tiger snakes (Notechis) and
Copper Heads (Austrelaps) (Serpentes: Elapidae).
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103(2): 125 -135.

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Nannoscincus Gunther: a re-evaluation. Memoirs
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Taxonomy and distribution of the scincid lizard
Saprosdncus challengeri and related species in
southeastern Australia. Memoirs ol the
Queensland Museum 34:138-158.

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R a m phot yph l ops broomi (Squamata.

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1 995b. A taxonomic revision of the Cyclodomorphus
casuarinae complex (Squamata: Scincidae).
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SIMPSON R D. & STANISIC. J. 1986. Faunal survey
of New England. II. The distribution ot gastropod
molluscs. Memoirs of the Queensland Museum

22(2): 115-139.

>MITH WJ S., OSBORNE, W.S., DONNELLAN,
SC & COOPER, P.D. 1999. The systematic
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(Reptilia: Agatnidae), in south eastern Australia.
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reptiles, amphibians and mammals of the Sara

ELlSTwI WELLINGTON, C R. .985. A
classification of the Amphibia and reptiles of

Suppl. series (1): 1-65.

ETNABATRACHUS MAXIMUS GEN. ET SP. NOV., A PLIO-PLEISTOCENE FROG
FROM MOUNT ETNA, CENTRAL EASTERN QUEENSLAND

SCOTT A. HOCKNULL

Hocknull, S.A. 2003 06 30: Etnabatrachus maximus gen. et sp. nov., a Plio-Pleistocene frog
from Mount Etna, central eastern Queensland. Memoirs of the Queensland Museum 49 ( 1 ):
327-330. Brisbane. ISSN 0079-8835.

Etnabatrachus maximus gen. et sp. nov. is the First record of fossil frogs from the Mount Etna
region of central eastern Queensland. E. maximus ’ very large size, divided prominences and
lateral mid-shaft ridge distinguish it from all other Australian genera. The associated fauna
from the type locality indicates a Plio-Pleistocene rainforest environment, making E.
maximus the largest known species of rainforest frog from Australia. □ Etnabatrachus
maximus, Plio-Pleistocene, Australia, Ilium, Cainozoic.

Scott A. Hocknull, Queensland Museum, PO Box 3300, South Brisbane, 4101 (e-mail:
scotth@qm.qld.gov.au); 20 May 2003.

The first fossil frogs from Australia were
described by Tyler in 1974 (Tyler, 1974). Since
then, Tyler has published on taxa from the
Eocene of southern Queensland through to the
late Holocene deposits of South and Western
Australia (Tyler, 1999). The dominating faunas
exhibiting extinct taxa have been the Oligo-
Miocene of Riversleigh Qld, Bullock Creek NT
and Lake Palankarrina SA. The Pliocene to
Holocene record is marked by the exclusively
extant taxa illustrating an early establishment of
the modem frog faunas.

Several new local faunas have been excavated
from deposits from the Mount Etna Caves,
Rockhampton, CE Queensland. These local
faunas span the Pliocene to Holocene and
preserve abundant frog remains, including
species of Lechriodus , Kyar annus, Litoria and
Limnodynastes. The age of the type locality for
Etnabatrachus maximus gen. et sp. nov. is
interpreted as late Pliocene to early Pleistocene
based on taxa occurring across this boundary,
including: Petauroides sp., close to P. stirtoni
(Turnbull & Lundelius, 1970), Pseudokoala sp.,
close to P erlita (Turnbull & Lundelius, 1970),
Pa torches tes sp., close to P. parvus (Tedford,
1994), Perameles sp., close to P. bowensis
(Muirhead et ah, 1 997) and Kurrabi sp. (Flannery
et al., 1992).

The type locality for E. maximus gen. et sp.
nov. is interpreted as rainforest on the basis of
sediments being at a time of major speleothem
development, and taxa with distinct rainforest
affinities, including: pseudocheirids -
Pseudochit-ulus mayeri , Pseudocheirops spp.
petaurids; Dactylopsila spp.; macropodids -
Dendrolagus sp.; peramelimorphians -

peroryctid spp.; murids - Uromys sp. and
Pogonomys sp. nov. (H. Godthelp, pers.comm.);
and anurans -Lechriodus sp. and Kyarannus sp.

Specimens described herein are deposited in
the Queensland Museum fossil collection
(QMF). Ilial terminology and nomenclature
follow Tyler (1976) and taxonomy follows
Cogger (2000).

SYSTEMATICS

Family 7HYLIDAE Gray, 1825

Etnabatrachus gen. nov.

TYPE SPECIES. Etnabatrachus maximus sp. nov. from
die Plio-Pleistocene of Mount Etna and Limestone Ridge,
localities QML 1385 (Type) and QML1284.

ETYMOLOGY. Etna- in reference to Mount Etna, type
locality and -batrachus for frog.

DIAGNOSIS. Large, robust ilium. Ilial crest
absent, dorsal acetabular expansion poorly
developed, rising slightly above dorsal margin of
ilial shaft. Preactabular zone broad and divided
laterally by a mid-shaft ridge. Ridge runs anterior
to the acetabular rim, orientated anterodorsally
approximately halfway along its length. Ridge
terminates at ventral margin of shaft, at
approximately anterior Va of ilium length. Dorsal
prominence divided into an anterior and a
posterior prominence, bisected by a shallow
valley running from the anterio-medial side to the
postero-lateral side. Anterior-most prominence
laterally directed with elongate-ovoid
protuberance in lateral profile. Anterior-most
prominence placed well anterior of anterior
acetabular rim. Posterior-most prominence
smaller than anterior prominence with a more

328

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 1. Etnabatrachus maxim us gen. et sp. nov. A, left ilium, Holotype QMF44207; B, left ilium, Paratype
QMF44208. Scale = 5mm. apt, anterior-most protruberance; pt, proximal protruberance.

rounded proturberance in lateral profile.
Posterior-most prominence placed superiorly
and central to acetabular fossa, llial shaft broad
posteriorly, tapering sharply to anterior end.

DESCRIPTION. Holotype. QMF44207, is a
large, robust left ilium preserving approximately
half the posterior end, including a portion of the
shaft, acetabulum, dorsal and ventral expansions
and prominences. Dorsal acetabular expansion
small and rounded superiorly, making a short

ilial/ischial suture. Ventral acetabular expansion
missing, broken on ventral margin, however,
inferred as small with short ilial/pubic suture.
Acetabular fossa large and well differentiated
from ilial shaft. Acetabular rim narrow and
produced laterally from the antero-posterior axis
of the ilial shaft. Ilial shaft incomplete, broken
approximately half way along its length,
compressed medially and rounded laterally when
viewed in cross-section. Preacetabular zone
broad, broken postero-ventrally of acetabular

PLIO-PLEISTOCENE FROG FROM QUEENSLAND

329

rim, superior to ventral acetabular expansion.
Ridge runs antero-dorsally from ventral margin
of acetabular rim, curves anteriorly half-way
along its length positioned on the midline of the
lateral side of the shaft. Curves antero-ventally
along last 1/3 of its length, terminating on ventral
margin of ilial shaft. Acetabular fossa eroded
along posterior margin of fossa and anterior
margin of acetabular rim. Dorsal prominence
divided into two by a large groove running
medio-laterally across ilial shaft. Groove
terminates in a small basin, or insertion point,
superior to dorsal acetabular rim. Anterior-most
protuberance elongate-ovoid and laterally
produced. Posterior-most protuberance ovoid in
shape and antero-posterior in orientation,
prominence terminating at the base of the dorsal
acetabular expansion. Ilial shaft broad along its
length, broken superiorly and perpendicular to
the long axis of the ilium. Ilial length: 13.10+mm,
dorsal acetabular expansion to ventral acetabular
expansion height: 6.38+mm. Paratype QMF44208
is an incomplete left ilium, preserving the
acetabulum, portions of both the dorsal and ventral
acetabular expansions, divided prominences and
preacetabular zone. Ilium large, robust along its
length, broken approximately 1/3 its length.
Ventral acetabular expansion unknown, inferred
as diminutive in antero-posterior length and
extending only slightly past ventral margin of
acetabular rim. Acetabular fossa large with
prominent narrow and high rim. Dorsal
prominence divided into an anterior and a
posterior prominence, bisected by a shallow
valley running from the anterio-medial side to the
postero- lateral side. Valley terminates as a small
basin (muscular insertion point) positioned
superior to the dorsal acetabular rim. Anterior-
most prominence laterally directed with an
elongate-ovoid protuberance in lateral profile.
Anterior-most prominence placed well anterior
of anterior acetabular rim. Posterior-most
prominence smaller than Anterior prominence
with a more rounded protuberance in lateral
profile. Prominence less laterally positioned,
terminating at the base of the dorsal acetabular
expansion. Posterior-most prominence placed
superiorly and central to acetabular fossa. Ilial
shaft broad posteriorly, tapering sharply to
anterior end. Ilial length: 10.20+mm, dorsal
acetabular expansion to ventral acetabular
expansion height: 5.14+mm.

REMARKS. The large size, divided prominence,
lateral mid-shaft ridge and reduced dorsal
acetabular expansion distinguish Etnabatrachus

from all other known Australian genera. The
rounded protuberance and lack of a dorsal crest
ally this taxon closest to Litoria of Australia.
However, specific placement in the Hylidae can
only be tentative at this point. Etnabatrachus is
considerably larger than the Oligo-Miocene
Litoria magna Tyler, 1991, which Tyler (1991)
considered to attain sizes of at least 120mm SVL.
This puts L. magna in a size range greater than the
largest extant Litoria (Cogger, 2000). It has been
hard to estimate the SVL of E. maximus based on
the lack of a complete ilium, however, the overall
size of the ilium indicates a frog of considerable
size, larger than 120mm SVL. The associated
faunas recovered from the type locality indicate a
palaeoenvironment of rainforest. Such a large
species of frog in a rainforest deposit is not
unusual as a number of species in present day
rainforests attain large sizes (e.g. Mixophyes
1 15mm SVL and Litoria 120mm SVL).

Etnabatrachus maximus sp. nov.

(Fig. 1)

ETYMOLOGY. Latin, maximus , alluding to the large size
of the species.

MATERIAL. HOLOTYPE: QMF44207, Paratype:
QMF44208 from Plio-Pleistocene deposits. Mount Etna
and Limestone Ridge (QML1284 & QML1385).

DIAGNOSIS. As for genus.

DESCRIPTION. As for genus.

ACKNOWLEDGEMENTS

I thank Rick Bell, Patrick Couper, Andrew
Amey, Gilbert Price, Noel Sands and Paul
Tierney for access to specimens, encouragement
and help in preparing the material for this study.

LITERATURE CITED

COGGER, H. 2000. Reptiles and Amphibians of
Australia. 5th Ed. (Reed: Melbourne).
FLANNERY, T.F., RICH, T.H., TURNBULL. W.D^&
LUNDELIUS, E.L. Jr 1992. The macropodoidea
(Marsupialia) of the early Pliocene Hamilton
Local Fauna, Victoria, Australia. Fieldiana
Geology NS 25: 1-37.

MUIRHEAD, J., DAWSON, L. & ARCHER. M. 1997.
Perameles bowensis , a new species of Perameles
(Peremelamorphia: Marsupialia) from Pliocene
faunas of Bow and Wellington Caves, New South
Wales. Proceedings of the Royal Society of New
South Wales 117: 163-173.

TEDFORD, R.H. 1 994. Succession of Pliocene through
medial Pleistocene mammal faunas of
southeastern Australia. Records of the South
Australian Museum. 27(2): 79-93.

330

MEMOIRS OF THE QUEENSLAND MUSEUM

TURNBULL, W.D. & LUNDELIUS, E.L. Jr 1970. The
Hamilton Fauna: a late Pliocene mammalian
fauna from the Grange Bum, Victoria, Australia.
Fieldiana: Geology NS 19: 1-163.

TYLER, MJ. 1974. First frog fossils from Australia.
Nature 248 (5450): 711-712.

1976. Comparative osteology of the pelvic girdle of
Australian frogs and description of a new fossil
genus. Transactions of the Royal Society of
South Australia 100(1): 3-14.

1982. Tertiary frogs from South Australia.
Alcheringa6: 101-103.

1985. Quaternary fossil frogs from Skull Cave and
Devil’s Lair in the extreme south-west of
Western Australia. Records of the Western
Australian Museum 12(2): 233-240.

1988. Neobatrachus pietus (Anura: Lepto-
dactylidae) from the Miocene/Pliocene boundary
of South Australia. Transactions of the Royal
Society of South Australia 1 12: 91 .

1989. A new species of Lechriodus (Anura:
Leptodactylidae) from the Tertiary of
Queensland, with a redefinition of the ilial
characteristics of the genus. Transactions of the
Royal Society of South Australia 113: 15-21.

1990. Limnodynastes Fitzingcr (Anura:
Leptodactylidae) from the Cainozoic of
Queensland. Memoirs of the Queensland
Museum 28(2): 779-784.

1991a. Crinia Tschudi (Anura: Leptodactylidae)
from the Cainozoic of Queensland, with
description of a new species. Transactions of the
Royal Society of South Australia 1 1 5(2): 99- 101.

1991b. Kyarannus Moore (Anura, Leptodactylidae)
from the Tertiary of Queensland. Proceedings of
the Royal Society of Victoria 103(1): 47-51.

1991c. A large new species of Litoria (Anura:
Hylidae) from the Tertiary of Queensland.
Transactions of the Royal Society of South
Australia 115(2): 103-105.

1994. Hylid frogs from the mid-Miocene Camficld
beds of Northern Australia. The Beagle, Records
of the Museum and Art Galleries of the Northern
Territory 11: 141-144.

1999. Australian Frogs: a natural history. (Reed:
New Holland).

TYLER, M.J., ASLIN, F.W. & BRYARS, S. 1992.
Early Holocene frogs from the Tantanoola Cave,
South Australia. Transactions of the Royal
Society of South Australia 1 16(4): 153.

TYLER, MJ. & GODTHELP, H. 1993. A new species
of Lechriodus Boulenger (Anura: Lepto-
dactylidae) from the early Eocene of Queensland.
Transactions of the Roval Society of South
Australia 117(4): 187489.

TYLER, M.J. & WALKLEY, R.W. 1996. First fossil
record of the hylid frog Litoria raniformis
(Keferstein). Transactions of the Royal Society of
South Australia 120(2): 69.

TYLER, MJ., DAVIS, A.C. & WILLIAMS, C.R. 1998.
Pleistocene frogs from near Cooma, New South
Wales. Proceedings of the Linnean Society of
New South Wales 119; 107-113.

TYLER, M,J., GODTHELP, H. & ARCHER, M. 1994.
Frogs from a Plio-Pleistocene site at Floraville
Station, Northwest Queensland. Records of the
South Australian Museum 27(2): 169-173.

MACROINVERTEBRATE FAUNA OF AN IRON-RICH STREAM IN THE WET
TROPICS OF AUSTRALIA: A COMPARATIVE ANALYSIS OF COMMUNITIES USING
A RAPID BIOASSESSMENT PROTOCOL

PIETER T.J. JOHNSON AND EUAN G RITCHIE

Johnson, P.T.J. & Ritchie, E.G 2003 06 30: Macroinvertebrate fauna of an iron-rich stream in
the Wet Tropics of Australia: a comparative analysis of communities using a rapid
bioassessment protocol. Memoirs of the Queensland Museum 49(1): 331-338. Brisbane.

ISSN 0079-8835.

The present study examined an iron-rich stream in the Wet Tropics of Queensland, Australia,
and used a Rapid Bioassessment Protocol to compare its macroinvertebrate fauna with that
of (1) a nearby, undisturbed stream and (2) the second order stream formed from their
junction. The undisturbed stream supported significantly greater levels of macroinvertebrate
abundance and taxonomic richness than either the iron-rich or junction stream. The latter two
streams did not differ significantly for either measure, suggesting that iron-layering effects
maintain potency even at increased distances from the source (>lkm). Percent Similarity
(PSy) and Shannon- Weiner Diversity (IT) indices were used to compare the streams’
macroinvertebrate communities, while two biotic indices (SIGNAL) were employed to
estimate water quality. The undisturbed stream exhibited greater invertebrate diversity and
higher water quality relative to the other two streams. Species assemblage patterns were
comparable to iron-rich stream studies from the temperate region: mayfly and caddisfly
nymphs were almost completely absent from the sites of iron-deposition while they
comprised the majority of the invertebrates in the undisturbed stream. The conservation
issues and management implications surrounding the release of water from stratified dams
and reservoirs are discussed. □ Iron-rich, rapid hioassessment protocol, macroinvertebrate
communities, Wet Tropics, dams.

Pieter T.J. Johnson * c& Euan G. Ritchie (e-mail: Euan.Ritchie@jcu.edu.au), School of
Tropical Biology \ James Cook University, Townsville, Queensland 4811, Australia.
(* Present address: Centre for Limnology, University of Wisconsin, Wisconsin 53706,
U.S.A.); 26 March 2002.

Freshwater organisms confront a unique set of
problems that are largely absent from terrestrial
and marine environments. Bodies of freshwater,
whether rivers, streams, ponds, or lakes, tend to
be ( 1 ) small with respect to occupied volume and
surface area, (2) patchy pattern across the land-
scape, and (3) sinks for pollutants and terrestrial
run-off. Consequently, freshwater systems are
highly sensitive to disturbance and pollution. The
most frequent forms of aquatic pollution include
thermal effluent, cultural eutrophication, acid-
ification, build-up of sediments or suspended
solids, and leakage of pesticides, petroleum
products, and heavy metals (Hynes, 1960;
Wiederholm, 1 984; Kim & Chon, 200 1 ; Mascaro
et al., 200 1 ). Construction of dams and other flow
impediments may accelerate or intensify these
processes. In addition to changing normal flow
regimes, dams increase bank erosion, disturb
flood cycles, impede migrations, displace native
species, and alter pH, temperature, salinity, and
natural chemistry of the waterway (Lake, 1967;
Walker, 1981; Palmer & O’Keefe, 1990).

Anthropogenic sources of iron that may affect
lotic systems include mining, logging, peat
production, and agricultural run-off (Vuori,
1995). Less commonly, iron deposition may
occur downstream from a stratified dam or
reservoir. Thermal stratification in reservoirs
creates an anoxic hypolimnion, and older dams
with deep-release outlets may send ammonium-
rich, oxygen-poor water downstream (Walker ct
al., 1978; Krenkcl et al., 1979; Marcus, 1980).
High quantities of iron can dissolve and
concentrate within the anoxic water and, upon
re-exposure to the oxygen-rich lotic environment,
precipitate out to form a layer of ferrichydroxide.
This deposition can seriously impact the resident
flora and fauna, and iron deposition has been
associated with diminished diversity and
abundance for most recorded taxa (Hildrew &
Townsend, 1976; Wellnitz et al., 1994; Wellnitz
& Sheldon, 1995; Vuori, 1995). The occurrence
of any anthropogenic impacts within the
relatively pristine Wet Tropics is a major concern.
The region is known as an area of significant
ecological importance sustaining high

332

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Location of the study site. (Numbers indicate
locations of sampling sites)

biodiversity, including many endemic and rare
species. To illustrate this point, the area is known
to contain 40% of Australia’s fish species (Pusey
& Kennard, 1996) and 25% of the terrestrial
vertebrates (Williams et al., 1 996) and 37% of the
plants (Keto & Scott, 1986) are known to be
regionally endemic.

We examined the macroinvertebrate fauna of a
stream with heavy ferric-hydroxide deposition
within the Wet Tropics of Australia. The stream
flowed from a base-level leak in a peripheral
saddle dam, wherein the iron is dissolved and
concentrated. As a natural ‘control’, samples
were also collected from a nearby stream which
neither originated from the dam nor showed any
signs of iron-deposition, but flowed over
identical topography, soil type, and vegetation.
The two streams eventually united into a second
order stream, and the fauna below this junction
was also sampled. Taxonomic richness, total
abundance, and species composition were com-
pared among the three areas. Two biotic indices
of water quality were also calculated: the Stream
Invertebrate Grade Number (SIGNAL) average
family level and the weighted index (Chessman,
1 995). The primary objective of this study was to
use a rapid bioassessment protocol (RBP) to assess
the effects of iron deposition on macroinvertebrate
communities in an Australian tropical stream.

MATERIALS AND METHODS

Paluma is situated in the Wet Tropics of North
Queensland (146°13’E, 19°S) and supports both
rainforest and eucalypt forest areas. Study sites
were located in an area of rainforest approx-
imately 500m downstream from the Paluma dam

(Fig. 1 ). At 890m above sea level, the dam itself
has a maximum capacity of 9.5 billion litres and
an average depth of approximately 15m. The
‘iron stream’ stems from a small leak at the base
of a peripheral saddle dam (R. Pearson, pers.
comm.) and is characterised by a distinctive,
ochre deposit covering all rocks and leaf litter
within the stream. An ostensibly undisturbed stream
(henceforth referred to as the ‘dear-stream’)
flowing from a nearby tributary was also sampled.
The third sample site was located in the second
order stream formed via the junction of the clear
and iron streams.

In each stream, two sites were sampled: one at
50m and one at 100m from the point of junction.
At each sample site, flow rate, canopy cover,
mean width, mean depth, temperature, and
substrate composition (percentages of cobble and
leaf-litter) were estimated. For each sample, five
similarly sized rocks were selected. Area was
estimated by measuring the two longest axes of
the rock; minimum and maximum recorded^ areas
among the samples were 10,400mm 2 and
51,300mm 2 , respectively. Each rock was lifted
rapidly from the substrate and placed in a 400pm
mesh dipnet, wherein the rock’s surface was
rigorously scraped free of all sediment and
organisms. After a final visual inspection, the
rock was replaced to the substrate while contents
of the net were transferred to 70% ethanol and
returned to the laboratory for sorting and
identification. Macroinvertebrate specimens
were identified to the level of family. A total of 30
rocks was examined.

Differences in macroinvertebrate abundance
and taxonomic richness (family level) among the
three streams were tested using Kruskal-Wallis
tests followed by paired Mann-Whitney U-tests
to isolate differences. To compare the structure
and assemblage of macroinvertebrate com-
munities, Shannon-Wiener Diversity (//’) and
Percent Similarity (PSy) indices (see equation 6.5
Jongman et al. 1995) were calculated. In
accordance with Chessman’s (1995) river rapid
assessment protocol, the weighted index
(SIGNAL-W) and the average family index
(SIGNAL) were estimated. In these indices,
specimens are keyed to family level and given an
interim pollution sensitivity grade value (1-10).
Chessman ( 1 995) provided such grades for common
families of Australian macroinvertebrates. For the
weighted index, each family is also assigned an
occurrence value (1-4), and the product of the
occurrence value and the pollution sensitivity
grade are summed across the sample. The

MACROINVERTEBRATE FAUNA OF AN IRON-RICH STREAM

333

resulting SIGNAL values give an indication of
water quality: clean water (value greater than 6),
doubtful quality (values between 5 and 6),
probable moderate pollution (values between 4
and 5), and probable severe pollution (values less
than 4). The protocol was designed to yield rapid
results and minimise time spent sampling and
sorting. Chessman et al. (1997) has revised the
grade numbers for a suite of macroinvertebrate
families. We decided against using this updated
SIGNAL index on the basis that the well
represented families in our data set have not
changed in grade number, and in addition some
families do not have an updated grade number,
hence allowing only a partial reanalysis.

RESULTS

The substratum of each stream was composed
of rocks and leaf litter. The mean size of rocks
(iron - 1 101cm 2 ±46, junction - 1609cm 2 ±252
and clear - 1555cm 2 ±96) did not differ
significantly between sites (ANOVA, d.f. = 2, 5,
F = 3 . 1 1 , p = 0. 1 9). The abiotic characteristics of
each stream are presented in Table 1 .

The three streams differed significantly with
respect to both macroinvertebrate abundance and
taxonomic richness (Kruskal Wallis test, df=2,
p<0.0002). The clear stream had significantly
higher levels of both macroinvertebrate abund-
ance and taxonomic richness than either the iron
or junction stream (Mann-Whitney U test, ni=
n 2 = 1 0, p<0.005; see Figs 2 and 3). No differences
were noted for either measure between the iron
and junction streams (Mann-Whitney U test, ni=
n 2 = 10, p>0.05). Thus, at the scale of study, no
increase in faunal abundance or taxonomic rich-
ness was noted with increasing distance from the
anoxic source. Accordingly, visual observations
showed no obvious decline in the layering of
ferrichydroxide, even beyond the junction of the
clear and iron streams. In both streams, several
rocks were completely devoid of macroinvertebrates,

TABLE 1. Abiotic descriptors for each stream. All
values represent the mean for each stream.

Iron Stream

Clear Stream

Junction

Stream

% Leaf

37.5

16

22-5

II % Cobble

45

60

60

% Canopy Cover

91

90

85

Width (m)

1.5

1.75

1.9

Depth (cm)

7.5

28.75

15

1 Flow (m/s)

0.24

0.03

0.13 |

in marked contrast to the densely populated rocks
of the clear stream.

The similarity, diversity, and water quality
analyses supported the trends noted for
abundance and taxonomic richness. With respect
to the collected families, percent similarity
calculations (PS,y) demonstrated a 40% similarity
between the iron and junction communities,
compared to 10.8% similarity between the
junction and the clear streams and a 4.4%
similarity between the clear and iron streams. For
the Shannon-Weiner diversity index and the
biotic indices of water quality, the clear stream
had considerably higher values than either the
junction stream or the iron stream (Table 2). The
SIGNAL values classified the water of the clear
stream as ‘clean’, while both the iron and junction
streams fell into the ‘moderately polluted'
category (Chessman, 1995). Species assemblages
from the different streams also differed sub-
stantially. The fauna list for the iron stream had
no representatives of either Trichoptera or
Ephemeroptera, which comprised approximately
50% and 25% of the clear stream fauna,
respectively. The junction sample included one
family in Trichoptera but lacked Ephemeroptera
(Table 3).

DISCUSSION

Despite the temporal and spatial limitations
inherent to the Rapid Bioassessment Protocol
used in this study, it appears that iron deposition
had substantial negative impacts on macro-
invertebrate abundance and family-level
richness. All indices had high values in the clear
stream and low values in the iron and junction
streams. This trend was most extreme for
abundance, and clear stream samples supported
more than ten times the number ol macro-
invertebrates found in either of the other streams.
Another intriguing aspect of the results was the
persistence of these effects with distance
downstream. The fauna of the uppermost iron

TABLE 2. Results of the three indices calculated for
each stream: Shannon-Weiner Diversity Index,
SIGNAL (Stream Invertebrate Grade Number -
Average Level per family), and SIGNAL-W
(weighted by the occurrence of each family).

1

Iron Stream

Clear Stream

Junction

Stream

Shannon-Weiner

2.07

2.19

2.07

SIGNAL

4.14

6.44

4.71

1 SIGNAL-W

4.20

6.50

4.23

334 MEMOIRS OF THE QUEENSLAND MUSEUM

Clear Jinction Iron

Stream

FIG. 2. Mean number of macroinvertebrates per
stream. (Letters indicate significance groupings)

sampling site and the lower junction site were
equally depauperate, despite more than a kilometre
separation, and supported similar macroinvertebrate
communities. An exceptionally large distance may
be necessary to observe any indications of
‘recovery’ . A study by Palmer & O' Keefe ( 1 990)
of the downstream effects of impoundments on
water chemistry, has shown polluted streams may
require up to 100km before regaining natural
assemblages. In their study potential pollutants
included agricultural runoff and urban effluents.

Iron-rich streams negatively impact macro-
invertebrate abundance and diversity and alter
the remaining species assemblage by affecting
taxa differentially (Hynes, I960). Previous
studies of iron-rich streams correlate increases in
iron deposition with the disappearance of most
flora and fauna (Wellnitz et al., 1994). While still
requiring experimental confirmation, the
sediment layer created by the iron is the most
likely causative agent. After leaving the
reservoir, water from the iron stream is no longer
anoxic, as the stream is shallow, fast flowing, and
the iron has oxidised out of solution. The ochre
layer of ferrichydroxide may negatively
influence macroinvertebrates in at least five
ways: by (1) reducing habitat complexity and
available shelter, (2) interfering with holdfast
mechanisms of certain species, (3) limiting
periphytic algae colonisation, thus decreasing
primary productivity and disturbing trophic
relationships, (4) coating and blocking animals’
respiratory surfaces, and (5) inhibiting proper ion
exchange and osmoregulation (Hynes, 1960;
Hildrew & Townsend, 1976; McKnight & Feder,
1984; Vuori, 1995). The possibility of toxic

Clear Junction Iron

Stream

FIG 3. Mean number of macroinvertebrate families
per stream. (Letters indicate significance groupings)

effects of iron traces in the water cannot be
discounted, especially when levels are well
above what is perceived as normal for rainforest
streams in an undisturbed state. Unfortunately,
few of these possibilities have been satisfactorily
tested and their relative importance remains
unknown. Presumably, the vertebrate fauna of
streams is also affected by iron deposition,
whether directly or indirectly. Within this study at
least, Myxophyes schevilli tadpoles were found
within the clear stream but not in either the iron or
junction sites.

The impacts observed in the current RBP study
parallel those of previous studies of iron-rich
streams: a diminished fauna consisting of select
macroinvertebrate orders. While the order
Ephemcroptera is usually well represented in
pristine, lotic streams, its absence from iron-rich
streams is a universal trend in the literature. In
general, mayfly nymphs are among the most
sensitive aquatic insects to pollution ( Wiederholm,
1984). Chessman (1995) gave Leptophlebiidae,
three genera of which were represented in the
clear stream but not in neither iron-rich stream, a
pollution sensitivity grade of ten. Wellnitz et al.,
(1994) found three mayfly species showing
increased mortality when caged within an area of
high ferrichydroxide deposits. Similarly,
Harding & Winterbourn (1995) noted poor
mayfly representation in pastoral streams with
high levels of iron relative to low-iron forest
streams. The heightened sensitivity of mayflies is
probably related to (1) the exposed nature of

MACROINVERTEBRATE FAUNA OF AN IRON-RICH STREAM

335

Ephemeroptera gills and (2) their dietary
dependence on periphytic algae (Roback, 1 974).

The paucity of Trichoptera representatives in
iron and junction streams also mirrors results
from past studies in temperate areas (Wiederholm,
1984; Harding & Winterboum, 1995). The two
trichopteran families abundantly represented in
the clear stream, Helicopsychidae and Lepto-
ceridae, have sensitivity grades of ten and seven,
respectively (Chessman. 1995). In contrast, the
one caddisfly family found in the junction
stream, Ecnomidae, is ranked with a grade of four
(Chessman, 1995).

The report of an iron-rich stream within the
relatively undisturbed Wet Tropics of north
Queensland is potential cause for concern. The
volume of water contributed by the iron and clear
streams into the junction stream are comparable,
but the biotic parameters of the second order
stream are clearly more aligned with the iron
stream. This finding was supported by all tested
indices. In many respects, streams represent the
veins of the surrounding landscape, upon which
the health of the entire ecosystem may depend.
Disturbances impacting waterways can create a
type of ‘ripple effect’, influencing aquatic and
teirestrial systems for a considerable distance. In
this study, the ripple created by the anoxic
stratification of Paluma dam maintained a nearly
constant, negative amplitude for over a kilometre
downstream. Mono- and biphasic aquatic
vertebrates, not to mention terrestrial flora and
fauna could suffer direct or indirect harm in areas
close to iron-enriched streams. A number of
stream breeding frogs have disappeared or are in
decline in the Australian Wet Tropics, and further
deleterious anthropogenic effects must be
minimised (Williams & Hero, 1998).

The absence of multi-depth release valves in a
dam or the presence of a fracture leak in a
reservoir may seem of minor consequence, but
the ecological reverberations can be intense. Iron-
rich streams have an altered macroinvertebrate
community structure and a shorter trophic chain
than their unaffected counterparts (Hildrew et al.,
1985). An iron sheath bacteria, Leptothrix
ochracea , predominates in the presence of iron
sediment but is absent in streams undisturbed by
ferric hydroxide deposition. Leptothrix ochracea
reduces naturally important populations of
diatoms upon which many aquatic insects and
other invertebrates graze' (Sheldon & Skelly,
1990). Much of the natural periphyton is
similarly lost following an influx of iron

sediment. In England, several studies have
focussed on iron streams due to the simplicity of
their trophic structure (Hildrew et al., 1985;
Lancaster & Robertson, 1995). Although little
work has been done on iron-rich streams in
tropical regions, the Rapid Bioassessment
Protocol used in this study yielded the results one
would expect with a more quantitative procedure,
as illustrated by previous studies of temperate
iron streams. Chessman et al. (1997) highlighted
the uncertainty around the application of
SIGNAL to other river systems, as the relation-
ship between pollution-sensitivity grade
numbers and types of polluting agents and
disturbances remains unclear. We therefore
acknowledge that the results of this study should
be considered with appropriate caution until
grade numbers developed for specific disturb-
ances become available.

Results of this study are of importance as they
demonstrate one of many impacts dam con-
struction can have on local freshwater systems.
This is especially disturbing if we consider the
future plans for water resources in Queensland. In
a recent release from the Department of Natural
Resources (1997), 93 proposals for waterway
development were recommended as having merit
for inclusion in the Queensland Water Conserv-
ation Strategy. Considering that construction of
dams typically alters the flow regime and water
quality of local waterways, these proposals have
the potential to increase environmental degrad-
ation of Queensland’s aquatic habitats. This
warrants an increased effort to monitor the
impacts of dam construction on the health of
aquatic ecosystems and employ such information
in subsequent plans for further dam development.
Without such research, Queensland, and possibly
other regions of the world are in danger of jeopard-
ising the future of their natural waterways.

Continued study of these Paluma streams will
focus on three aspects: (1) persistence of the
effects of iron deposition with increasing
distance from the dam, (2) consequences of iron
deposition on aquatic macroinvertebrates and
streamside fauna, and (3) exact mechanisms
through which deposited iron acts as such a
potent disturbance.

ACKNOWLEDGEMENTS

We extend sincere appreciation to Gordon
Kovacs for considerable advice and assistance in
the keying of macroinvertebrate samples, and to
Richard Pearson, for initially directing us toward
the study of iron-rich streams and supplying

336

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3. Taxonomic list and relative abundance of macroinvertebrates in the Clear, Iron and Junction streams at
Paluma.

Order Suborder

Family

Genus

No. Individuals

% of Total

Iron

Arachnida

Porohalacaridae

3

0.20

Coleoptera

Elmidae

1

0.07

Diptera

Tanyderidae

2

0.13

Diptera

Thaumaleidae

1

0.07

Diptera

Unidentified

1

0.07

Gastropoda

Planorbidae

Segnitila

1

0.07

Hemiptera

Veliidae

Rhagovelia

4

0.27

Odonata

Anisoptera

Unidentified

1

0.07

Plecoptera

Gripopterygidae

1

0.07

Clear

Diptera

Ceraptopogonidae

2

0.01

Diptera

Chironomidae

19

0.11

Diptera

Dixidae

1

0.01

Diptera

Empididae

1

0.01

Diptera

Psychodidae

2

0.01

Ephemcroptera

Caenidae

7

0.04

Ephemeroptera

Leptophlebiidae

Atalophlebia

13

0.08

Ephemeroptera

Leptophlebiidae

A ustrophlebiodes

7

0.04

Ephemeroptera

Leptophlebiidae

Ulmerophlebia

8

0.05

Ephemeroptera

Leptophlebiidae

Unidentified

8

0.05

Ephemeroptera

Unidentified

2

0.01

Hemiptera

Veliidae

Rhagovelia

2

0.01

Odonata

Anisoptera

Corduliidae

2

0.01

Odonata

Anisoptera

Unidentified

1

0.01

Odonata

Zygoptera

Chlorolcstidae

2

0.01

Odonata

Zygoptera

Lestoideidae

2

0.01

Plecoptera

Eustheniidae

1

0.01

Trichoptera

C'alamoccratidae

5

0.03

Trichoptera

Ecnomidac

4

0.02

Trichoptera

Helicopsychidae

37

0.22

Trichoptera

Lcptoceridae

41

0.24

Trichoptera

Philorheithridae

1

0.01

Trichoptera

Polycentropodidae

1

0.01

Junction

Arachnida

Porohalacaridae

3

0.18

Diptera

Chironomidae

4

0.24

Diptera

Empididae

1

0.06

Diptera

Tipulidae

1

0.06

Diptera

Unidentified

1

0.06

Hemiptera

Veliidae

Rhagovelia

1

0.06

Odonata

Anisoptera

Corduliidae

2

0.12

Odonata

Zygoptera

Amphipterygidae

2

0.12

Trichoptera

Ecnomidae

2

0.12

MACROINVERTEBRATE FAUNA OF AN IRON-RICH STREAM

337

guidance in both experimental design and data
analysis for the project. Richard Rowe provided
critical information regarding the life histories of
some of the collected taxa, for which we are
likewise grateful. We are indebted to Zury Soto,
Sapna Khandwala, and Eric Beerbohm for
editing the manuscript. Marg Criniti deserves
special mention for furnishing the necessary
supplies and storage space during the course of
the study. Logistical support for this project was
provided by the Australian Centre for Tropical
Freshwater Research.

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CASE HISTORIES OF ATTACKS BY THE SOUTHERN CASSOWARY IN

QUEENSLAND

CHRISTOPHER P. KOFRON

Kofron, C.P. 2003 06 30: Case histories of attacks by the Southern Cassowary in Queensland.
Memoirs of the Queensland Museum 49(1): 339-342. Brisbane. ISSN 0079-8835.

The Southern Cassowary is Australia's largest bird and is capable of causing serious injuries
to humans. The incidence of cassowary attacks in Australia was reviewed recently by Kolron
(1999). The eight recorded incidents in which humans were seriously injured by Southern
Cassowaries are examined, and also other related incidents ot special interest. The eight
seriously-injured victims suffered puncture wounds, lacerations, cuts, and/or broken bone,
one of them subsequently dying. Five of the eight attacks were by cassowaries that had been
fed previously by people, and in the single fatal cassowary attack recorded in Australia the
victim was trying to kill the cassowary. Also, four of the eight seriously-injured victims were
crouching or lying on the ground. Consequently a person encountering a cassowary should
not crouch down, which puts a person's head and vital organs in closer striking range of the
cassowary's foot. Rather, a person should remain standing and move behind a tree, or quickly
leave the area but without turning one's back. Children should be supervised and kept at safe
distances from cassowaries, especially cassowaries habituated to people. People should be
aware that cassowaries can be dangerous in certain situations. □ Southern Cassowary,
Casuarius casuarius johnsonii, Queensland, attacks.

Christopher P. Kofron, Queensland Parks and Wildlife Service, Northern Regional Centre,
PO Box 2066, Cairns 4870, Australia; present address, US Fish and Wildlife Service, 2493
Portola Road, Suite B, Ventura . Ca . 93003. USA (e-mail: chris.kofron@fvs.gov); 28 August 2002.

The Southern Cassowary (Casuarius casuarius
johnsonii , family Casuariidae) is endemic to the
tropical rainforests of northeastern Queensland.
This species is Australia’s largest bird and holds a
reputation for being dangerous. Cassowaries and
Ostriches are the only birds worldwide that have
caused human deaths by physical attack.
Incidents occur every year in Queensland, most
at Mission Beach (110km SE Cairns) but
previously also at Lake Barrine (39km SW
Cairns) and Mount Whitfield in Cairns.

The incidence of cassowary attacks in
Queensland was reviewed recently by Kofron
(1999), who reported on data for 221 attacks
against humans and domestic animals. One
hundred fifty attacks were against humans, 75%
of these by cassowaries fed previously by people.
The feeding of cassowaries appears to change
their natural behaviour, making them bold and
aggressive. Victims were chased or charged in
71% of the incidents, and kicked in 15%. Less
frequent actions included pushing, pecking,
jumping on, butting with the head and snatching
food. The cassowaries appeared to be expecting
or soliciting food from humans (73% of the
incidents); defending food (5%); and defending
themselves (15%) or their chicks or eggs (7%).
The purpose of the present paper is to examine
the eight recorded incidents in which humans

were seriously injured, and also other incidents ot
special interest, with a view to identifying the
hazards and minimising the risks in human-
cassowary interactions.

METHODS

I questioned and interviewed persons likely to
have knowledge of or experiences with
cassowaries (e.g. rangers, field workers, farmers,
veterinarians, wildlife carers). In addition, I
examined official records of the Queensland
Parks and Wildlife Service and the Queensland
Police Service, and also newspaper reports. All
information presented here is based on first
person and eyewitness accounts, except one
incident known only from two newspaper
reports. An injury was considered serious if it
required treatment by professional medical
personnel, and these included deep puncture
wounds, lacerations and cuts needing stitches,
and fractured bone. Where possible, 1 spoke with
the victims and the medical personnel.

RESULTS

ATTACKS TO HUMANS RESULTING IN
SERIOUS INJURIES. Eight attacks by wild
cassowaries resulted in serious injuries to
humans in the following parts of the body: throat;
buttocks; chest; scrotum, chest and forearm;

340

MEMOIRS OF THE QUEENSLAND MUSEUM

upper arm; thigh and forehead; lower leg; and
middle finger. One victim subsequently died. The
cassowaries kicked or jumped on the victims,
four which were crouching or lying on the ground
(one tripped, one knocked to the ground). In three
of these attacks the cassowaries appeared to be
soliciting food and in three attacks acting in
self-defence; but two of the attacks remain
unexplained.

Boy killed near Mossman, puncture wound in
neck, 1926. On 6 April 1926, two dogs and two
boys (Phillip McLean 16 years, Granville
McLean 13) attacked a cassowary at their
paddock near Mossman (67km NW Cairns),
striking the bird and trying to kill it. The
cassowary kicked one dog and the younger boy,
who stumbled away, but the other dog and boy
pressed on with their attack. When the cassowary
advanced on the older boy, he ran but tripped,
falling to the ground face upwards. The
cassowary jumped on him, its spike-like claw
penetrating the boy’s neck (puncture wound 1 V 4
cm diameter) and severing a blood vessel. The
boy got up and ran, but soon collapsed and died
from loss of blood.

Woman kicked on Mount Whitfield track , wound
in buttock, 1988. Two newspaper articles
published on 13 March 1988 (Anonymous,
1988a) and 15 March 1988 (Anonymous, 1988b)
reported that a woman was attacked by a
cassowary at Mount Whitfield, and the wound in
her buttock required stitches. Although one
article claimed that the woman was bitten by the
cassowary, it is likely that she was kicked. The
recent cassowaries at Mount Whitfield were fed
by people.

Man kicked at Julatten, puncture wound in chest,
1990. On 2 October 1990, Barry and Jenny Tuite
walked along Clacherty Road in Julatten (57km
NW Cairns), bound by rainforest, orchards,
paddocks and houses. They did not notice the
cassowary at the rainforest edge until only
several metres distant from it. They walked away,
but the bird followed for about 50m. Suddenly the
cassowary charged, leapt and kicked the man in
his chest (puncture wound 1 2 cm deep), knocking
him to the ground. As he lay unconscious in a
drain, the cassowary attempted to kick him again.
The woman shouted; the cassowary advanced
towards her, and she ran up an embankment. The
injured man regained consciousness and also
climbed the embankment. The woman ran to a
nearby farmhouse for help, with assistance
arriving as the cassowary again was confronting
the injured man. The bird moved away only a

short distance as Mr Tuite was assisted into a
motor vehicle. When the cassowary approached
again, the fanner shot and killed it.

The reason for the attack is unknown. No dogs
or cassowary chicks were involved, and residents
denied feeding the female cassowary, although
one person alleged the contrary. One day prior to
the attack, two people on horses were chased by a
cassowary along Clacherty Road; and one or two
weeks prior in Julatten, cows were chased by a
cassowary.

Man kicked at Cape Tribulation, lacerations in
forearm, chest and scrotum, 1991 . On 1 1
September 1991 at 02:00, Bodo Wintergerst
walked out of his caravan at Bailey's Creek
( 88 km NW Cairns) to toilet in the rainforest. He
claims that a cassowary kicked him while
squatting, lacerating his forearm, chest and groin.
The linear cuts required 9 stitches: right forearm
2, right upper chest 3 and left side of scrotum 4.
The alleged attack cannot be explained.

Woman kicked on Mount Whitfield track,
puncture wound in upper arm, 1994. On 20
December 1994, Joanne Edwards was walking
the Blue Arrow Track at Mount Whitfield when a
cassowary approached her from behind. The bird
stopped 5m away and preened its chest feathers.
The woman stepped backwards, and the
cassowary advanced. The woman crouched
down and covered her head with an arm. The bird
kicked Ms Edwards in the arm, knocking her to
the ground, and then kicked her several times
more in the arm and stomach. Eventually the
cassowary walked away. The woman received a
puncture wound in the upper ami that required 1 0
stitches.

Woman kicked on Mount Whitfield track,
puncture wound in thigh, lacerations in forehead,
1996. On 30 March 1996, Jill Turner was walking
the Blue Arrow Track at Mount Whitfield when a
cassowary approached her from behind. The
woman continued walking, and the bird
followed. Eventually she stepped off the track to
allow the cassowary to pass, but it stopped at 1 m
distance. The cassowary briefly looked at the
woman and then kicked Ms Turner in the thigh,
knocking her to the ground. The cassowary
quickly kicked her again in the forehead and
twice in the back. The cassowary sat down next to
her ( 2 m distance) for 10 minutes; then got up,
walked 4m and sat down for 30 minutes; and then
got up and walked away down the track. The
woman received puncture wounds in her thigh
and lacerations in her forehead that required two
and nine stitches, respectively.

SOUTHERN CASSOWARY ATTACKS

341

Man kicked at Stratford, Mount Whitfield,
fractured lower leg, 1 996. On 2 November 1 996,
James Barry saw two dogs attacking a cassowary
on the residential lot adjacent to his, one biting
the bird’s neck and the other its leg. As he pulled
one dog away, the cassowary kicked Mr Barry in
the shin, causing a simple fracture of his lower
leg. The cassowary was obviously acting in self
defence (it subsequently died), not realising that
Mr Barry was attempting to help it. This bird had
a history of being fed by people.

Man kicked at Etty Bay, cut on finger, 1 999. On 25
December 1999, George Newport saw two
children on bicycles confronted by a cassowary
at his caravan park. To assist the children and
shoo away the bird, Mr Newport approached the
cassowary with an umbrella, holding it with two
hands in front while opening and closing several
times, which had proven effective on previous
occasions. As Mr Newport neared the cassowary,
the bird jumped up and kicked the umbrella, also
striking the man’s hand. The cassowary then
walked away. Mr Newport suffered a cut to his
middle finger that required five stitches. This bird
had a history of being fed by people.

SOME OTHER ATTACKS TO HUMANS. Man
kicked at Mission Beach, 1 986. In one incident in
1986 at Mission Beach, a tourist was viewing a
cassowary. The bird approached the person to
within lm, apparently expecting to be fed.
Instead of offering food, the tourist squatted
down to photograph the cassowary. Suddenly the
bird kicked the person in the stomach, knocking
him to the ground.

Cassowary? jumped up and down on man at
Aeroglen, Mount Whitfield, 1991. In November
1991, a friend was visiting Joseph Collins at his
residence on Mount Whitfield. The fig tree in his
yard was frequented by a cassowary, and the bird
was feeding during the visit. The two men viewed
the cassowary at close quarters, picked up figs
and fed it by hand. The visitor moved backwards
to photograph the bird, lost balance and fell to the
ground. The cassowary quickly jumped onto the
prone man's back, and then jumped up and down
on him about 10 times. This action of the
cassowary appears to have been in defence of its
food and feeding area.

Man head-butted on Mount Whitfield track,
1995. On 29 October 1995, three hikers on the
Blue Arrow Track at Mount Whitfield encountered
a cassowary preening itself. The hikers went off
the trail intending to circumvent the bird, but
upon their emerging back onto the track the

cassowary was waiting for them only 2m away.
The cassowary lowered its head, charged and
butted Clive Skarott in the back. While falling he
grabbed a narrow tree trunk, the momentum
spinning him around the tree and back again to
face the cassowary. The man quickly retrieved an
apple from his pocket and offered it to the
cassowary, which the bird ate from hand. The
man offered a second apple, which was also
eaten. A third apple was offered, but then rolled
down the slope, which the cassowary pursued.
Boy pecked on testicles at Aeroglen, Mount
Whitfield, 1995. In another incident in October
1995 at Mount Whitfield, the son of Ann Keating
went outdoors to play. A cassowary was feeding
at a fig tree in their garden, which the bird
frequented regularly. The little boy (age four)
climbed onto a fence post about 1 / 4 m height, and
the cassowary walked to him. The cassowary
pecked the boy, striking him on the testicles. I
attribute this action of the cassowary to defence
of its food and feeding area.

SERIOUS INJURIES TO DOMESTIC
ANIMALS. Although injuries to dogs are included
here, the dogs were usually the aggressors and
were threatening the cassowaries. In 29 of 35
recorded incidents, the cassowaries were acting
in self defense because they were being attacked
by dogs. Conversely and most importantly, many
cassowaries have suffered injuries and even
death because of dogs, which will be the subject
of a separate paper.

Dog killed at the Tully River, puncture wound in
chest, 1976. One day in 1976 Les Henry drove
with his dog to the Tully River ( 1 24km S Cairns).
Upon getting out of the motor vehicle, the dog
sighted a cassowary, then barked and ran to it. As
the dog drew near, the cassowary took several
steps toward the dog, and then kicked the dog in
its chest making a deep puncture wound. The dog
died shortly after.

Dog killed at Bramstom Beach, ruptured
intestine, 1995. In one incident in 1995 at
Bramston Beach (54km SE Cairns), a dog was
chasing a cassowary. The cassowary turned to
confront the dog, and then kicked the dog in its
belly. Although there was no puncture wound,
there was massive bruising. The dog
subsequently died, apparently from a ruptured
intestine.

20 dogs kicked, puncture wounds, lacerations
and cuts, 1976-1996. Six veterinarians in
Innisfail (72km SE Cairns), Mission Beach and
Tully treated 20 other dogs with puncture wounds,

342

MEMOIRS OF THE QUEENSLAND MUSEUM

lacerations and cuts caused by cassowaries. All
injuries were in the torso (10 in the chest), and
each dog required 2-20 stitches.

Horse kicked at South Mission Beach, laceration
in buttock, 1987. Rangers Brent Vincent and Karl
Siener conducted a situation assessment of a race
horse that had been kicked by a cassowary at
South Mission Beach in 1987. This particular
cassowary was being fed fruit (pineapples,
bananas, apples) on the adjacent farm as part of
the wildlife rescue effort following Cyclone
Winifred. While the horse was feeding at its own
trough, a cassowary kicked the horse twice, in its
ribs and buttock. The horse suffered a deep
laceration about 1 8cm long in the upper thigh that
required stitches.

DISCUSSION

Eight attacks against humans resulted in
serious injuries and subsequently one death,
caused by cassowaries kicking or jumping on
victims. Five of these eight attacks were by
cassowaries that had been fed previously by
people. In the single fatal cassowary attack
recorded in Australia, the victim was trying to kill
the cassowary. In consideration of all the data, I
conclude that feeding cassowaries changes their
natural behaviour, making them bold and often
aggressive towards humans. Cassowaries that are
fed become habituated to humans, subsequently
recognising humans as a source of food. For
example, of the 90 cassowary attacks recorded at
Mount Whitfield (Kofron, 1999), I suspect that
no more than four cassowaries were involved,
and some of the serious injuries were probably by
the same cassowary.

Four of the eight human victims that suffered
serious injuries were crouching or lying on the
ground, and 22 dogs were seriously injured when
kicked. Consequently a person encountering a
cassowary should not crouch down, which puts a
person’s head and vital organs in closer striking
range to the cassowary’s foot. Rather, a person
should remain standing and move behind a tree,
or quickly leave the area but without turning

one’s back. Children should be supervised and
kept at safe distances from cassowaries, especially
cassowaries that are habituated to people.

Much of the Southern Cassowary’s tropical
rainforest habitat is now highly fragmented, and
human-cassowary interactions will continue to
occur, especially in the Mission Beach/Tully/
Innisfail area. People must be aware that
cassowaries are capable of causing serious
injuries, and they can be dangerous in certain
situations.

The Southern Cassowary is an endangered
species, and one of the threatening factors is
dogs. Owners should not allow dogs to roam at
large because of their impact on native Australian
wildlife, including cassowaries.

ACKNOWLEDGEMENTS

1 thank everyone who engaged in conversation
about cassowaries, especially Angela Chapman,
Clifford Frith, Stephen Garnett, Geoff Kelly, Roy
McKay, Les Moore, Michael Prociv, Mary
Ritchie and the Community for Coastal and
Cassowary Conservation (C4), Peter Saleras,
Karl Siener, Daryn Storch, Keith Smith, Rusty
Smith, Jim Thompson, Michael Trenerry and
Brent Vincent.

The views expressed in this paper are those of
the author and do not necessarily reflect those of
the Queensland Parks and Wildlife Service or
those of the Queensland Government.

LITERATURE CITED

ANONYMOUS 1988a. Forest killer bites joggers
Sunday Sun 13 Mar.: 29.

ANONYMOUS 1988b. Cassowary charges walkers.
Cairns Post 15 Mar.: 3.

KOFRON, C.P. 1999. Attacks to humans and domestic
animals by the Southern Cassowary ( Casuarius
casuarius johnsonii) in Queensland, Australia.
Journal of Zoology ( London) 249: 375-381.

The Board of the Queensland Museum gratefully
acknowledges the financial contribution of the
author in the publication of this paper.

TWO NEW SPECIES AND OTHER RECORDS OF SEA SPIDERS (PYCNOGONIDA:

ARTHROPODA) FROM TROPICAL NORTH QUEENSLAND

A.C. LEE AND C.P. ARANGO

Lee, A.C. & Arango, C.P. 2003 06 30: Two new species and other records of sea spiders
(Pycnogonida: Arthropoda) from tropical North Queensland. Memoirs of the Queensland
Museum 49 ( 1 ): 343-348. Brisbane. ISSN 0079-8835.

Five species of sea spiders found among the green intertidal algae Cladophora prolifera
(Roth) and the coastal soft bottoms of Cleveland Bay off Townsville, are reported. Two new
species are described; Ammothella fistella sp. nov., a closely related species ol the
appendiculata-group, and Callipallene catulus sp. nov., an extremely small species
differentiated by its size and general body shape. Three species ot Anoplodactylus are also
recorded, A. glandulifer a widely distributed species, A. cribellatus , a recent synonym ot the
Australian A. simplex , and A. tubiferus , a long-known species from the West-Pacific. The
finding of undescribed species in the alga C. prolifera encourages the interest on intertidal
habitats that might serve as shelter for possibly many others small to tiny pycnogonids. □
Pycnogonida, Ammothella, Callipallene, tropical sea spiders.

A.C. Lee, School of Marine Biolog y and Aquaculture, James Cook University’, Townsville
4811 (present address: Tropical Marine Science Institute, National University of Singapore,

14 Kent Ridge Road, Singapore 119223: e-mail: tmsleeac@nus.edu.sg); C.P. Arango,
Department of Zoology and Tropical Ecology’, James Cook University’, Townsville 4811,
Australia, (present address: Division of Invertebrate Zoology , American Museum of Natural
History, Central Park West at 79 Street, New York, NY 10024, USA; e-mail:
carango@amnh.org): 21 February 2002.

This is a report of a small collection of
pycnogonids from Townsville, in tropical North
Queensland. It comprises three named species
plus the description of two new species, adding
them to the list of 34 species known from
Queensland. Studies on pycnogonid fauna from
the North Queensland region are few and
sporadic. Reports on the fauna have rarely been a
result of active search. Instead, many of the
specimens found were primarily taken from
dredge and grab, or trawled samples, usually with
expeditions (Clark, 1963; Child, 1975). In the last
20 years the only reports on sea spiders from
Australia are those by Staples (1982), on
pycnogonids from the vicinity of Gladstone,

Child ( 1 990) on the 1 5 species reported from the
Great Barrier Reef, and Arango (2003) also
recording new species from North Queensland.

The present note constitutes a complement to that
larger report on pycnogonids from tropical
Queensland and nearby areas (Arango, 2003).

The discovery of two new species of pycno-
gonids in a very small collection of green algae is
an indication that the pycnogonid fauna of this
region is much more diverse than previously
thought, especially of very small forms of
pycnogonids that predominate in a diversity of
shallow water habitats.

Rowes Bay (19°15’S, 146 °47’E) and Cleveland
Bay constitute the area of collection ot the
present report. The intertidal habitat of Rowes
Bay, composed of small boulders and a mixture
of muddy and sandy sediment and a large
diversity of benthic organisms and various algal
species, was explored by the authors in different
times of the year in 1998 and 1999. Aside from
this, dredge and grab collections of the soft
benthos of Cleveland Bay were obtained.
Anoplodactylus species were dominant, lollowed

by Ammothella and Callipallene, Anoplodactylus
glandulifer Stock, 1954, Ammothella fistella sp.
nov. and Callipallene catulus sp. nov. were
collected among the algae Cladophora prolifera
(Roth); while A. cribellatus Caiman, 1923 (syn.
A. simple * Clark, 1963) and A. tubiferus Haswell,
1884 were collected by dredge and grab in the
muddy bottoms of Cleveland Bay.

Although the area of North Queensland might
be a well-studied region regarding the
pycnogonid fauna compared to other Australian
regions, the two previously undescribed species
in such a small collection suggests that little
collecting has been done, especially of very small
forms of sea spiders that predominate in a
diversity of shallow water habitats.

344

MEMOIRS OF THE QUEENSLAND MUSEUM

SYSTEMATICS

AMMOTHEIDAE Dorhn 1881
Ammothella fistella sp. nov.

(Fig. 1)

ETYMOLOGY. Latin, fistella . diminutive of fistula,
meaning a pipe, tube or ulcer; suggested by Dr C. Allan
Child. It refers to the various tubular spines on the lateral
processes and appendages.

MATERIAL. Townsville, Rowes Bay, intertidal rocky,
found on the green alga, Cladophora pmlifera (Roth) in
0.5m, April-November 1998,1 <3, holotype, (QM
SI 05865). coll. Lee; April-November 1998. Paratypes, 1 6
with eggs, 3 9, 16juv. paratypes(QMS105866),4d,29, 1
juv., coll. Lee; April-Novl998, 1 d with eggs, 19,1 juv.,
(QMS 105866), coll. Arango; Apr. 1999, 1 d, 49, 1 juv.
(QMS 1 05867), coll. Arango.

DISTRIBUTION. Known only from Rowes Bay,
Townsville, Qld, 0.5m tidal height.

DESCRIPTION. Relatively small, leg span of
6.84mm. Trunk robust, spinose, with numerous
short tubular and long pointed spines. Lateral
processes separated by half their diameter or less,
ornamented with three similar dorso-distal
tubular spines, last pair of lateral processes with
one single tubular spine: tubular spines all taller
than their diameters. Abdomen long, curving
posteriorly, bending horizontally at midpoint,
ornamented with four short and two long tubular
spines and four short pointed lateral spines.
Ocular tubercle of moderate length, pointed
anteriorly, 2.5 x longer than distal diameter.
Eyes large, darkly pigmented, located distally.
Neck robust, expanded distally, without
ornamentation. Proboscis typical, with swollen
medial section, with proximal and distal con-
strictions. Chelifores slender, lightly spinose,
-with double jointed-scape, first segment slightly
shorter than second, ornamented with one
dorso-median tubular spine, one dorsodistal, two
latero-distal tubular spines and two pointed
spines of similar length. Second segment
ornamented with 5-8 dorsal and lateral tubular
spines and 3-4 dorsal pointed spines. Chelae
reduced, distal end resembling club. Palps long,
slender, without tubular spines, consisting of 9
segments, with ventro-distal spines slightly longer
than their segment diameters. Ovigers typical,
strigilis segments denticulated, with many lateral
denticles. Two large and long setae located
laterally on seventh segment. Legs slender,
spinose, with large number of short and long
pointed spines and numerous tubular spines; first
coxae similar to anterior lateral process, with 3-4

dorsodistal tubular spines and 4-5 lateral and
ventral pointed spines; second coxa has 3
dorso-median short tubular spines and very small
ventro-distal spines; third coxae with 3 short
ventro-distal spine; femur slightly shorter than
subequal tibia; all leg segments armed with
randomly located tubular spines, longer pointed
spines dorsal ly and distally and short ventral
spines. Cement gland dorso-distal ly, tube of
slightly longer than segment diameter, carried
horizontally; tarsus short, propodus well curved,
with 4 heel spines and 6-7 sole spines, several
dorsal setae located randomly; claws half as long
as propodus, auxiliary claws long, roughly 0.8
claw length.

Female. Easily identified, with fewer tubular
spines on legs and lack of conspicuous femoral
cement gland. Females slightly larger, however,
possess smaller oviger than males. Juveniles
have fully functional chela, crossing at tips, with
simple teeth; chelifores slender and spinose.
Juveniles are also easily identifiable with oviger
buds.

Measurements . Holotype (in mm). Trunk length
(chelifore insertion to tip of fourth lateral
processes), 0.77; trunk width (across second
lateral processes) 0.67; proboscis length 0.6;
abdomen length 0.52; third leg, coxa 1 0.2; coxa II
0.3; coxa III 0.23; femur 0.57; tibia I 0.6; tibia II
0.58; tarsus 0. 1 ; propodus 0.38; claw 0. 1 6.

REMARKS. This new species is closely related
to two other species known from the Pacific: A.
tippula Child, 1 983 and A. paciftca Hilton, 1 942.
The primary difference between this new species
and A. tippula is seen in the tubular spines on the
lateral processes. A. tippula has low dorso-distal
knobs and short lateral spines on lateral pro-
cesses, lacking tubular spines on these regions. A.
fistella has three articulated short tubular spines
on the dorsodistal edge of the first three pairs of
lateral processes and a single tubular spine on the
last lateral process pair. A. paciftca , on the other
hand, lacks any form of tubular spines on its
lateral processes but possesses short postero-
lateral spines bearing lateral setules (/feathered
spines’) on the two anterior pairs and no spines on
the two posterior pairs. A. paciftca also lacks
spines of any form on the antero-lateral comers of
its trunk, while the same comers of A. tippula and
A. fistella each have a short tubular, inarticulate
spine.

The appendages of A. tippula and A. paciftca
have tubular spines that are uniformly longer
than those found on A. fistella. The first segment

NEW PYCNOGONIDA FROM TROPICAL QUEENSLAND

345

0.2 mm

FIG. 1. Ammothella fistella sp. nov. A,
dorsal view (ovigers omitted); B,
lateral view, showing curved
abdomen, palp with numerous
ventro-distal spines and oviger; C,
third leg, arrow indicating femoral
gland tube.

on the chelifore of these two species has 1 dorsal
tubular spine while the same segment on A.
fistella has 4. The second scape segment of A.
fistella has numerous short tubular spines, more
than any of the other two related species.

has fewer tubular spines and lacks spines or setae
on its lateral processes.

CALLIPALLENIDAE Hilton, 1942
Callipallene catulus sp. nov.

(Fig. 2)

Ammothella fistella is less closely related to its
nearest geographic neighbour, A. prolixa Child,
1990, collected from Orpheus Island, north of
Townsville, although they appear slightly
similar. Ammothella prolixa has a more stretched
or attenuated habitus with more slender
appendages than the other species discussed. It

-TYMOLOGY. Latin, catulus , young ot an animal;
Suggested by Dr Allan Child. It refers to the inflated,
uvenile appearance of the trunk.

VlATERIAL. Townsville, Rowes Bay, intertidal rocky
mnd, on the green alga, Cladophora pmlifcra (Koih) m
).5m, 1998, Id, holotype, (QMS105868) April-
siovember 1998, coll. Lee; Paratypes, 1 <J with eggs, 3 V

346

MEMOIRS OF THE QUEENSLAND MUSEUM

C

0.1mm

0.1 mm

B

0.1 mm

0.1 mm

FIG. 2. Callipallene catulus sp. nov. A, dorsal view,
B third leg, arrow indicating cement gland and
mid-femur inflation, inset showing enlarged femur
of ovigerous female with ova; C, ten-segmented
oviger, inset showing denticulated spine; D,
chelifores with one segmented scape, chela fingers
crossing at tips.

ovigerous, 2 juv. (QMS 105869); Paratypes 1 6 with eggs,
9 ovigerous, coll. Lee, Syntypes 26 , 2 9 (QMS 1 05870 ).

DISTRIBUTION. Known only from Rowes Bay,
Townsville, Queensland, in 0.5m tidal height.

DESCRIPTION. Body tiny, leg span 5.6mm.
Trunk without ornamentation, segments inflated,
first two segments fully segmented, third seg-
ment lacking complete segmentation lines. Lateral
processes glabrous, short, widely separated by
twice their diameter, no ornamentation. Abdomen
small, low, almost glabrous. Ocular tubercle low,
rounded dome, large eyes filling entire tubercle,
lightly pigmented. Neck distinct, moderately
long, very narrow proximally, expanded distally.

Proboscis short, semi globular, tapering to a
small oral surface. Chelifores moderately small,
scape one-segmented, about 2.5 X longer than
wide, armed with three or four distal and lateral
setae longer than or as long as scape diameters.
Chelae with similar dorsal setae, fully formed
with denticulate immovable finger with 6
well-formed teeth, movable finger without teeth.
Fingers cross at tips. Ovigers typical for the
genus, strigilis segments with rounded
denticulate spines in the formula 6:4:4:4,
terminal claw absent, fifth segment the longest,
with tiny dorsodistal tubercle having two short
setae. Legs long and slender, cuticle transparent;
few setae, all shorter than segment diameter

NEW PYCNOGONIDA FROM TROPICAL QUEENSLAND

347

except dorsodistal seta; tarsus very short, with
one heel spine and 2-3 short setae; propodus
slender, moderately curved with 4 heel spines and
10-12 shorter sole spines and setae; main claw
long, about 2/3 propodal length; auxiliary claws
long almost as long as dactyl. Single cement
gland, extruding ventrally through a single pore
causing slight inflation at mid-femur.

Female. Slightly larger, neck slightly longer than
male, with latero-distal apophysis on fifth
segment of oviger lacking in female, as typical.
Legs are similar except in ovigerous females
where the femora are enlarged with ova in
various stages of development.

Measurements. Holotype (in mm). Trunk length
(chelifore insertion to tip of fourth lateral pro-
cesses) 0.64; trunk width (across second lateral
processes) 0.27; proboscis length 0.21 ; abdomen
length 0.15; third leg, coxa I 0.09; coxa II coxa
III, 0.13; femur 0.64; tibia I 0.55; tibia II 0.65;
tarsus 0.05; propodus 0.25; claw 0. 13.

REMARKS. This new species is rather similar to
Callipallene tridens Nakamura & Child, 1988.
These similarities are seen primarily in the
inflated posterior trunk segments of each species.
In both, the ocular tubercle is placed on a slight
elevation and the segments are sutured the same
way, and the third and fourth segments lacking
the posterior sutures. Many species of this genus
inflated posterior trunk segments but it is usually
seen in juveniles, while the adult trunk segments
are more slender. Many also lack the third trunk
suture, as do C. tridens and C. catulus. The main
differences between C. catulus and C. tridens lie
in the lack of the specialised trident-shaped auxil-
iary claws of C. tridens. Callipallene catulus also
has longer and more slender appendages and
different chelae, which serve to distinguish the
two species. Callipallene catulus is extremely
tiny, almost smaller than any Callipallene species
described, except C. tridens , which is possibly
the smallest known species at 3.5mm in leg span
(Nakamura & Child, 1988).

PHOXICHILIDIIDAE Sars, 1891

Anoplodactylus gland u lifer Stock, 1954

Anoplodacty'lus spec.. Caiman, 1923: 289.

Anoplodactylus gl an du lifer, Stock, 1954: 16-17. Child 1982:

273-274; 1988: 58-59. Nakamura & Child, 1988: 813.

Muller, 1990: 74. Child 1990: 331. Muller, 1992: 166, figs

27-30. Stock 1992: 94-95.Bamber, 1992: 193-194. Stock

1994: 59.

Anoplodactylus multiclavus Child, 1977: 593-596, fig. 4;

1979: 58, fig 19d; 1982: 272. Muller, 1992: 166.

MATERIAL. Rowes Bay, rocky band on algae
Cladophora prolifera (Roth) in 0.5m, April-November
1998, 1<3 with eggs, 29 (QMS 105873); 3d, 39.

DISTRIBUTION. This species is fairly widely
distributed throughout the Indo-West Pacific,
from the Red Sea to Kenya, Singapore, Hong
Kong, the Marshall Islands, the Great Barrier
Reef and also the Caribbean Sea (where it was
described as A. multiclavus).

REMARKS. This species is one of a group of
over 20 in this genus with multiple femoral
cement gland openings (Child, 1988: 293-294,
for a partial key). Femora have from 2-5 tiny cups
located dorsally, with the numbers varying
among legs of the same male specimen and also
among specimens of a single population. Other-
wise, this species is quite robust, with lateral
processes closely set. Both chelae lingers are
armed with teeth, and the movable finger is
longer than its partner. Hints ol palp buds on the
anterior side of the first lateral processes and a
long propodal lamina are characteristic of the
species.

Anoplodactylus cribcllatus Caiman, 1923

Anoplodactylus cribellatus Caiman, 1923: 285. - CAir
Anoplodactylus simplex Clark, 1963: 50-51, fig. *-5A-r.

Stock, 1979: 158. Staples, 1982: 461, fig. 2A-B, pi. 1. figs

C-D. Bamber. 1997; 46-49, fig. 2.

Anoplodacty'lus perforatus , Nakamura & Child, 1983.289,

1988:813,1991:29. Bamber, 1997: 46-49, fig. 2.

MATERIAL. Cleveland Bay, Townsville, dredge sample,
3-1 5m depth, sandy-muddy bottom, 1998 2 9,
(QMS 105872) coll. J. Cruz.

DISTRIBUTION. The individuals here collected
would have been reported as A. simplex for which
there were only two previous records from the
east coast of Australia (Clark, 1963; Staples,
1982; but see Arango, 2003). However, Bambei
(1997) examined the Indo-west Pacific Anoplo-
dactylus species with very numerous cement
gland pores concluding the synonymy ol three
species: A. cribellatus from
Islands, A. perforatus Nakamura & Child. 1983
from Japan and A. simplex from Australia. All
specimens found in rather shallow waters.

REMARKS. The specimens collected here agree
with the description of A. simplex syn. Clark
showing 22-30 cement gland pores, a short
abdomen and elongate propodus without a
typical propodal lamina along its sole. After
Bamber’s examination of three similar species in
which the number of cement glands pores overlap
but the structures of the legs and ovigers are

348

MEMOIRS OF THE QUEENSLAND MUSEUM

identical among them (Bamber, 1997) it is
reasonable to report these specimens as A. cribellatus
and not A. simplex.

Anoplodactylus tubiferus (Haswell)

Phoxixhilidium tubiferum Haswell, 1884: 1032, pi. 57, figs
1-5.

Anoplodactylus tubiferus Cole, 1904: 288. Flynn, 1919:
79-81, pi. xx., figs 12-14, pi. xxi., fig. 15. Nakamura &
Child, 1991: 32. Stock, 1994: 67.

MATERIAL. Cleveland Bay, Townsville, dredge sample.
3- 1 5m depth, sandy-muddy bottom, 4 6 , 1 9 , (QMS 1 0587 1 )
coll. J.Cruz.

DISTRIBUTION. This species was first
described in 1884 and has been collected from
Madagascar and the Persian Gulf to Australia,
Philippines and Japan. It is known to inhabit
different depth strata, from 2-235m.

REMARKS. The extremely long and conspicuous
cement gland makes the males of this species
easily identifiable. It has a single cement gland
tube more than twice as long as the femur,
protruding almost vertically. This species has a
very tall ocular tubercle and abdomen, very long
distal spines on the lateral processes, and
sometimes dorsally on the trunk. These features
facilitate the identification of A. tubiferus
(Staples, 1982: 457-459, fig. 2C). The species
shows a general elongate habitus shared with
other species from the western Pacific. A key for
this species can be found in Nakamura & Child
(1983:41-42).

ACKNOWLEDGEMENTS

This short report is dedicated to Dr C.A. Child
who helped us enormously with the taxonomic
identification and the production of earlier
versions of this manuscript. Thanks to J.J. Cruz
for the collection of specimens ffom Cleveland
Bay. This study was funded by James Cook
University, Queensland, Australia through the
IRA grants.

LITERATURE CITED

ARANGO, C.P. 2003. Sea spiders (Pycnogonida, Arthro-
poda) from the Great Barrier Reef, Australia: new
species, new records and ecological annotations.
Journal of Natural History (posted on-line as
preview article on December 2002), 1-50 p.
BAMBER, R.N. 1992. Some pycnogonids from the
South China Sea. Asian Marine Biology 9:
103-203.

1997. Two pycnogonids from the Coral Sea region:
a new Callipallene and a reassessment of the
Anoplodactylus cribellatus complex. Species
diversity 2: 43-50.

CHILD, C.A. 1 975. Pycnogonida of Western Australia.
Smithsonian Contributions to Zoology 1 90: 1 -29.
1979. Shallow- Water Pycnogonida of the Isthmus
of Panama and the coast of middle America.
Smithsonian Contributions to Zoology 293: 1 -86.
1982. Deep-Sea Pycnogonida from the North and
South Atlantic Basin. Smithsonian Contrib-
utions to Zoology 349: 1-54.

1988. Pycnogonida of the western Pacific Islands
III. Recent Smithsonian-Philippine Expedition.
Smithsonian Contributions to Zoology 468: 1 -32

1990. Pycnogonida of the western Pacific Islands
VIII. Recent-collections from islands of the
Great Barrier Reef, Australia. Proceedings of the
Biological Society of Washington 103:31 1-335.

CLARK, W.C. 1963. Australian Pycnogonida. Records
of the Australian Museum 26: 1-81.

1970. New Pycnogonida from Queensland.
Transactions of the Royal Society of New
Zealand, Biological Sciences 12: 13-20.
HASWELL, W.A. 1884. On the Pycnogonida of the
Australian coast with descriptions of new species.
1021-1034.

NAKAMURA, K. & CHILD, A. 1983. Shallow-water
Pycnogonida from the Izu Peninsula, Japan.
Smithsonian Contributions to Zoology 386:1-71.

1991. Pycnogonida from waters adjacent to Japan.
Smithsonian Contributions to Zoology 512:1 -74.

STAPLES, D. 1 982. Pycnogonida of the Calliope River
and Auckland Creek, Queensland. Memoirs of the
Queensland Museum 20: 455-71.

THE GREEN TURTLE, CHELONIA MYDAS , POPULATION OF RAINE ISLAND AND
THE NORTHERN GREAT BARRIER REEF: 1843-2001

COLIN J. LIMPUS, JEFFREY D. MILLER. C. JOHN PARMENTER AND DUNCAN J. LIMPUS

Limpus, C.J.. Miller, J.D., Parmenter, CJ. & Limpus, D.J. 2003 06 30: The Green Turtle,
Chelonia my das, population of Raine Island and the northern Great Barrier Reef: 1 843-200 1 .
Memoirs of the Queensland Museum 49(1): 349-440. Brisbane. ISSN 0079-8835.

European history of the use of Raine Island and nesting green turtles is reviewed. Results of a
diverse mosaic of investigations of the reproductive biology of the nesting population of
green turtles using Raine Island and the adjacent islands are presented. These studies have
focused on defining the fundamental characteristics of the breeding population, with more
in-depth studies being conducted elsewhere. Studies at Raine Island have defined the species
composition, morphology of adults, eggs and hatchlings, hatching and emergence success,
sources of mortality of adults, eggs and hatchlings as well as the nesting environment. □
Green turtle, Raine Island, population history, Great Barrier Reef

Colin J. Limpus , Jeffrey ’ D. Miller, Duncan J. Limpus, Queensland Parks and Wildlife
Service, PO Box 155, Albert Street, Brisbane, 4002; C. John Parmenter, School of Biological
Sciences, Central Queensland University \ Rockhampton, 4702, Australia; 20 April 2000.

There arc four major green turtle, Chelonia
my das, breeding concentrations in Australia:
Capricorn - Bunker Groups and adjacent islands
in the southern Great Barrier Reef (GBR); Raine
Island - Moulter Cay and adjacent islands in the
northern GBR; Wellesley Group - eastern
Arnhem Land in the Gulf of Carpentaria; and, the
northwest shelf of Western Australia. These
appear to be the only major rookery areas for the
Australasian region where significant population
declines are not already in progress for this
species (Groombridge & Luxmoore, 1989).
Raine Island, Moulter Cay and the other sand
cays of the northern GBR collectively support the
largest remaining breeding assemblage of green
turtles in the world.

While world-wide there were once many other
C. mydas rookeries of the magnitude of Raine
Island - Moulter Cay, there may now only be a
few: Tortuguerro rookery of Caribbean Costa
Rica; Ras al Had in Oman; Northwest Shelf of
Western Australia. Australia and, in particular the
GBR. is one of the last strongholds for a once
nourishing species worldwider

Within a few years of commencement of our
tagging at Raine Island it became apparent that
many of the turtles were migrating from feeding
areas outside of Australia. In addition, large
numbers of C. mydas were being hunted for food,
both at the village level and for commercial
market, within the feeding areas supplying turtles
to the Australian rookeries. So, while there has
been almost total protection of the nesting
females at the GBR rookeries for many years, the

population is under a large but imprecisely
determined harvest pressure. In many areas this
harvest targets large females but in Torres Strait
gravid females migrating to breed are preferred
where they are available. The pattern that has
developed' this century with turtle harvests in
other countries is that marine turtle population
numbers will crash following several decades of
intense harvesting. Given that the large harvests
that target the Raine Island turtles have only been
operating in the past 2 or 3 decades, concern must
be held for the future stability of this population.
Improvements in our basic understanding of
turtle biology and significant improvements in
marine turtle management are surely needed in
these times of declining turtle populations in
Southeast Asia and the Western Pacific Region
(Limpus, 1 997). This is particularly so, given that
sustainable management of marine turtle
populations subjected to large scale harvests has
not been achieved in the 300 years since the
elements of our modem understandings of green
turtle biology were first described by William
Dampier in 1717 (Masefield, 1906).

The turtle research at Raine Island in the years
1975-1979 was initially part of exploratory
studies by the Queensland Government
Conservation Agency (currently called
Queensland Parks and Wildlife Service [QPWS])
to map the species composition, size and
distribution of marine turtle breeding populations
in Queensland. At that time it became apparent
that there was a lack of fundamental information
required to effectively manage a marine turtle

350

MEMOIRS OF THE QUEENSLAND MUSEUM

population in the face of large scale harvesting
and guarantee its long term continuity. During the
period 1979-1985, the Queensland Turtle
Research Project undertook the development of
new methodologies to address marine turtle
population dynamics. These included: tagging
methodology to enable turtles to be recognised
over decades rather than a few years (Limpus,
1992a); capturing techniques for studying the
turtles in their home feeding areas (Limpus &
Read, 1985a); methodologies for sexing and
assessing the reproductive status of turtles in their
feeding areas (Limpus, 1 992b); genetics research
to determine methods of recognising the dis-
tribution and number of genetically discrete
breeding units within our C. mydas herds
(Norman et al., 1994a); and mark-recapture
studies to provide the quantified life table
parameters including fecundity, growth data,
survivorship and recruitment, (Limpus &
Chaloupka, 1997; Chaloupka & Limpus, 2002)
that are necessary for developing good models
for describing marine turtle population dynamics
(Chaloupka & Limpus, 1 996; Chaloupka, 2002).

This latter issue of understanding population
dynamics is vital if we are to manage our turtle
herds on a sustainable basis in the face of large
scale mortality from harvest and other sources.

Recognising the potential threats to the con-
servation of the GBR C. mydas breeding
populations posed by large scale harvesting
within the Australasian region, QPWS placed a
priority on studies of C. mydas in 1985. An
integrated set of field studies was developed,
bringing together a team of researchers with
diverse skills, to investigate the population
dynamics of eastern Australian green turtles. The
study commenced in 1986 and became fully
operational in 1987. At the core of the study has
been the simultaneous study of C. mydas at each
of its three eastern Australian rookery regions
and at a series of feeding areas distributed
throughout eastern Australia.

For logistical considerations. Heron Island
within the southern GBR nesting area was
selected as the primary study site for in-depth
research of the reproductive biology of C. mydas.
Access to the major nesting beaches of the
northern GBR has been more difficult because of
the remoteness of Raine Island and Moulter Cay
and large expense associated with maintaining
on-site studies there. Therefore, studies at Raine
Island were planned as short term sampling
studies rather than total season tagging census

studies with an emphasis on quantifying those
parameters considered to have unique values for
Raine Island. It was also recognised that results
of past intense studies from Bramble Cay in the
extreme northern GBR would supplement results
for the Raine Island area.

It is in the context of this complex mosaic of
diverse, yet complementary investigations, that
the present study of the C. mydas nesting
population at Raine Island and adjacent rookeries
is presented. It is part of a planned, long-term,
integrated regional study of green turtles in the
Australasian region designed to provide sound
information that can assist in improving the
conservation management of C. mydas.

METHODS

Data for this study was drawn from the
historical and published accounts of Raine
Island, from the results of interviews with
fishermen and others who visited the island in
recent years and from field studies undertaken at
Raine Island in the period 1 974 to February 2002.
While the principal emphasis will be on Raine
Island and its turtle population, the green turtle
populations at Moulter Cay and the other islands
in the northern Great Barrier Reef will also be
considered.

SPECIES IDENTIFICATION. Turtle species
identification was made from the available keys
(Bustard, 1972 and Cogger & Lindner, 1969).
These data have been modified into the field key
used in the QPWS Queensland Turtle Research
(QTR) Project (Limpus, 1992c).

TAGGING. Table 1 summarises the types of tags
and their positions of application used in the
present study. All tags were self-piercing
self-locking flipper tags, serially numbered with
a return address on the opposite side to the
number. Monel tags: National Band and Tag
Company, Kentucky; style 1005, #49 and #19;
weight = 7g. Three series of monel tags were
used, each being identified by the prefix to the tag
number (Table 1). Titanium tags: Stockbrands
Company Pty Ltd, Western Australia; #standard;
weight = 4g, commercially pure titanium, grade
125, 1mm thick. These titanium tags have a
locking mechanism resembling that of the monel
tag style 1 005 # 1 9. Return address was to Queens-
land Parks and Wildlife Service. All titanium tags
carry a T, I or K prefix to the tag number.

Tags have been applied to various positions on
the trailing edge of the front flippers. Standard
tagging positions (Limpus, 1992a) have been

GREEN TURTLE POPULATIONS OF RAINE ISLAND

351

TABLE 1 . Summary of tag designs, return addresses,
tagging positions and the number of tags applied per
turtle. See Limpus (1992a) for further definitions. *
Recaptured turtles from previous seasons or other
studies were double tagged L3 and R3.

Tag series

Metal type
or style

No. tags per
female

Tagging

position

Return

address

No prefix

Monel #49

1

LI

Applied

Ecology

Pty Ltd

A prefix

Monel #49

2

LI & R1

Applied

Ecology

Pty Ltd

lx prefix

Under

X35000

Above

X35000

Monel #49

Monel #19

1

1

1*

LI prior to

1978

L3 alter
July 1978

L3 after

1979

QPWS

T prefix,

I prefix,

K prefix

Titanium

1*

L3

QPWS

coded as follows: L = left front flipper; R = right
front flipper; 1 = tag applied immediately distal to
phalange 5 (after Bustard, 1968); 3 = tag applied
through or adjacent to the most proximal axillary
scale (= axillary tagging scale, Limpus & Reed,
1985a). Thus a tag can be described as having
being applied at the LI, L3, R1 or R3 position.

COUNTING TURTLES. Counts or indices of the
number of turtles ashore per night were
established in one of the following ways:

a) Tally count. An instantaneous count of all
turtles was conducted by several persons walking
line abreast across the beach, for one circuit of the
island between 2000-2300hr (a tally count
usually took 40-60min. to complete). Counts
commenced at least one hour after turtles had
swimming access to the nesting beach across the
reef Hat after dark. During the summer nesting
season the tides are such that on most of the
nights, the majority of the turtles for the night will
have beached within a few hours of darkness. A
green turtle rarely lays and returns to the water in
less than 3hr on the beach.

b) Tagging census. A count of all turtles ashore
for the night based on tagging records. This was
only attempted during periods of low density
nesting (<400 females ashore per night).

c) Track count. A count of all turtle tracks for the
night by species, based on a count just after dawn
of the tracks that start below the previous nights
high tide mark. Each track equals an ‘emergence’

and a ‘return’ component. This method has only
been applicable on nights with a low density of
turtles. At Raine Island, if there is an error it will
usually involve some under-estimation. At
localities visited for a single day only, tracks
accumulated over several nights may be
indistinguishable, resulting in the potential for
overestimates.

d) Stepping index. Commencing with the Raine
Island during the 1987 summer expedition, the
opportunity was taken to evaluate nesting density
equivalents to old reports from fishermen who
spoke of having walked across the backs of many
nesting turtles without having to step down onto
the sand. When dense aggregations of nesting
turtles were encountered on Raine Island beach
between 2 1 00-2400hrs, an agile average sized
person stepped from turtle to turtle for as many
steps as possible without stepping onto the beach.
The number of turtles stepped on is defined as the
stepping index of the nesting density. This was
repeated at several places around the island on
each night. It was not attempted on nights with
rain when the carapaces were slippery.

MEASUREMENTS OF TURTLES, EGGS
AND NESTS. The adult turtles were measured as
follows. Curved carapace measurements were
taken with flexible fibre-glass tape measure
(±0.5cm prior to 1983; ±0.1 cm thereafter)
stretched tightly from edge to edge. Straight
carapace measurements were taken with a steel
tape measure (±0.1 cm) and large wooden
dividers with recurved ends. Measurements of
standard length were made along the midline
from the anterior skin-carapace junction to the
apex of the posterior midline notch between the
supracaudal scutes (curved and straight carapace
length = CCL and SCL, respectively). Width was
measured at the widest part of the carapace
(curved and straight carapace width - CC W and
SCW, respectively). These measurements
correspond to those used in other major studies of
marine turtles in Australia (Limpus, 1 ,

Bustard, 1972; Limpus etal., 1983a,b 1984a) To
allow for comparison with turtle studies in other
areas, adults were also measured for the standard
length measurement used by Professor Archie
Carr, i.e. from the most anterior projection of the
carapace over the shoulder to the tip of the most
posterior projection of the carapace (= C arr s
straight carapace length [CSCL] [Carr and Ogren
I960]: this is not a midline measurement). Adult
turtles were weighed on Salter dial balances
(±0.5kg). Females were weighed after completion
of oviposition.

352

MEMOIRS OF THE QUEENSLAND MUSEUM

Clutches were counted and eggs measured
within 2 hours of being laid and with a minimum
of rotation and vibration of the eggs (Limpus et
al., 1979; Parmenter, 1980a). Eggs and hatch-
lings were measured for mean diameter and
straight carapace length, respectively, with
vernier slide callipers (±0.0 1 cm) and weighed on
Pesola spring balances (±0.5g). No more than 10
eggs or hatchlings were measured per clutch. The
emergence success of a clutch was assessed by
excavation of the nest and recording the fate of
the eggs by six classes: undeveloped = egg
contained no obvious embryo (< Stage 17 of
Miller, 1985); unhatched = egg contained a
recognisable embryo (> stage 17 of Miller, 1985);
predated = egg broken by crab; dead in nest =
hatchling died after hatching, but below the
beach surface; live in nest = hatchling was alive,
but had not emerged from the nest (groups of live
hatchlings just below the surface and not tangled
in vegetation were counted as having emerged);
emerged = hatchlings which successfully
hatched and escaped from the nest to the beach
surface (Limpus et al., 1983a,b).

Nest depths were measured with a fibreglass
tape measure (±0.5cm) from the beach surface to
the top of the upper most egg and to the bottom of
the egg chamber (sinsu Bustard & Greenham,
1969 ).

Sand temperature was measured with a
mercury-in-glass thermometer (±0.5°C) inserted
horizontally into the sand 50cm below the
surface. Core body temperatures of nesting
turtles were measured by the Mrosovsky &
Pritchard ( 1971 ) method. This required catching
an egg immediately as it was oviposited and
. inserting a quick reading mercury-in-glass
thermometer into the middle of the yolk.

NESTING SUCCESS AND CLUTCH
DESTRUCTION. Two methods were used for
estimating nesting success, here defined as the
proportion of females ashore for the night that lay
a clutch.

a) Following individual turtles. Nesting turtles
were selected at random as they came ashore on
the western end of the island at night and were
followed until they returned to the sea. Their
nesting success was recorded. This method of
measurement was time consuming, as it required
a person committed to maintain observation of an
individual turtle, irrespective of how long it was
ashore. Usually only one turtle could be followed
on any one night. Each turtle had to be followed at

such a distance that she did not respond to the
observer's presence.

b) Using fixed sampling sites. Standard study
sites were chosen on opposite sides of the island.
Unless otherwise stated, each site was 100m in
length and extended from the water line to the
cliff line. On selected nights, within one or both
of these sampling sites, a count was made of the
number of turtles which entered the sampling site
from across the waterline, the number of clutches
laid within the area by these turtles, and the
number of existing clutches that were dug into by
the nesting turtles. Each data set required one
person monitoring a sampling site for approx-
imately 14hr continuously commencing at about
1600hr. Two rate parameters can be calculated
from these data: nesting success (NS) and clutch
disturbance (CD) by nesting turtles.

NS = Number of clutches laid / total number of
turtles entering area

Nesting success was calculated on the number
of turtles crossing the high water line on the
assumption that lateral movements into and out
of the sampling sites along the beach by turtles
attempting to nest are approximately random and
will cancel each other.

CD = Number of existing clutches disturbed /
Number of new clutches laid.

When a turtle was found digging into an
existing clutch, irrespective of where it was on
the island, the number of eggs dug from the nest
was counted. These counts provide a measure of
the egg mortality per clutch disturbed (EMpDC)
by nesting females. When compared to the mean
clutch count for the rookery, this egg destruction
can be expressed as the proportion of a clutch
destroyed per clutch laid (clutch equivalent egg
mortality = CEEM) CEEM = EMpDC x CD /
mean clutch count.

ESTIMATION OF THE SIZE OF THE
ANNUAL NESTING POPULATION.

Petersen Estimate. If a sample of the turtles
ashore on Raine Island at night is marked so that
they can be recognised in the water over the
following days, and if the marked turtles are
uniformly distributed among the unmarked
turtles in the waters surrounding Raine Island,
then mark-recapture data can be analysed using
the Petersen estimate with Bailey’s correction
(Caughley, 1977) to provide an estimate of the
number of adult green turtles in the waters
surrounding Raine Island at that time.

In a population where M animals are marked

GREEN TURTLE POPULATIONS OF RAINE ISLAND

353

out of a total population of N animals and m
marked animals are recaptured in a subsequent
sampling of n animals:

N = M(n+l)/(m+l)

This estimate has a formal standard error of
approximately:

SE = [M 2 (n+1 )(n-m) / (m+1 ) 2 (m+2)] l/2

The principal assumptions made in using this
method are: 1) the marked turtles are uniformly
distributed among the unmarked turtles that form
the Raine Island breeding assemblage within the
intemesting habitat on that day; 2) there is a
negligible rate of loss of the mark between
marking and recapture; 3) the marks are easily
seen on the swimming turtle; 4) all the females
aggregated on Raine Island Reef are there to
breed i.e. there are insignificant resident
non-breeding female and immature male
populations on Raine Island Reef; 5) there is
equal probability of the tagged and untagged
turtles moving outside the sampling area; and 6)
there is negligible mortality of the adult turtles
during the sampling period.

Two methods have been used to mark the Raine
Island nesting turtles for mark recapture studies.

1) Standard turtle tags. Within a study trip, after
several nights of tagging the nesting females on
the beach, a search of sections of the surrounding
Raine Island Reef was made to locate, count and
visually check females for the presence or
absence of a tag. Only those females checked
were counted. Those seen in the distance but not
checked were excluded from calculations. Monel
tags applied to the LI or R1 tagging positions
(towards the distal ends of the front flippers) were
readily visible on a swimming turtle. In 1979 the
place of application of the metal tags was
changed to the axillary area of the trailing edge of
the front flippers (L3 or R3 tagging positions) to
improve tag retention. This "axillary tag
placement resulted in a reduced visibility of the
metal tag on the swimming turtle, especially for
those not viewed at very close range. The poor
retention rate of monel tags (Limpus, 1992a)
made it unlikely that there was a significant
proportion of remigrant turtles that would be
counted in the intemesting habitat as part of n but
which were not part of M.

2) Paint marks on the carapace. Commercially
available paint was use to mark the dry carapace
of nesting turtles on Raine Island at night. Paint
marks were at least 10cm long, applied by a
2.5cm wide brush or by a pressure pack can, were
applied to the dry carapace of nesting turtles on

Raine Island at night. Oil based enamel paints
were used initially. Commencing in 1995, the
paint used to mark the turtles was changed to
white Wattyl Airpave™ (504-40100.
BN:-45 1695-09) paint applied as single stripes
with 5.0cm wide brushes. On the first night of
painting, the stripes were longitudinal along the
midline, on the posterior half of the carapace. If
turtles were painted on a second night in the same
trip, those marks were transverse stripes on the
posterior half of the carapace. A turtle was
selected for painting if: the carapace was dry; the
carapace did not have a thick coating of algae;
and the turtle was inland of the beach crest, and
well within the nesting habitat. When applied in
these conditions, the paint was observed to
adhere well to the carapace surface for at least
48hr (enamel paint) or 96hr ( Airpave). Using this
marking system, the goal was to paint-mark as
many turtles as possible on the island in a single
night. A search was made for marked turtles
among the turtles in the waters surrounding the
island on subsequent days. Turtles seen were
recorded by species and tail length. Those with
long tails were scored as male, adult-sized short
tailed turtles were scored as presumed adult
female and those smaller than adult female (CCL
<90cm for C. my das; CCL <75cm for E.
imbricata) were scored as immature (Limpus &
Reed, 1985a; Limpus, 1992b). Counts were made
of those females for which the presence/absence
of the tag/paint mark could be definitely established.

Two types of in-water searches were made for
the turtles.

1 ) Diving transects. Transects using SCUBA, or
snorkel, were swum along the reef edge above the
10m depth terrace. Written records were kept on
waterproof paper. During a diving transect, two
persons were in the water, one counting and one
on the lookout for sharks. The lookout earned a
hand spear for warding off sharks and the
recorder carried a line attached to a buoy at the
surface. A third person maintained a dinghy
adjacent to the buoy for rapid retrieval of the
divers from the water, if required.

2) Speedboat transects. On days with clear,
relatively calm water, transects were driven
across the reef flat at the upper part of the tidal
cycle and along the outer edge of the reef in
search of turtles, using an outboard powered
dinghy. By driving adjacent to the swimming
turtle, it was possible to score it by species/
size/sex/mark presence. A minimum of two
persons were required in the speedboat, one
recording and the other driving and counting. The

354

MEMOIRS OF THE QUEENSLAND MUSEUM

acceleration capability of the speedboat was
vital. The boat had to be able to accelerate quickly
to reach the turtle before its swimming disturbed
others nearby, or before it escaped to deep water.
Boats about 4-5m in length, powered with a
40-60HP outboard motor were best suited to this
use in shallow water. Conventional 3.5m
aluminium dinghies and inflatable dinghies fitted
with 10-15HP outboard motors proved unsuited
to the task.

In high density nesting seasons, it was im-
possible to obtain an absolute measure of the
nightly number of females emerging on Raine
Island, so the mean nightly tally count has been
used in lieu of the number of turtles emerging per
night. This under-estimates the actual number of
turtles emerging nightly.

MORTALITY STUDIES. The number of dead
and moribund turtles at each studied rookery was
recorded on arrival. Subsequent adult mortality,
particularly on Raine Island, was recorded daily.
Previously encountered carcasses were identified
by flagging them with coloured surveying tape or
by paint marks. Moribund turtles were rescued
where possible, especially those that had fallen
onto their backs. These rescued turtles were counted
as mortalities for the purposes of the study.

In December 1 989, a group of turtles that had
remained on the beach until at least mid-morning,
and were in a location where all such turtles died
naturally of heat exhaustion were selected for
detailed study. Changes in core body temp-
eratures of these moribund turtles was assessed
by inserting a cannula through the inguinal
pocket anteriodorsally, to the movement arch of
the hind leg (as is done during laparoscopic
examination of turtles). The cannula provided
access to the abdominal cavity and allowed a
thermometer probe, attached to a 50cm rod, to be
inserted to lie above the gut and ventral to the
lungs just posterior to the heart. The lead and
readout unit were taped to the carapace and the
turtles were allowed to move undisturbed on the
beach until they died. Temperatures were read at
half hourly intervals. Necropsies of these turtles
confirmed the correct placement of the probes,
and that no damage had been done to organs or
blood vessels. The insertion of the probes did not
contribute to the death of the turtles.

The daily census of dead and moribund turtles
was conducted between 1500 and 1900hr. On
occasion, freshly dead turtles were necropsied to
assess oviducal contents and ovarian condition.

TABLE 2. Circumference of sand islands used by
green turtles for nesting in the Far Northern Great
Barrier reef. Circumference was measured at
approximately mean high water spring tide level
using a pedometer.

Island

Date

Circumference

Raine Island

1977 Dec

1799m

1978 Dec

1764m

1980 Jun

1808m

Moulter Cay

1976 Dec

1179m

1977 Dec

1138m

Maclennan Cay

1978 Nov

885m

No. 7 Sandbank

1976 Nov

1755m

No. 8 Sandbank

1976 Nov

1816m

Bramble Cay

1977 Dec

748m

In December 1989 the core body temperature of
some dead turtles was measured during necroscopy.

RECRUITMENT RATE STUDIES. We have
introduced a new method for marine turtle studies
for quantifying recruitment of new adults into the
breeding population. Traditionally, recruitment
of new adults into the breeding population has
used long term total tagging census to identify all
turtles with a past breeding history. After a total
tagging census at a nesting beach that spans
several mean remigration intervals, turtles that
have not been previously tagged are identified as
new recruit (Hughes, 1989; Richardson &
Richardson, 1982; Limpus, 1985; Limpus et al.,
1984b; Parmenter & Limpus, 1995). This has
been an inappropriate method for the Raine
Island population given the difficulties in tagging
the entire nesting population, even in a single
year. Within foraging area studies in eastern
Australia, marine turtles have been assessed for
recruiting into the adult breeding population
using ovarian features, in particular scoring for
the presence or absence of corpora albicantia. A
corpus albicantium is a healed corpus luteum and
a corpus luteum is formed by the release
(ovulation) of a mature follicle from the ovary.
The presence of corpora albicantia indicates that
a female has bred in a past nesting season.
Similarly the absence of corpora albicantia on the
ovaries of a female with oviducal eggs indicates
that she has not bred in a past breeding season and
that she is in her first breeding season. Therefore
the absence of corpora albicantia in the ovaries of
females that have recently ovulated can be used
to identify those females that are new recruits to
the breeding population (Limpus & Limpus, 2003).

Ovaries were examined by direct examination
via necropsy of nesting females that had freshly

GREEN TURTLE POPULATIONS OF RAINE ISLAND

355

died on the nesting beach or by endoscopy
(Limpus & Reed, 1985b; Limpus et al., 1994c).
For endoscopy of the breeding female, it was
necessary to examine females ashore as they
completed the laying of a clutch. Otherwise the
presence of oviducal eggs made it very difficult
to view the surface of the ovary clearly. Limpus &
Limpus (2003) provide a key for scoring sex,
maturity, breeding status and recruitment from
gonad examination.

GENETIC STUDIES. Tissue samples were
collected for genetic research in a variety of
ways. Non-sibling hatchlings were sampled by
collecting one hatchling per clutch from clutches
which emerged within a two week period at any
given rookery (Norman et al., 1994a). This
ensured that no more than a single hatchling was
sampled from any one female within the breeding
season. To ensure that individual adult turtles
were sampled only once, adult turtles with unique
tag numbers were sampled for either muscle,
blood or skin (Norman et al., 1 994b; FitzSimmons
et al., 1 995). The methods for genetic analysis are
described in these separate papers.

RESTRICTIONS ON STUDIES AT RAINE
ISLAND. Raine Island and the adjacent islands
have proven to be logistically difficult for long
term and exrended period study due to
remoteness. The number of personnel that can be
deployed on the island for the turtle studies has
been limited because of the need to minimise
disturbance to the nesting colonies of sea birds
that Raine Island supports.

The first trips to Raine Island for this study
were opportunist. The Queensland Parks and
Wildlife Service (QPWS) study was supported in
part by research grants from the Australian
National Parks and Wildlife Service during
1976-1979. In 1977 and 1978 the expeditions to
Raine Island were supported by Applied Ecology
Pty Ltd (AE) who provided a vessel to visit the
island. In 1979 with the aid of a WWF grant a
study of seas birds at Raine Island was com-
menced and one turtle researcher was able to
accompany the bird study group. In 1981 the
Raine Island Corporation became responsible for
the conservation of Raine Island and commenced
funding research expeditions to Raine Island.
From 1981 to 1985 the Raine Island Corporation
chose to place an emphasis on sea bird studies
and the turtle research team was limited to one
person on the island in any one trip except during
December 1 984, when 2 persons were present for
the turtle studies. The sea bird research team

assisted with turtle tally counts and incidentally
with the other aspects of the study. Since 1 985 the
Raine Island Corporation has reduced its
emphasis on sea bird studies at Raine Island and
has funded the summer expeditions primarily for
the turtle research with teams of 4-5 persons per
trip. To maximise the data gathered on a trip,
small numbers of other islands were surveyed for
marine turtle nesting during transit to and from
raine Island. Thus from 1975-1989, the majority
of the islands between Princess Charlotte Bay
and Cape York were surveyed at least once.
These data provided a measure of the regional
distribution of marine turtle nesting by species.

CLIMATE DATA. Regional rainfall and
temperature data from the three long-term Coral
Sea weather stations nearest to Raine Island
(Torres Strait, Willis Island and Lockhardt River)
were obtained from the Australian Bureau of
Meteorology.

STUDY AREA

RAINE ISLAND. Raine Island (11°36 , S,
144°01’E) is a coral cay on the leeward north-
western end of a small detached reef on the outer
edge of the northern GBR (Fig. 1 ). The island is
approximately 80km offshore lrom the mainland,
north east of Cape Grenville. Raine Island Reef
lies in the eastern approaches at the Raine Island
Entrance, a deep water passage through the outer
barrier reef. The reef rises abruptly out of deep
water with near precipitous cliffs to a narrow
terrace at approximately 8m depth and is topped
by an approximately uniform height reef flat that
dries at the lower tidal ranges.

Raine Island, as described in recent times
(Warham, 1977; Stoddart et al., 1981; King,
1986), approximates to the island as it was
described in 1843 (Jukes, 1847). The island is
approximately 1800m in circumference at the
mean high water spring (H WS) tide level (Fig. 2,
Table 2). The beach substrate is a white calcium
carbonate sand composed of mostly fragmented
coral and mollusc and foramimferan skeletons (
star sand). Turtle and sea bird bones areobvious
but at low density within this sand. I he beach
width measured from the high tide line to the low
cliff' line of the central rock platform varies from
approximately 15m near the beacon to 90m on
the southern side. Storms wash over the outer
beach rim to flood the beach depression, as
indicated by accumulated pumice in the floor of
this depression.

356

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 1 . Raine Island lies on the north-western end ofRainc Island Reef. This small detached reef outside the main outer
barrier reef marks the eastern boundary of the Raine Island Entrance. (8 April 1980). A, Raine Island and its reef,
viewed looking north. B, this view ofRainc Island shows a reduced beach vegetation cover outside the cliff line.

The island is devoid of trees. The inland
portion of the beach is covered by grasses and
herbs that vary in density and extent from year to
year (Fig. 3). In breeding seasons with high density
turtle nesting, the beach can be completely
dug-over almost nightly and, as a result, can be
essentially devoid of surface vegetation, e.g.
following the 1974, 1984 and 1996 breeding
seasons (Fig. 3 A,F). Following such dense nesting
seasons, the surface vegetation regenerates
during and following the wet season, in the early
to mid year. In a low density breeding season,
there may be little disturbance to the beach
vegetation (Fig. 3C) and it may be up to 0.5m
high and extend over a half of the beach width at
the commencement of the wet season. On all
visits, the beach surface was uneven and cratered
with turtle body pits of varying age.

The typical water line to cliff tracsect on the
island, demonstrates a beach crest as an
unvegetated outer ridge approximately 1 m
above the mean HWS tide level. From this ridge
the beach slopes downward into a depression
before again rising to the base of the cliff line
edging the central rock platform. The beach
depression encircles the island except tor a
narrow rocky area adjacent to the tower. The
floor of the beach depression can be up to 1 m
below the outer beach ridge (i.e. at approximately
the same horizontal level as the mean HWS tide)
and slopes upwards in the seaward direction by as
much as 13°.

The cliff line around the central rock platform
(Figs 3, 4) ranged 1 -2m in height and was usually
a barrier to turtles climbing from the beach onto
the surface of the central platform. At times,
nesting turtles at the base of the cliff threw up
sand rampways that provided temporary access
onto the central platform by other nesting turtles.

The position of the ramps constantly changed in
high density nesting seasons as turtles dug away
some ramps and formed others. These ramps
provided access to the raised surface of the rock
platform, especially in years of high density turtle
nesting. This resulted in digging turtles creating
patches of disturbed vegetation and bare sand on
the adjacent rock platform surface adjacent to
natural ramps (Fig. 3A). From time to time,
nesting turtles would also undermine sections of
the cliff to form small caves under the edge of the
central platform.

The central rock platform consists of a cemented
phosphatic limestone and has an approximately
horizontal upper surface approximately 2m
above the HWS tide level. A vegetated sand ridge
of uncemented calcareous sand, rising to a
variable height up to approximately 5m above
HWS tide level and covered with grass and herbs,
rims the rock platform and surrounds the central
depression (Fig. 2). The floor of the central
depression is the upper surface of the phosphatic
limestone platform and has been modified by past
guano mining.

MOULTER CAY. Moulter Cay, formerly called
Pandora Cay (1 1°27’S, 144°00’E) (Fig. 5 A), is a
vegetated sand cay on the extreme outer barrier of
the Great Barrier Reef, lying 14.5km north of
Raine Island about 82km from the mainland coast
at Cape Grenville (King et al., 1 983b). The island
measured approximately 1150m in circum-
ference at the HWS tide level (Table 2). The
calcareous sand beach was of very similar
composition to that of Raine Island. The island
had an outer unvegetated and an inner vegetated
portion similar in width, elevation and
topography to that of Raine Island. The small
central rock platform of phosphatic limestone is
partially bordered by a low cliff less than 1 m high

GREEN TURTLE POPULATIONS OF RAINE ISLAND

357

FIG 2. Map of Raine Island, prepared by the late Brian King, showing the 100m interval grid lines. Study sites, 100m
wide, for quantifying nesting success and clutch disturbance are designated by letters A to E.

at its maximum. As a result of this incomplete and
low elevation cliff, no part of this island is
protected from nesting turtle incursion. In the less
dense turtle nesting seasons and following the
wet season, the rock platform and the inner beach
area has by a grass and herb coverage to
approximately 0.5m high. There have been no
trees established on this island in historic times.

MACLENNAN CAY. Maclennan Cay (1 l 0 22’S,
143°48’E) is a very small vegetated sand cay on a
small reef inside the outer barrier of the GBR
32km northwest of Raine Island (King et al,
1983a) (Fig. 5B, Table 2). Its elevation is
approximately 0.5m above the mean HWS tide
level. The sand substrate of the island resembles
that of Raine Island and Moulter Cay but there is
no central rock platform. It is sparsely vegetated
with grass and herbs and does not have
established trees. The island is frequently washed
over by storm surges.

NO. 7 SANDBANK. No.7 Sandbank (13°27’S,
143°59’E) is a small unvegetated sand cay on the
extreme outer barrier of the Great Barrier Reef
approximately 37km east of the mainland at Cape
Sidmouth and 7km south southeast of No. 8
Sandbank (King & Limpus, 1983) (Fig. 5C,
Table 2). The beach is composed of uncemented
calcareous sand rising to less than lm above
HWS. The island does not have a central rock
platform. The island has been devoid of trees
during historical times.

NO.8 SANDBANK. No.8 Sandbank (I3°22’S,
143°58’E) is a small vegetated sand cay on the
extreme outer barrier of the Great Barrier Reef
approximately 35km east of the mainland at Cape
Sidmouth (King et al, 1 983c) (Fig. 5C, Table 2).
As with No.7 Sandbank, the beach is composed
of uncemented calcareous sand rising to
approximately lm above HWS. There is no
central rock platform on the island. The central
portion of the island is covered with grass and
herbsand the island has been devoid of trees
during historical times.

GENERAL OBSERVATIONS ON THE OUTER
BARRIER CAYS.

Trees. Seedling trees naturally germinated from
beachwashed seeds (including Barringtonia
as ia tic a. Cocos nucifera and Nypa fruticans)
(Fig. 6) were found on the outer beach rim of
Moulter Cay in December 1977 and Raine Island
in December 1 978. By December these seedlings
were stressed and dying in the dry substrate of the
outer beach rim. In most summers nesting turtles
destroy any surviving plants on the outer beach
rim of these islands. Any young tree that survived
the digging of the nesting turtles would be used
by perching sea birds and probably killed as a
result.

Although Raine Island has been devoid of
naturally occurring trees, some have been planted
at various times by crews of passing boats,
commencing with the visit of HMS Fly (Jukes,

358

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 3. Annual fluctuations in beach vegetation cover of Raine Island. Southern beach looking towards the beacon. A,
1974-75 breeding season, 1 March 1975. The beach is almost totally devoid of vegetation. Adjacent to sand
rampways that gave access to the top of the rock platform for nesting turtles, the vegetation above the cliffline is also
greatly reduced. B, 1 976-77 breeding season, 3 December 1 976. C, 1 977-78 breeding season, 5 December 1 977. This
view illustrates the low level of disturbance of beach vegetation that can occur in years of very low density turtle
nesting. D, 1978-79 breeding season, 2 December 1978. The same log is in the foreground as in B. E, 1980-81
breeding season, 5 December 1980. F, 1984-85 breeding season, 19 December 1984. This view illustrates the
extreme level of disturbance of beach vegetation that can occur in years of very high density turtle nesting.

1847). The BBC documentary Zoo Quest: The
Birds of Paradise II. Raine Island documents one
such planting in July 1957. These trees have not
survived in the long term. On the last occasion,
six coconut palms ( Cocos nucifera) were planted
on the western sand dune of the central rock
platform during 1980. Since 1976, any coconut

trees found planted on these islands have been
removed by QPWS staff.

Vertebrates. There are few species of animals
resident on these islands that have an impact on
the marine turtles. These outer barrier cays
support no native terrestrial mammalian or
reptilian inhabitants, except for Bramble Cay
which supports a gecko and an endemic native

GREEN TURTLE POPULATIONS OF RAINE ISLAND

359

FIG 4. Margin to the phosphatic limestone platform,
Raine Island adjacent to the beacon where the rock
platform had a well defined cliff margin.

rodent, Melomys rubicola. This is a small herb-
ivorous rat that does not prey on turtle hatchling
or their eggs (Limpus et al., 1983d). The gecko is
Hemidactylus frenatus , a small insectivorous
lizard that was probably introduced with the
building materials or food parcels supplied to
Bramble Cay during the 1970s when it was a
research study site and egg collection site for the
Torres Strait turtle farms. Goats were released
onto Raine Island in the 1840s but disappeared
from the island sometime late last century
(Stoddart et al., 1981).

The extensive avifauna of these outer barrier
coral cays has been reviewed in recent years:
Raine Island (King, 1986), Moulter Cay (King et
al., 1983b), Maclennan Cay (King et al., 1983a),
No. 8 Sandbank (King et al., 1983c), No.7
Sandbank (King & Limpus, 1983). However,
most of the sea bird species, and in particular
those that occur in very high density are not
predators of the turtles. Observations of bird
predation of the turtle hatchlings will be
discussed in the section dealing with hatchling
predation.

An estuarine crocodile, Crocodylus porosus ,
was observed at Raine Island on one occasion; it
is commonly seen in low numbers on the inner
shelf islands (Limpus, 1980c) and has been
recorded preying on marine turtles elsewhere
(Limpus et al., 1983b).

Invertebrates . Terrestrial crabs, all of which
recruit to the islands from aquatic larval stages,
have been scarce on Raine Island. Two species of
ghost crab, Ocypode cordimana and O.
ceratophthalma , occur at low density. These
species occur on al most all islands in this area and
have been recorded preying on turtle hatchlings
and eggs. O. pallidula was collected from No. 8
Sandbank. Two species of terrestrial hermit crab

occur on these outer barrier reef cays. Coenobita
nigosus, has been found at low density on Raine
Island, Moulter Cay and No.8 Sandbank and in
high density on No.7 Sandbank. The larger reddish
coloured species, C. perlatus has been particularly
abundant on No.7 Sandbank. These Coenobita
spp. have not been observ ed interacting with the
turtle hatchlings or eggs although C. perlatus is
large enough to prey on hatchling turtles.

Oligochaete worms were recorded among
hatched and unhatched eggs in nests at Raine
Island in January 1997, December 1999 and
December 2001. Specimens (N39131) collected
29 January 1 983 by B. King were lodged with the
Queensland Museum.

Climate Data . The north-western Coral Sea
regional climate recorded at the three Bureau of
Meteorology weather stations in closest
proximity to Raine Island is characterised by
relatively uniform maximum and minimum air
temperatures year round. The warmer months
coincide with a distinct summer wet season from
December to April (Fig. 7). Mean monthly daily
air temperatures are above 25°C year round and
the mean monthly maximum air temperature did
not exceed 35°C.

Eastern Australia and the Coral Sea region are
subject to considerable variability in annual
climate. This is illustrated by the variability in the
monthly Southern Oscillation Index (SOI) (Fig.
8). Region rainfall, which has impacts on marine
turtle nesting (Limpus et al., 2001), is under the
influence of the regional climate fluctuations.
Periods with a negative Southern Oscillation
Index (SOI) are usually characterised by low
rainfall while positive values are usually
associated with elevated rainfall levels
(Couper-Johnston, 2000). Examination of SOI
fluctuations over the last 31 summers (Fig. 8)
indicates that during the last six summers
(since 1996) there has been an atypical sequence
of wet seasons. Indeed, based on positive SOI
values, Raine Island since 1 996 should have been
exposed to 5 of the 6 wettest summers in the last
25yr (Figs 7, 8). It has been during this same
period that flooding of the nesting habitat has
become an obvious and regular event at these
northern GBR rookeries. Photographs (Fig.
9A,B) demonstrate clearly that the water table
can rise higher than the depth at which the turtle
eggs have been laid. At this time we are not
prepared to speculate as to whether this pattern of
more regular wet summers is the result of long
term climate fluctuation or of global warming
(Manton et al., 2001).

360

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 5. Other coral cays of the far northern Great Barrier
Reef. A, Moulter Cay, 8 April 1980. This view of the
island shows it with reduced beach vegetation outside
the cliff line. B, Maclcnnan Cay, 8 April 1980. C, No.7
Sandbank (foreground) and No.8 Sandbank, 19
December 1984.

HISTORICAL REVIEW OF PRE-1974
OBSERVATIONS OF TURTLES IN THE
RAINE ISLAND REGION

The Raine Island Entrance from the Coral Sea
through the Great Barrier Reef to Torres Strait
was the focus of considerable navigational
activity during the early to mid 19th Century.
This, along with the large green turtle and sea bird
populations associated with the adjacent islands
(Raine Island and Moulter Cay) has resulted in
numerous visits to the region by early Australian
explorers, historians, sailors and naturalists. As a

FIG 6. Seedling coconut palm. Cocos nucifera , growing
on the outer beach rim of Moulter Cay, 3 December
1977.

consequence there has been a considerable
volume of natural history writings and
observations on these islands.

Raine Island and adjacent Moulter Cay support
one of the few remaining large breeding
aggregations of Chelonia mydas in the world. In
most areas where the species has bred in
abundance in the past, the breeding populations
have been greatly reduced, or taken to extinction
by over harvesting at places such as Grand
Cayman Island, Bermuda and Reunion Island
(Groombridge & Luxmore, 1989). The current
status of the turtle population that nests at Raine
Island and Moulter Cay will be, in part, a function
of the past history of human utilisation of the
turtles and the islands. The following review of
marine turtle records from these islands provides
an historical context against which results of
recent research can be compared and future
change assessed.

The Queensland Turtle Research Project
expanded to include systematic studies of the
Raine Island Chelonia mydas rookery during the
1974-1975 breeding season. This then has been
taken as the end point of the historical review of
marine turtles in the Raine Island area. In this
review, information has been brought together
from diverse sources: published accounts of
visits to the islands, interviews with fishermen

GREEN TURTLE POPULATIONS OF RAINE ISLAND

361

and other visitors to the area,
newspaper accounts of visits to the
islands, and old photographs and
documentaries. Together they contain
a considerable body of knowledge,
spanning 131 years of turtles at Raine
Island and Moulter Cay. Also high-
lighted are some past misconceptions
concerning marine turtle biology,
including the belief that marine turtles
i jul acg sep oct nov dec jan feb mar apr may jun could be systematically exploited on a

grand scale.

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» Willis Is: Max T ■ Willis Is: Min T
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□ 1950-1991 (TI)
-1996-1997 (Horn)
*■ 1997-1998 (Horn)
-1998-1999 (Horn)
- 1999-2000 (Horn)
— 2000-2001 (Horn)
♦ 2001-2002 (Horn)

JUL AUG SEP OCT NOV DEC JAN FEB MAR APR MAY JUN

FIG. 7. North-western Coral Sea Regional climate data from
Australian Bureau of Meteorology stations. A, monthly minimum
and maximum air temperature data, Willis Island. B, monthly
rainfall data from Lockhart River, NE Cape York Peninsula.
C, monthly rainfall data from Horn Island and Thursday Island (Tl),
Torres Strait. D, monthly rainfall data from Willis Island.

HISTORICAL REVIEW. Raine Island
was discovered by Thomas Raine on
the HMS Surry in 1815 (de Salis,
1969). He did not land on the island
and no reference was made to turtles.

Jukes (1847: 130-131) provided the
first description of Raine Island and its
natural history during the visit by HMS
Fly on 29-30 July 1843. While the
primary interest in the turtles was to
provision the ship. Jukes provided
some interpretation of the behaviour of
the turtles at the island. They recorded
a few turtle tracks on the beach, but did
not capture any. Jukes was impressed
by the large numbers of dead turtles
and turtle bones on the island. He
interpretation of the cause ol their
death was ‘that when feeble, from
sickness or age, the turtles have come
here to die, and that those lying on their
backs had died in a vain attempt to
crawl up the broken bank into the
interior of the island’ (Jukes, 1847:

1 30). On 1 August 1 843 he expressed a
similar interpretation ot why large
numbers of turtles died at Moulter Cay
(Jukes, 1847: 131).

HMS Fly and HMS Bramble
returned during 29 May - mid
September 1 844 with a labour force of
convicts to construct a navigation
beacon on Raine Island (Jukes, 1 847).
The quarries dug during this
construction work provided insights
into the geology of the island and
observations of fossil turtles and turtle
eggs in considerable abundance
(Jukes, 1847). Based on his observ-
ations of the fossil turtle eggs in nests,
Jukes concluded that the rock forming
the central elevated platform of Raine

362

MEMOIRS OF THE QUEENSLAND MUSEUM

Jin .Inn Jin Jin Jan Jan Jan Jan Jan Jan Jan Jan Jan Jan Jan Jan Jan
1971 1973 1975 1977 1979 1981 1983 1985 1987 1989 1991 1 1993 i 1995 1997 1999 2001 |

FIG. 8. Fluctuations in Southern Oscillation Index (SOI) over the 31
summers 1971 - 2001 (Bureau of Meteorology Australia, 2002). Summer
SOI values (November - February) are shown in black.

Island had not been formed underwater and
subsequently elevated.

MacGillivray (1846: 1474-1478), the
naturalist on board HMS Rattlesnake , also visited
Raine Island at this time and commented on the
fossil turtle eggs in nests, citing them as evidence
of the recent formation of this bed of sandstone
‘only a few feet thick’ where the rock for the
tower was quarried. MacGillivray (1846) also
made a number of original observations on the
biology of the nesting turtles. All turtles visiting
Raine Island and adjacent islands were ‘ green '
turtles which came to lay eggs, mostly at night.
During June to August, single turtles came ashore
at irregular intervals while the nesting density
increased during September. He expressed
surprise at the number of dead turtles on their
backs which he interpreted as turtles that had

fallen on their backs while
endeavouring to climb the low
rocky border of the island. He
makes no reference to
Dipterans associated with the
turtle carcases, but notes the
presence of ‘several
necrophagous Coleoptera’.
Turtle bones were abundant
enough on the island for him
to note that ‘noddies, Anous
stolidus , used fragments of
turtle shell and turtle bones
among their nest building
material’. While he wrote that
hatchling turtles were ‘preyed
upon by various kinds of
sea-fowls, from the
frigate-bird down to the sooty
tern’, he only reported on one
observation of a ‘young turtle from the stomach
of a tern which had swallowed it entire’.

An immature hawksbill turtle, Eretmochelys
imbricata , was presumably collected during the
1844 visit by HMS Fly : British Museum
specimen, register no. 46.7.27.6; immature with
curved carapace length = 35.5cm; collected at
Raine Island and presented by Lieut. Ince in July
1846 (A.F. Stimson in litt., 26 May 1977). This is
the Boulenger ( 1 889) specimen V.

The crew of the Heroine went ashore on Raine
Island on 28 January 1845 at daylight and
‘obtained fourteen large turtles, each averaging
four cwt.; also an immense number of eggs, and
the crew killed birds out of number’ (Mackenzie,
1845:494).

FIG. 9. Raine Island during early December 1 999 showing: A, the elevated water table level on the southern beach
platform; B, a perched water layer in the central depression (flooding at this level is not tidal).

GREEN TURTLE POPULATIONS OF RAINE ISLAND

363

Sweatman (in Allen & Corris, 1977) was a
clerk on board HMS Bramble with a
responsibility for keeping records of provisions
for the ship throughout its voyages and, as a
result, his diary provides many quantified details
of the occurrence of turtles. Crew members from
HMS Bramble were on Raine Island during 28
April - 4 May 1845 when they caught 14 turtles.
One was a live turtle which had fallen into the
well near the tower and been trapped there for
some time. He describes nightly patrols of the
island at 2 hourly intervals to locate the nesting
turtles, the walking of the turtles on tethers in the
shallows to move them around to the anchorage
end of the island by day and the novelty of riding
on the turtles back in the shallows. Sharks were
reported to attack the turtles being walked on
tethers. Sweatman also describes abundant remains
of dead turtles on their backs along the cliff.

Sweatman (in Allen & Corris, 1977: 55), in
commenting on the food shortage on board HMS
Bramble as she returned to Port Jackson on 9 July
1845, noted that ‘Had it not been for the turtle
caught at Raine’s It. and the fish caught on the
reefs we would have been destitute for food 3
weeks before’.

Stoddart et al. (1981) reported that the HMS
Heroine was again in the vicinity of Raine Island
on 5 August 1 846 but there is no recorded attempt
to collect turtles. However, the date of this visit is
probably incorrect because the Heroine was
recorded sunk on 24 April 1846.

On 24 July 1850 the HMS Enchantress was
wrecked on Raine Island Reef. The crew went
ashore on Raine Island and was rescued on 25
July 1 850 by the by HMS Lady Margaret (Lack,
1 953). The ship HMS Constant was also wrecked
on Raine Island Reef on 12 July 1858 (Loney,
1980). There were no records of turtles being
encountered by the survivors of either of these
wrecks.

Rattray ( 1 869) visited Raine Island in 1 860 on
board HMS Salamander and interpreted the
phosphatic limestone rock of Raine Island as ‘a
thin layer of guano deposited by the numerous
turtle and flocks of terns, gannets, and other
aquatic birds that, like the former, make this their
headquarters and favourite breeding-place’. The
authors are unaware of any other report that
attributes guano formation to turtle nesting
activity. Given that turtles do not del'aecate while
on the nesting beach, this interpretation is
presumed to relate to the incorporation of
decaying turtles into the substrate.

In 1860, HMS Herald visited Raine Island
twice (Denham, 1861): in April when supplies
were left on the island; and 19 September to
check on the condition of the supplies. There was
no reference in Denham’s diary to turtles or any
other aspect of the island from either visit.

In 1862, Raine Island was leased for guano
mining but there appears to be no reference to
turtles in respect to this (Crowther, 1939).

In about 1 874 a beche-de-mer fishery camp
was operating on Raine Island and staffed by 2
whites, 2 or 3 Chinese, several Kanakas and some
aboriginals from the adjacent mainland (Ellis,
1936). The duration of this operation was not
stated but Ellis implied that they harvested turtles
from the island to provision the crew.

Moseley (1879), in reporting on the 3 1 August
1874 visit of HMS Challenger to Raine Island,
commented on the death of large turtles on the
island. He interpreted the death of many turtles at
the base of one part of the cliff as being the result
of turtles coming ashore to lay eggs. He believed
that they were unable to turn around or go
backwards and hence died at the foot of the small
cliff.

During 1890-92, Raine Island was extensively
mined for guano with approximately 1 1 0 persons
resident on the island at times and turtles were
taken for food (Ellis, 1 936). The total harvest was
not documented: one turtle was killed daily
during the nesting season to feed the European
staff (about 10 people); eggs were also
occasionally eaten; but the consumption rate by
the approximately 1 00 Chinese labourers was not
recorded. Ellis, the manager of the mining
operation, provided the first account ot the
massed nesting by green turtles at the island: The
coral sand beaches of Raine Island were visited
by incredible numbers of green turtles during the
summer months, for the purpose of laying their
egus. In the height of the season it was difficult to
find a few yards of beach free from their broad
tracks made the previous night, w r hen perhaps
more than a thousand would have come up’.
Ellis’s description also includes an account of the
nesting behaviour of the turtles that agrees well
with the recent technical description of green
turtle nesting behaviour by Bustard & Greenham
(1969). He gives a clutch count of ‘eighty to a
hundred or more’ and an incubation period of
‘about six weeks’. He provides a vivid account of
the co-operative digging of hatchlings to reach
the beach surface, the resting of masses of
hatchlings near the beach surface until after dark,

364

MEMOIRS OF THE QUEENSLAND MUSEUM

the synchronous emergence of hatchlings from a
nest and their vigorous dash for the sea. Ellis
makes no reference to birds as predators of
hatchlings but gives the impression that sharks
took large numbers of hatchlings as they sped
across the reef flat. The sport of riding turtles over
the reef flat is commented on, and support is
given to De Rougemont’s claims that people
could ride turtles in the sea (Maslen, 1977). The
large numbers of nesting turtles were regarded as
a nuisance as they dug up paths and disturbed
people’s sleep when they threw sand against the
sides of houses during the night. Ellis also
described turtles entering the central depression
and dying of heat exhaustion during the day.

W. MacGillivray, an ornithologist, made
several trips to Raine Island and the adjacent area
during 1910-1913. On 30 October 1910 he
recorded ‘great numbers’ of turtles on the beach
at Raine Island and in the adjacent shallows
(MacGillivray, 1910). During his 9-15 July 1911
visit to Raine Island he made no reference to
turtle nesting or to bird predation on turtle
hatchlings (MacGillivray, 1914). However,
during his 4- 1 2 December 1913 visit he provides
a description of Raine Island and Moulter Cay
under high density green turtle nesting conditions
(MacGillivray, 1917). In this account, the crew
were described leaping from the boat onto turtles
for rides as hundreds of turtles swam past the
vessel and, after the party landed, riding turtles
down the beach into the water. MacGillivray
described himself as ‘absorbingly interested in
the wonderful bird-life’ of the islands during the
day and ‘tormented at night by the nesting turtles,
who would persist in trying to walk over or under
our sleeping shelter’. This negativity towards the
turtles was not evident in his graphic word picture
of massed nesting at Moulter Cay: ‘In the
moonlight we noticed thousands of turtles
climbing the sand-bank, their wet backs
gleaming with phosphorescence as they leave the
water’. The whole surface of Moulter Cay was
described as ‘ploughed up by the tracks and pits
of the reptiles, and the air is redolent with the
odour of the dead’. At Moulter Cay he also
described the destruction by nesting turtles of
bird eggs in a large crested tern. Sterna bergii ,
nesting colony. S. bergii and silver gulls, Lams
novaehollandiae , were recorded scavenging on
turtle eggs dug up by nesting turtle during the
previous night at Raine Island. Rufous
night-herons, Nycticorax caledonicus, were
recorded as numerous on Raine Island and
identified as predators of the hatchlings

(MacGillivray, 1918a). In his summary of brown
booby, Sula leucogaster, nesting, MacGillivray
(1918b) again voiced negativity towards the
turtles when he described Moulter Cay as ‘a large
turtle-infested sand-bank’.

On 7 September 1950, the Order in Council of
15 December 1932 under the Queensland
Fisheries Act was repealed. The 1932 regulation
had put in place a closed season for the harvest of
green turtles and their eggs during the months of
October and November south of 17°S. in its
place, this 1 950 Order in Council declared a year
round closed season for green turtles and their
eggs throughout all Queensland. This was the
first conservation management measure applied
to the north Queensland turtle populations.

Our discussions with several people, including
D. DufTey in 1974 and J. Dagie in 1978, who had
been crew members of trochus luggers that
operated out of Torres Strait, indicated that
trochus luggers travelling south for trochus
gathering in Great Barrier Reef waters, regularly
travelled via Raine Island. Here they took on live
green turtles as deck cargo as food for the crews.
Many of these islanders have left their names
inscribed in the interior wall of the beacon as the
only record of their visit to the island (Bairstow,
1983).

Vince Vlassoff (pers. comm., 2 April 1981)
visited Raine Island during a summer turtle
nesting period in February - March of 1956 (the
same year that Cyclone Agnes impacted north
Queensland). During one night, his party counted
turtles in several hundred yards of beach and
extrapolated for the entire beach. They estimated
10-12 thousand green turtles for the night. They
rescued about 10 turtles per morning from on
their backs at the base of the cliffs. It was
estimated that they ‘could smell dead turtles from
about 2 miles (3km) out to sea’ (Fig. 1 0). The well
was about 4ft (~1.3m) deep but had been filling
up with dead turtles (Fig. 10D). Photographs
taken on this trip provide the opportunity for the
first independent validation of the species
identification of the nesting turtles at Raine
Island. Up to this time, it appears that no
specimen of a green turtle from Raine Island or
Moulter Cay had been lodged in a Museum.
During this trip Vlassoff was accompanied by an
Italian film crew from the Astoria Film Society.
They filmed for an Italian documentary titled La
Grand Barrier. This documentary, if it still exists,
would provide the oldest film record of the island

GREEN TURTLE POPULATIONS OF RAINE ISLAND

365

FIG 1 0. Raine Island in February to March of either 1 954 or 1 956. Photographs by V. vla ss°ff showing > evidence of
extremely high density turtle nesting for one or both of these breeding season. These are c o t es ' fe
for Raine Island. A, numerous recently dead adult Chelonia mydas that had died Irom heat ex aus .

central depression. B, the surface of the rock platform adjacent to the beacon was almos co p > _ y
vegetation. C, a group of dead or dying adult female Chelonia mydas that had fallen over e c 1 c •

Note the relatively bare sand surface of the rock platform above. D, remains ol 2 recen > c '
mydas in the well at the western end of the central depression of the rock platform.

and its wildlife, including footage from a very
high density turtle nesting season.

During 20-25 July 1957, Vince Vlassoff
returned to Raine Island on his vessel. Tropic
Seas , with a BBC Film Unit including producer
David Attenborough (D. Attenborough, in litt. 4
April 1984; V. Vlassoff, pers. comm., 2 April
1981). The BBC documentary Zoo Quest: The
Birds of Paradise II. Raine Island was filmed
during this visit. This documentary provides a
visuaF description of an island lush with grass
cover across the rock platform. The images of
dead turtles in the documentary were consistent
with few turtles having died in the turtle breeding
season some six months earlier. All except one
carcass was a year or more old. There was low'
density turtle nesting during this winter visit with
about twenty sets of tracks from nesting turtles
counted in four nights. This documentary and
photographs by D. Attenborough provide the first
positive identification that the turtle species breed-
ing at Raine Island in mid-year (winter) is C. mydas.

Changes in fisheries regulations in 1958 opened
the way for commercial harvesting of turtles from
the northern Great Barrier Reel. Records con-
tained in the 1956-1958 Queensland Department
of Harbours and Marine files indicate that the
Cairns based Whittaker brothers (Snowy and
Neil) of the fishing vessel Trader Horn, lobbied
during 1956 and 1957 via their local member ot
parliament, GW. Wallace MLA, to have permits
issued to harvest turtles in north Queensland.
They claimed that there were so many turtles
nesting in areas of the Northern Great Barrier
Reef that the turtles were unable to find places to
deposit their eggs and consequently they dig out
eugs previously laid by other turtles, and that on
the following morning these are eaten by the
birds. They further claimed that ‘over a period ot
six to eight weeks, as many as 1,500 turtles visit
Raine Island nightly’ (B.W. Dick, in litt. 26 Feb.
1957, Brisbane). They suggested that the permit
embrace the area from Olinda Entrance to No. 7
Bank, including Pandora (= Moulter Cay), Raine

366

MEMOIRS OF THE QUEENSLAND MUSEUM

Island, Ham Reef, Derry and Nos. 8 and 7
Sandbanks. Their concept of the biology of green
turtles included the following ideas: turtle laying
season extended from about November to March;
less than 1% of the turtles hatched are males and
therefore, unlike most other animals eaten by
man, it was better that the females should be
taken; it was appropriate to harvest the female
turtle on the nesting beach as she headed back to
the water because as long as the females lay once
the turtles reproduce themselves; very few
hatchlings ever live to become adults which
weigh about 1 14kg; green turtles take only about
four years to grow to adults (Anon, 1 959a); there
were so many green turtles in the area that they
really needed thinning out; and the crew would
save more than they killed, because they would
take any hatchling that they found to the water
and save them from death by birds (Anon,
1959b). Their political lobbying was successful,
and on 4 September 1 958, the Order in Council of
7 September 1950 under the Queensland
Fisheries Act was repealed and a new Order in
Council declared which restricted the year round
closed season for harvesting green turtles to south
of 15°S. The year round closed season for turtle
egg harvest throughout Queensland was retained.
This meant that turtles could be harvested
without restriction in Queensland north of 15°S.

In addition to published material and corres-
pondence, information regarding the Whittaker’s
turtle harvesting business was obtained from
interviews in Cairns in August 1 983 by CJL with
Neil Whittaker himself and E. Gray who was a
crew member of the Trader Horn for the first
major turtle harvesting trip to Raine Island in
January 1959. Neil and Snowy Whittaker’s
vessel for the turtle harvesting from Raine Island
was the Trader Horn : displacement 70 tons,
engine 115HP, overall length 72ft, beam 17ft,
draft 7ft, wooden hull, built by Alf Hansen of
Cairns (N. Whittaker, pers. comm.). She carried 7
dories and a crew of 12 (Anon, 1959a).
According to Neil, the Trader Horn had a freezer
capacity of about 50,0001b (22,700kg); most of
the turtles were taken from Raine Island but some
came from Moulter Cay; Moulter was hard to
work when a swell was running; they had no
problem capturing 100 turtles per night for
slaughter on every trip; turtles were turned on the
beach at night, killed and partly butchered on the
beach next morning, loaded on a dory (Fig. 1 1 A),
transferred to the Trader Horn where the meat
was washed on the deck and butchering
completed (Fig. 1 1 B); only meat (= muscle) was

taken. According to E. Gray: the turtles were
butchered at the island, at first by using an axe;
the white crew did not like this job and were slow;
the boat went to Thursday Island to employ extra
native crew to slaughter the turtles; in the latter
part of the first harvest many turtles were shot for
butchering; and the meat was marketed through
Cairns. Gray also observed that the green turtles
that they harvested were nesting in sufficient
numbers so that ‘at times you could walk a long
distance along the beach from turtle to turtle’;
most nesting was at night and on the eastern half
of the island. Gray did not go on the second trip in
February 1959. According to G. E. Rowell (in litt.
to Queensland Department of Harbours and
Marine, 21 April 1959), once the refrigerated
holds were filled with turtle meat, killing ceased
and the Trader Horn s deck was loaded with live
turtle for the return trip to Cairns (Fig. 1 1C). The
live turtles from the first major catching trip were
slaughtered at the Smith’s Creek Wharf in Cairns.
Rowles, as Harbour Master, advised Whittaker
that this was not permitted. Whittaker took the
live deck cargo turtles from the last trip out to
Michelmas Reef where they were butchered and
the meat returned to Cairns.

The Whittaker’s turtle harvesting enterprise
was initiated with a small unquantified harvest
from an unstated origin from early in the nesting
season in late 1958 (Anon, 1959b). Some of this
turtle meat was used to promote sales to Meyer
Emporium of Melbourne, Victoria, and to
presumably supply the demands of local hotels.
The first substantial harvest occurred during a
two week trip to Raine Island in January 1959
(Anon, 1959b). The second major expedition to
harvest turtles occurred during February 1959.
The itinerary of Trader Horn for this February
trip was: 14th Raine Island, 14-16 Pandora Cay
(= Moulter Cay), 17-18 Murray Island Sandbank,
19-22 Pandora Cay, 22 Raine Island. Whittaker
met a visiting ornithologist, Dr Warham, on
Raine Island by chance and invited him to travel
with them for the remainder of the trip (Warham,
1963). On this latter trip, 508 turtles were taken
that yielded about 36,000 lb of turtle meat (Anon,
1959c). G. Rowell (in litt. to Qld Dept, of
Harbours and Marine, 10 April 1959) provided a
summary of the combined summer’s turtle
harvests by the Whittakers: ‘Turtle fishing was
commenced in early January, and discontinued
during March. During that period approximately
1,200 turtles were taken which produced
approximately 80,000 lbs of meat (about 60 to
701bs of meat per turtle)’.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

367

FIG. 1 1 . Photographs by Dr J. Warham documenting
some of the procedures of the Whittaker brothers’
turtle harvest at Raine Island in February 1959. A,
dory and crew at the beach at Raine Island. B,
cleaning and dressing turtle meat on the deck of the
Trader Horn. C , once the freezers were full,
additional turtles were stacked live on the deck for the
return trip to Cairns.

In preparation for the commencement of turtle
harvesting from the Raine Island area, Mr T.A.
Hiley was reported in the Cairns Post in Decem-
ber 1 958 discussing a proposed amendment to the
Fish Supply Management Act that would remove
turtles from the definition of fish under the
Queensland Fisheries Act. The purpose of this
was to encourage development of the turtle meat
industry north ot Cooktown by allowing people
engaged in the industry to operate outside the
Fish Board and sell direct to the public. This
change in regulations did not occur.

Turtle meat was not novel to the Cairns
community at this time. The Strand Hotel had
been serving turtle steaks at the hotel for six or
seven years (Anon, 1959b). The chef, Mr. Leigh
Kingston, acknowledged that it was very popular
with the southern tourists during the tourist
season but the local people did not take to it too
well at the beginning (Anon, 1959b). As part of
the promotion for sale of turtle meat locally, this
hotel staged at least one cooking demonstration
for the media and outlined styles of preparation of

turtle meat meals prior to the first major
consignment of turtle meat arriving in Cairns
(Anon, 1 959b). A local wholesaler in Cairns, Mr.
M. Stock, was organised to distribute the
Whittakers’ turtle meat at 3/- per lb (Anon,
1959b). Snowy Whittaker promoted it as
reasonably priced meat for the ‘working man’
(Anon, 1959b).

On 15 January 1959, L. Jorgensen, Manager of
the Cairns Fish Board, announced (Anon, 1 959d)
that under the Fish Supply Management Act,
because turtles were defined as fish under the
Act, it was illegal for the Whittakers to sell their
turtle meat in Queensland to anyone except the
Fish Board. They were however, entitled to sell
their turtle meat direct to buyers outside
Queensland. The Fish Board's commission on
the sale of turtle meat was 15%. Snowy
Whittaker responded that ‘the housewife would
have to pay at least 4 d per lb more for it' (Anon,
1959e). Some of the turtle meat was passed
immediately into the local Fish Board coldrooms
for local sales. By early February, turtle meat

MEMOIRS OF THE QUEENSLAND MUSEUM

368

REAL TURlLfc

Add T**"'

*** * r~>

,Cc. o« Umon. **
Br wln °
if S * fV *

SL-T-HWAV.9

Win..**.**"’

«rd S««*>" ,n 9'

@)Foo<^

FIG 1 2. Label from a can of Master Foods turtle soup made from Raine Island Chelonia mydas harvested by the
Whittaker brothers in January - February 1959. Master Foods continued to market turtle soup under this label for
approximately another two decades even though their turtles did not originate from the Great Barrier Reef.

from Cairns at 4/6 d per lb was selling in
Townsville in favour of more expensive beef
(Anon, 1959f)- The Whittakers canvassed
markets interstate in Sydney and Melbourne with
the assistance of the Queerah Meatworks
providing storage of the majority of the meat
without surcharge (Anon, 1959g; N. Whittaker,
pers. comm.). Additional markets were also
sought internationally in Singapore (Anon,
1959g) and New York (N. Whittaker, pers.
comm.). As a result of a Cairns visit by Dr V.
Lewis of the Sydney canning company of Lewis
and Sons to inspect the Whittakers’ catch (Anon,
1959h), most of the turtle meat was to be sold to
Lewis and Sons at l/9 d per lb (N. Whittaker, pers.
comm. ). This meat was processed at their Sydney
cannery into turtle soup and marketed under the
‘Masterfoods’ label (Fig. 12) (N. Whittaker, pers.
comm.). This, association did not continue as
expected because of financial issues (N.
Whittaker, pers. comm.).

When sales of turtle meat did not increase as
expected, a publicity campaign to counter the
conservative eating habits of the north Queens-
land community provided recipes in the local
news paper for cooking turtle meat, including
fried turtle steak, crumbed steak, turtle hot pot,
turtle rolls, turtle meat sate, turtle saute, turtle
roast and curried turtle (Anon, 1959i; N.
Whittaker, pers. comm.). Additionally, the
Whittakers campaigned for the sale of turtle meat
as an alternative to fish. This led to a debate as to
whether turtle meat could be eaten by Roman

Catholics on Fridays which resulted in
Archbishop Duhig ruling that turtle was fish and
therefore suitable for Roman Catholics to eat on
Fridays (N. Whittaker, pers. comm.). This
apparently initiated further discussion between
the Fish Board and the Abattoir authority.
Because turtles were fish under the Queensland
Fisheries Act and under the Abattoir’s Act, fish
and meat could not be stored in the same
coldroom, the turtle meat had to be removed from
the meatwork’s storage. As of 10 April 1959, of
the original estimated harvest of 80,0001b of
turtle meat, about 58,0001b remained in Cairns:
23,0001b of the 25,0001b received at the Fish
Board, Cairns, remained unsold; and 35,0001b
remained in other Cairns cold storage facilities
although some of this had been purchased by a
Sydney firm (G Rowell, in litt. to Qld Dept, of
Harbours and Marine). The turtle harvest from
the Raine Island area did not continue beyond this
one season because the Whittaker brothers con-
sidered that it was not profitable (N. Whittaker,
pers. comm.).

One of the turtles brought back to Cairns by the
Whittaker brothers was presented to the Cairns
Ambulance Brigade. After preparation by a local
taxidermist, this turtle was to be displayed in the
first Cairns Ambulance Wildlife Exhibit at the
Cairns show in 1 959 in conjunction with ‘Snakes
Alive’ (N. Whittaker, pers. comm.).

N. Whittaker (pers. comm.) provided some
observations concerning turtle breeding in the
1950’s: all the sandbanks of the northern Great

GREEN TURTLE POPULATIONS OF RAINE ISLAND

369

Barrier Reef carried a few nesting turtles, but
none with numbers like Raine and Pandora (=
Moulter); at Raine Island the turtles dug up lots of
eggs while nesting; Raine Island was the only
area with lots of frigatebirds and the frigates
would prey on hatchling turtles by day off the
beach and in the water; and Neil walked the beach
one night after turning 100 turtles for killing and
counted ‘a thousand turtles in one walk of the
island'.

Warham (1963) visited Raine Island on 7-14
and 22 February 1959. The crew of the fishing
boat Galeru that delivered him to the island
harvested a nesting turtle for food. Warham
identified that many previously laid eggs were
dug out by nesting turtles; landrails scavenged on
these destroyed eggs; dead turtles were
numerous; and some nesting turtles still ashore
on the beach after daylight died from over
heating. Warham (1961) provided a description
of the topography and vegetation of the island,
described green turtles as abundant, and
identified the rufous night-heron, Nycticorax
caledonicus , as a predator of turtle hatchlings.
Warham ( 1961 ) also visited Moulter Cay during
14-16 and 1 9-22 February 1 959. He described the
island's topography and vegetation with a ‘turtle
churned level area surrounding a shrub covered
centre. Crested terns, Sterna be/*gii , which were
nesting and roosting on the island, patrolled the
beaches at daylight and preyed on hatchling
turtles on their way to the sea.

Both Neil Whittaker and E. Grey during their
above interviews concerning the turtle harvest
reported that some hawksbill turtles were seen
nesting on Raine Island but not harvested.

Another turtle harvesting business started in
Torres Strait at about the same time as the
Whittaker brothers began their venture out of
Cairns. This Torres Strait based enterprise started
in a small way using Islander labour but the
Department of Native Affairs disagreed with this
and insisted that £15 per week be paid to the
Islanders. This and unsuitable weather appear to
have stopped the project (Harbour Master,
Thursday Island, in litt. 9 June 1959).

There was at least one protest from the
conservation lobby against this newly energised
turtle harvesting industry. The Townsville and
District Natural History Society wrote to the
Queensland Government protesting against the
slaughter of turtles for their meat (Anon, 1959j).
John Warham, the ornithologist who was at Raine
Island during one of the Whittaker’s harvesting

trips, while not critical of the Whittaker’s
operation, expressed the need for caution with
regard to the number of licences operating in the
harvest if the turtle resource was not to be
substantially reduced (Anon, 1959k).

Another proposal to create industry out of the
Raine Island turtles in 1959 does not appear to
have progressed to implementation. A Cairns-
based mercantile firm, Cairns Coasters, had
acquired samples of turtle bones from ‘a vast
natural grave yard ... on the outer reef six or
seven hundred miles north of Cairns’ to test for
their phosphorus content (N. Whittaker, pers.
comm.). The plan had been for their vessel, the
West Aussie, to bring a shipload of bone back to
Cairns every time it returned from Thursday Island.

HMAS Gascoyne visited Raine Island on 1 1
November 1961 and her crew carried out some
repairs on the tower. There are no records
referring to turtles from this visit.

Paul Watson (pers. comm. 18 June 1979) first
visited Raine Island with V. Vlassoff in about late
November 1965. The area of the island with the
densest turtle nesting was at the tower end and
south side. Most turtle nesting was by night. On
one night Watson had been able to step or jump
from turtle to turtle while Vlassoff walked 100
paces beside him. He reported that he
subsequently visited Raine Island on a number of
occasions in October - December in different
years but never saw it again with turtles as thick
as on his first visit.

On 1 8 July 1 968 an Order in Council under the
Queensland Fisheries Act declared an all year
round closed season overall marine turtle species
and their eggs through out all of Queensland.
This marked the end of a 20yr window of
opportunity for commercial harvesting of green
turtles in the northern Great Barrier Reef.

In the early 1970s, Dr Robert Bustard, a
Scottish herpetologist working out of Canberra
who established the Applied Ecology Torres
Strait Turtle farms, listed Raine Island as one of
the important G. mvdas rookeries for Queensland
(Bustard, 1971) and identified Raine Island as
one of the 15 most import ant green turtle nesting
beaches worldwide (Bustard, 1972). In neither
publication did he elaborate on how he arrived at
these conclusions.

The James Cook University of North Queens-
land (JCUNQ) 1973 James Kirby Expedition
conducted biological studies at Raine Island
during 3 1 October - 4 November (Stoddart et. al.
1981). During this visit, Raine Island had a

370

MEMOIRS OF THE QUEENSLAND MUSEUM

measured high tide circumference of 2100m
( 1 600m of sand and discontinuous narrow beach
rock; 500m continuous beach rock). They
recorded a nightly green turtle track count =
133.8(SD= 17.8, range = 1 15-162, n = 4nights).
When corrected for turtles emerging across beach
rock that did not result in tracks at the water line,
their results indicated that about 200 nesting
turtles were coming ashore nightly. Their mean
curved carapace length = 109cm (SD = 5.29cm,
range = 90- 122cm, n = 124). At the time of this
early nesting season visit, there was still a
substantive Lepturus tussock grassland
extending out from the cliff to the bare sand
beach. In their effort to estimate the size of the
total nesting population, they have assumed a
1 00% nightly nesting success and that the nesting
habitat does not include that area of grassland
below the cliff. Based on the unquantified
observation that some animals ‘differed
markedly in degree of curvature of carapace’,
Stoddart et. al. (1981) postulated that some of the
nesting turtles may have been flatback turtles, N.
depressus. It appears that none of the observers
had prior experience with the flatback turtle and
no photographs were recorded of any of the
presumed flatback turtles. We therefore reject this
identification in light of prior and subsequent data.
CJL examined the skull collected from the dried
carcass of an adult sized E. imbricata found in
central depression of Raine Island by J. Collins
during this JCUNQ visit (Limpus, 1980a) and
while not reported by Stoddart et. al. ( 1981 ), this
skull remains the only conclusive evidence that
occassional E. imbricata come ashore to nest on
Raine Island.

Commenting on his first visit to Raine Island in
late January 1974, Ben Cropp (pers. comm. 3
March 1975): expressed the opinion that the
abundance of track each morning in early 1974
was of the same order of magnitude as he saw on
his next trip to the island in February 1975 when
he was accompanied by CJL, i.e. tracks from
several hundred turtles per night.

Dr G Goedon of the Queensland Fisheries
Service (pers. comm. 1975) counted about 50
recent sets of tracks below the high tide level in
one walk of Raine Island in early October 1975.
Later in the same breeding season. Dr J.
Kowarsky of Applied Ecology Pty Ltd recorded
numerous sets of green turtle tracks on Raine
Island from the night of 13 January 1976
(Kowarsky, 1978). These tracks were observed
during an aerial survey and were not accurately
countable. A Torres Strait Islander reportedly

counting 1 60 tracks (? = 80 turtles) freshly made
in one night on 29 January 1976 (J. Kowarsky, in
litt. 3 March 1976). While Kowarsky was unsure
of the accuracy of the count, it is clearly
consistent with few turtles nesting on Raine
Island that night.

HISTORIC SUMMARY. The authors are
unaware of any written records or evidence at the
island of use of Raine Island by indigenous
people prior to its discovery by Europeans in
1815. However, it is highly likely that at least the
inhabitants of Murray. Damley and Stephen
Islands of eastern Torres Strait, with their large
dugout sail canoes would have visited these
islands prior to European arrival. Similarly there
is the possibility that aboriginals from the
adjacent mainland may have also ventured out to
Raine Island under favourable weather
conditions, given that the island is visible from
the top of some inshore islands (e.g. Sir Charles
Hardy Islands) on a clear day. Since its discovery
by Europeans, Raine Island has been inhabited
for extended periods on three occasions: during
the building of the beacon in 1844, by a
beche-de-mer fishery crew in the 1870’s and by
guano miners for 3 yr from 1890. During the
same period, neither Raine Island nor Moulter
Cay have been continuously inhabited by
indigenous peoples.

From the earliest records of these islands in
1843 until the present times, it is apparent that
Raine Island has been a significant C. mydas
rookery and that large numbers of nesting
females may die on the island each year from
natural causes.

Since European discovery of these islands,
there was only sporadic low intensity harvest of
the nesting C. mydas at the islands up until
January 1959. During January - February 1959
there was a harvest of approximately 1,200
nesting C. mydas from Raine Island and Moulter
Cay. This large scale harvesting of the nesting
turtles was not repeated. Raine Island and
Moulter Cay appears to be one of the least
disturbed major Chelonia mydas rookery regions
of the World, having escaped the ravages of
intensive turtle harvesting associated with
European colonial expansion into tropical seas
(Parsons, 1962; Hirth, 1997).

BREEDING DISTRIBUTION IN THE
NORTHERN GREAT BARRIER REEF

BREEDING SPECIES. The distribution and
abundance of nesting by marine turtles in the

GREEN TURTLE POPULATIONS OF RAINE ISLAND

371

FIG 13. Maps summarising the distribution and abundance of marine turtle nesting in Queensland. These maps were
generated from the QPWS marine turtle nesting database. A, Chelonia my das, green turtle. The groupings of
rookeries that represent each of the four genetically separated stocks that breed in northeastern Australia
tire identified. B, Eretmochelys imbricata , hawksbill turtle. C, Natator depressus, flatback turtle.

372

MEMOIRS OF THE QUEENSLAND MUSEUM

northern GBR and Torres Straight region was
undertaken throughout this study by teams
landing on beaches and counting turtles or tracks
for each species on a nightly basis (Tables 3-6).
During the late 1990s. additional data on the
temporal and spatial distribution of marine turtle
breeding in Queensland was recorded during
dedicated aerial survey of nesting beaches
(Limpus et al., 2000). Detailed studies of turtle
nesting biology from the region have also been
published for Bramble Cay (Limpus et al., 2001 ),
Milman Island (Dobbs et al., 1 999) and a northern
GBR-Torres Strait survey of E. imbricata nesting
(Miller, et al., 1995). These data have been
incorporated into the QPWS Marine Turtle Atlas.
This is a relational database that summarises the
breeding season, location and estimated size of
the annual breeding population by species for
marine turtles in Queensland. Analyses of these
data have been output as summary maps using
GIS software (Fig. 13).

Green turtle, Chelonia mydas. There are three
concentrations of C mydas rookeries in Queens-
land: southern GBR: northern GBR and adjacent
Torres Strait and southern Gulf of Carpentaria
with a fourth adjacent cluster of rookeries on the
Coral Sea Platform (Fig. 13 A). These data,
including the results of the on ground surveys of
turtle nesting throughout the northern GBR and
Torres Strait region (Tables 3-6), show that C.
mydas breeding is concentrated onto the cays of
the outer barrier reef from Bramble Cay to
Princess Charlotte Bay. The main concentration
within this outer barrier reef nesting zone occurs
on Raine Island (Table 3 ) and Moulter Cay (Table
4) with subsidiary concentrations on Bramble
Cay, Murray Islands (Fig. 13 A), No. 8 Sandbank
and No. 7 Sandbank (Table 5). C. mydas breeds
less frequently on the inner shelf sand cays, con-
tinental islands and adjacent mainland (Table 6).

Since 1974 hundreds of thousands of nesting
and recently dead adult C. mydas and their
hatchlings have been observed at Raine Island,
Moulter Cay, MacLennan Cay and No. 8 and
No. 7 Sandbanks. A total of 34,361 tagged female
C. mydas were processed while nesting at Raine
Island and 3,015 at the other outer barrier reef
islands (Tables 7-9). During the 32 trips on which
tagging and census studies were conducted by
QPWS and/or Applied Ecology, no adult turtle,
turtle track or hatchling of a species other than C.
mydas was recorded from Raine Island, Moulter
Cay, Maclennan Cay, No. 8 Sandbank or No. 7
Sandbank.

Hawksbill turtle, Eretmochelys imbricata.
There is one concentration of E. imbricata
rookeries in Queensland: on coral cays of the
inner shelf of the northern GBR from the Piper
Islands north to the sand islands and continental
islands of central and western Torres Strait (Fig.
13B: Table 10). Low density to sporadic nesting
occurs south of the Piper Islands to Princess
Charlotte Bay.

British Museum specimen, registration no.
46.7.27.6 is an immature specimen with CCL =
35.5cm collected at Raine Island (A.F. Stimson,
in litt., 26 May, 1977) which was presented to the
museum by Lieut, lnce in July 1846. This is the
Boulenger (1889) specimen V. It was on the
basis of this specimen that Stoddard et al. (1981 )
associated the species with Raine Island.
However, the latter study failed to recognise that
the turtle was an immature, presumably having
been caught on the surrounding reef at the time of
the construction of the beacon in 1 844. Immature
to adult-sized E. imbricata were occasionally
seen on Raine Island Reef during the present
study.

.1. Collins (JCUNQ) collected the skull from an
adult E. imbricata carcass within the central
depression of Raine Island in November 1973
(Limpus, 1980a). As only nesting female turtles
venture this far inland, this must be accepted as a
breeding record for the species from Raine
Island. During the January-February 1959
harvest of C. mydas from Raine Island by the
crew of the Trader Horn , ‘hawksbill turtles’ as
well as the abundant green turtles were identified
as being ashore nesting (E. Gray, pers. comm.,
1983; N. Whittaker, pers. comm., 1983). E.
imbricata breeding at Raine Island and adjacent
outer barrier cays during the present study period
may have been unrecorded because very little
census work has been conducted during
January-March which is the peak of the region’s
E. imbricata nesting season (Dobbs et al., 1999).
At Bramble Cay, E. imbricata nests at very low
density mostly during the January-March period
(D. Carter, pers. comm.; C.J. Parmenter, unpubl.
data). E. imbricata breeds at very low density on
Raine Island and probably also on the adjacent
outer barrier reef cays.

Flatback turtle, Natator depressus. There are
three concentrations of N. depressus rookeries in
Queensland: on continental islands and mainland
beaches of the central east coast from Bundaberg
to Townsville; sand islands and continental
islands of western Torres Strait and the mainland

GREEN TURTLE POPULATIONS OF RAINE ISLAND

373

TABLE 3. List of study trips to Raine Island and other islands of the northern Great Barrier Reef: November 1 974
- December 200 L summarising the nightly counts of breeding female turtles and their associated mortality
while ashore nesting on the islands. The nightly count of turtles is listed as the tally count or in parenthesis as a
tagging census count. The mean nightly count was calculated on half monthly sampling intervals. Nightly
mortality includes those turtles found alive but which would have died if left unaided and which resulted from
that night’s nesting attempt. Accumulated dead turtles were counted at the arrival of the study team for a census
period. Only Chelonia mydas was recorded. # denotes that the count may include more than one night’s
nestings, tr = tracks. Key to notes: 1 = JCUNQ, A. Birtles; 2 = C. Limpus & B. Cropp; 3 = G. Goedon (pers.
comm.); 4 = Kowarsky, 1978; 5 = QPWS; 6 = B. Cropp (pers. comm.); 7 = QPWS & AE; 8 = QP\VS aerial
survey; 9 = AIMS, D. Williams; 1 0 = QPWS/RIC; 1 1 = D. Williams; 1 2 = QPWS & GBRMPA; 1 3 = GBRMPA,
J. O’Dyer; 14 = Operation Raleigh; 15 = A. Fleay (in litt.).

Date

Nightly count

No. ashore

Nightly count

Death rate

-a

Mean nightly female
count

Notes

Accumulal

dead turtle

Mean

SD

n

9-23/6/80

.

0-1

_

14

5

8/12/80

71

(83)

0

0

59

17.5

5

5

9/12/80

68

(81)

10/12/80

58

11/12/80

69

12/12/80

29

(80)

28/2/81

_

-

(24)

12.7

7

9

1/3/81

(3)

2/3/81

-

3/3/81

o

Cl

_

4/3/81

(36)

.

5/3/81

(27)

.

6/3/81

(36)

_

7/3/81

(30)

.

8/3/81

-

9/3/81

(10)

_

7/6/81

0

0

_

(0.4)

0.5

9

10

8/6/81

0

0

9/6/81

0

0

10/6/81

(1)

0

1 1/6/81

(1)

0

12/6/81

0

0

13/6/81

(1)

0

14/6/81

(1)

0

15/6/81

0

0

16/6/81

(1)

0

(0.3)

-

3

17/6/81

0

0

18/6/81

0

0

12/11/81

1019

0

6

1009

-

3

10

13/11/81

0

14/11/81

1103

2

15/11/81

906

0

16/11/81

955

5

2038

446.7

13

17/11/81

0

18/11/81

1722

1

19/11/81

2074

4

1 20/11/81

1943

5

Date

Nightly count

No. ashore

Nightly count |
Death rate

1 .

Mean nightly female
count

j

Accumulal

dead turtle

Mean

SD

n

29/11/74

11565

281

11565

-

1

i

28/2/75

427 tr

>588

447 tr

-

4

2

1//3/75

517 tr

2/Z3/75

408 tr

3//3Z75

435 tr

22/10/75

-50 tr

3

29/1/76

-80 tr

4

2/12/76

-

872

-

2

5

3/12/76

-

4/12/76

898

5/12/76

-

6/12/76

845

Early/ 1 1/
77

-40-5

0

6

3/12/77

(23)

0

0

(32)

-

5

7

4/12/77

(23)

0

5/12/77

09)

0

6/12/77

(41)

1

7/12/77

(53)

-

1/12/78

(148)

0

1

(243)

-

5

7

2/12/78

(204)

0

3/12/78

(240)

1

4/12/78

(327)

1

5/12/78

(295)

-

29/10/79

-

0

18

416

-

2

5

30/10/79

381

0

31/10/79

451

1

1/11/79

410

0

427

-

2

2/11/79

-

0

3/11/79

-

0

4/11/79

444

0

5/11/79

-

0

6/11/79

-

0

7/1 1/79

-

1

8/11/79

-

1

9/11/79

-

0

13/12/79

1211

0

72

1347

-

2

5

14/12/79

1482

3

8 !

8/4/80

1-2 tr

374

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3 ( Cont .)

Date

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

Mean nightly female
count

Notes

Mean

SD

n

21/11/81

1986

0

22/11/81

1954

6

23/11/81

2565

2

24/11/81

2369

3

25/11/81

2307

2

26/11/81

1542

9

27/11/81

2263

7

28/11/81

2212

6

29/11/81

15

3011/81

2599

10

1/12/81

2757

6

2094

665.3

12

2/12/81

2345

13

3/12/81

1738

15

4/12/81

1287

8

5/12/81

1445

2

6/12/81

2

7/12/81

1394

2

8/12/81

1832

2

9/12/81

1795

8

10/12/81

2654

5

11/12/81

1817

8

12/12/81

2599

8

13/12/81

3463

8

21/2/82

350

350

1

11

13/7/82

0

0

189

(0.3)

-

3

10

14/7/82

0

0

15/7/82

(1)

0

16/7/82

0

0

(0.5)

0.8

14

17/7/82

(1)

0

18/7/82

0

0

19/7/82

(1)

0

20/7/82

0

0

21/7/82

(1)

0

22/7/82

(2)

'0

23/7/82

(2)

0

24/7/82

0

0

25/7/82

0

0

26/7/82

0

0

27/7/82

0

0

28/7/82

0

0

29/7/82

0

0

13/11/82

460

0

6

492

-

3

10

14/11/82

525

0

15/11/82

490

0

16/11/82

452

1

718

183.4

13

17/11/82

470

1

18/11/82

553

0

19/11/82

670

0

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

Mean nightly female
count

Notes

| Date

Mean

SD

n

20/11/82

639

1

21/11/82

716

0

22/11/82

674

0

23/11/82

670

1

24/1 1/82

822

0

25/11/82

854

0

26/11/82

760

0

27/11/82

0

28/11/82

0

29/11/82

1095

1

30/11/82

959

0

1/12/82

1000

2

925

112.5

6

2/12/82

0

3/12/82

878

0

4/12/82

810

0

5/12/82

809

1

6/12/82

960

2

7/12/82

1091

2

8/12/82

2

25/4/83

2 tr

0

82

3.2 tr

-

5

10

26/4/83

2 tr

0

27/4/83

8 tr

0

28/4/83

3 tr

0

29/4/83

1 tr

0

15/11/83

Several hundred tracks seen from air

12

3/4/84

8

0

92

9

4.2

13

10

, 4/4/84

9

0

i 5/4/84

6

0

6/4/84

11

0

7/4/84

5

0

8/4/84

3

0

9/4/84

15

0

10/4/84

17

0

1 1/4/84

8

0

12/4/84

7

0

13/4/84

7

0

14/4/84

6

0

15/4/84

14

0

16/4/84

3

0

5

1.9

9

17/4/84

5

0

18/4/84

5

0

19/4/84

4

0

20/4/84

2

0

21/4/84

3

0

| 22/4/84

7

0

23/4/84

5

0

24/4/84

8

0

1 25/4/84

0

GREEN TURTLE POPULATIONS OF RAINE ISLAND

375

TABLE 3 ( cont .)

Date

Nightly count

No. ashore

Nightly count
Death rate

Accumulated

dead turtles

Mean nightly female
count

Mean

SD

n

Notes

1/9/84

7 tr

7 tr

-

1

13

21/1184

7739

24

256

6376

-

5

10

22/1184

2

23/1184

6459

14

24/1184

15

25/1184

7074

1

26/1184

23

27/1184

5428

33

28/1184

19

29/1184

5179

11

30/1184

12

1/12/84

7867

8

6493

1683

8

2/12/84

23

3/12/84

9301

22

4/12/84

17

5/12/84

7070

12

6/12/84

29

7/12/84

7099

19

8/12/84

31

9/12/84

5633

35

10/12/84

21

11/12/84

5764

11

12/12/84

47

13/12/84

5295

26

14/12/84

29

15/12/84

3916

34

16/12/84

40

8873

2429

5

17/12/84

5102

52

18/12/84

52

19/12/84

9776

44

20/12/84

11467

25

21/12/84

9960

22/12/84

23/12/84

8059

24/11/85

4

10

25/1 1/85

119

0

196

-

3

26/11/85

0

27/11/85

308

0

28/11/85

0

29/1 1/85

162,
202 tr

0

30/11/85

0

1/12/85

214

0

227

38

6

2/12/85

0

3/12/85

205

0

4/12/85

0

5/12/85

0

6/12/85

227

0

Jl

Date

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

Mean nightly female
count

Notes

Mean

SD

n

7/12/85

0

8/12/85

295

0

9/12/85

1

10/12/85

236

0

11/12/84

5764

11

12/12/84

47

13/12/84

5295

26

14/12/84

29

15/12/84

3916

34

16/12/84

40

8873

2175

5

17/12/84

5102

52

18/12/84

52

19/12/84

9776

44

20/12/84

11467

25

21/12/84

9960

22/12/84

23/12/84

8059

24/11/85

4

10

25/11/85

119

0

196

-

3

26/11/85

0

27/11/85

308

0

28/11/85

0

29/11/85

162,
202 tr

0

30/11/85

0

1/12/85

214

0

227

38

6

2/12/85

0

3/12/85

205

0

4/12/85

0

5/12/85

0

6/12/85

227

0

7/12/85

0

8/12/85

295

0

9/12/85

1

10/12/85

236

0

11/12/85

0

12/12/85

0

13/12/85

183,
323 tr

1

14/12/85

0

15/12/85

0

16/12/85

105,
264 tr

1

151

-

3

17/12/85

2

18/12/85

142

0

19/12/85

205

0

20/12/85

0

376

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3 ( cortt .)

Date

Nightly count

No. ashore

Nightly count
Death rate

Accumulated

dead turtles

Mean nightly female
count

Notes

Mean

SD

n

17/10/86

137

130

-

2

14

18/10/86

123

13/11/86

1439

0

1549

-

3

14/11/86

1705

0

15/11/86

1503

0

16/11/86

1284

0

3145

-

5

23/11/86

3142

11

90

10

24/11/86

25/11/86

3208

23#

26/11/86

7

27/11/86

3525

7

28/11/86

7

29/11/86

4565

8

30/11/86

8

1/12/86

4696

8

3957

686

8

2/12/86

13

3/12/86

4547

6

4/12/86

9

5/12/86

3991

8

6/12/86

9

7/12/86

4838

11

8/12/86

18

9/12/86

3648

12

10/12/86

7

11/12/86

3643

6

12/12/86

12

13/12/86

12

14/12/86

3382

22

15/12/86

2910

-

29/11/87

10

158

10

30/1 1/87

3296

10

3296

1

1/12/87

2312

9

4099

-

4

2/12/87

19

3/12/87

4489

Ur

4/12/87

4585

14

5/12/87

5011

29/11/88

1

52

10

30/11/88

2

1/12/88

516

1

731

-

4

2/12/88

1

3/12/88

1

4/12/88

1088

2

5/12/88

671

0

6/12/88

2

7/12/88

647

0

28/11/89

24

177

5

29/1189

5919

15

5919

-

1

30/1189

16

Date

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

Mean nightly female
count

Notes

Mean

SD

n

1/12/89

16

6144

-

3

2/12/89

18

3/12/89

19

4/12/89

14

5/12/89

5991

14

6/12/89

5925

14

7/12/89

6516

14

29/11/91

2289

20

-

3571

3

6

10

1 30/11/91

_

6

1/12/91

3360

8

2/12/91

.

4

3/12/91

3483

8

4/12/91

-

13

5/12/91

3492

16

6/12/91

_

6

7/12/91

3741

33

1 8/12/91

.

36

9/12/91

5062

20

10/12/91

.

6

29/11/92

850

3

-

1323

458

6

10

30/11/92

.

3

1/12/92

874

12

2/12/92

.

8

3/12/92

1048

8

4/12/92

7

5/12/92

1591

6

6/12/92

-

5

7/12/92

1936

8

8/12/92

.

8

9/12/92

1638

5

10/12/92

8

29/11/93

9245

11

-

8593

585

6

10

30/1 1/93

-

33

1/12/93

9012

27

2/12/93

-

11

3/12/93

7596

19

4/12/93

.

0

5/12/93

8384

10

6/12/93

-

20

7/12/93

8478

5

8/12/93

8

9/12/93

8841

20

10/12/93

_

22

30/11/94

_

0

-

1/12/94

586

0

645

-

5

l°J

2/12/94

.

0

3/12/94

836

0

_l

GREEN TURTLE POPULATIONS OF RAINE ISLAND

377

TABLE 3 (Cont.)

Date

Nightly count

No. ashore

Nightly count
Death rate

Accumulated

dead turtles

Mean nightly female
count

Notes

Mean

SD

n

4/12/94

-

1

5/12/94

729

1

6/12/94

.

1

7/12//94

404

0

8/12/94

_

0

9/12/94

671

0

5/12/95

4653

5

191

4648

-

5

10

6/12/95

_

8

7/12/95

4627

9

8/12/95

5765

15

9/12/95

4044

14

10/12/95

.

0

11/12/95

4150

4

12/12/95

-

21

13/12/95

-

6

30/11/96

11755

67

150

12100

1421

7

10

1/12/96

_

51

2/12/96

-

52

3/12/96

11454

75

4/12/96

11422

29

5/12/96

10677

58

6/12/96

-

65

7/12/96

11209

42

8/12/96

13667

62

9/12/96

14519

26

29/11/97

3184

11

186

4065

905

7

10

30/1 1/97

3279

6

1/12/97

3881

4

2/12/97

3292

10

3/12/97

4312

10

4/12/97

-

14

5/12/97

5284

7

6/12/97

5223

5

28/11/98

110

1

35

472

245

7

10

29/1 1/98

300

0

30/11/98

.

2

01/12/98

762

0

2/12/98

707

1

3/12/98

-

0

coast of north-western Cape York Peninsula and
the Wellesley Island of the southeastern Gulf of
Carpentaria (Fig. 13C). The on-ground surveys
of turtle nesting throughout the northern GBR,
recorded only sparse N. depressus nesting (Miller
et al„ 1995; Dobbs et al„ 1999) (Table 6).

Date

Nightly count

No. ashore

Nightly count

Death rate

T3

P C/5

Mean nightly female
count

Notes

Accumulat

dead turtle;

Mean

SD

n

4/12/98

567

0

5/12/98

583

0

6/12/98

_

1

7/12/98

.

0

8/12/98

-

0

9/12/98

273

0

28/11/99

5806

41

213

5617

-

3

10

29/11/99

5834

18

30/11/99

5212

13

1/12/99

.

9

7079

776

8

2/12/99

7010

14

3/12/99

20

4/12/99

6282

17

5/12/99

6848

25

6/12/99

6745

36

7/12/99

8101

25

8/12/99

_

35

9/12/99

6301

24

10/12/99

8408

32

11/12/99

7077

41

30/1 1/00

114

1

0

114

-

1

10

1/12/00

82

0

131

31

7

2/12/00

_

0

3/12/00

136

0

4/12/00

136

0

5/12/00

.

0

6/12/00

107

0

7/12/00

171

0

8/12/00

165

1

9/12/00

122

0

1/12/01

4539

20

71

3601

1401

8

10

2/12/01

3194

5

3/12/01

2462

6

4/12/01

_

10

5/12/01

1923

0

6/12/01

2675

2

7/12/01

3072

21

8/12/01

4988

15

9/12/01

_

15

10/12/01

5952

4

Stoddart et al. (1981) suggested the possibility
that N. ( =Chelonia ) depressus nested at Raine
Island. Their description of the turtles in question
is not consistent with that of adult N. depressus
but fits well within the normal variation in
carapace shape of C. mydas. In view of the lack of

378

MEMOIRS OF THE QUEENSLAND MUSEUM

supporting evidence for the presence of N.
depressus at Raine Island, the species should not
be considered part of the nesting community of
Raine Island or the adjacent outer barrier cays.

Loggerhead turtle, Caretta caretta. C. caretta
has not been recorded breeding in the far northern
GBR. An unsuccessful nesting attempt by the
species was recorded from Lizard Island prior to
this survey (Limpus 1 982). This remains the only
eastern Australian breeding record for the species
from north of Townsville.

Misidentijied species . The Leatherback turtle,
Dermochelys coriacea was recorded ashore at
Moulter Cay and Raine Island on 8th and 16
November 1986 among the nesting C. mydas
(Anon., 1986). The turtles were not described in
detail, nor were they measured or photographed.
Given that the recorders were naive with respect
to marine turtle identification, that during the
same breeding season the species was not
encountered by experienced observers at these
islands, and that the species has not been
recorded by any other observers at these islands
at any other time, D. coriacea will not be
considered as part of the marine turtle fauna of
the Raine Island area. No records ofD. coriacea
breeding were obtained during the surveys of
turtle breeding throughout the remainder of the
northern GBR and Torres Strait (Miller et al.,
1995; Limpus et al., 2000) (Tables 5-6).

CHELONIA MYDAS BREEDING SEASON.
Ellis (1936) in summarising his stay on Raine
Island from late 1890 to 1892. identified
pedominantly summer nesting by green turtles.
The next longest reported habitation of the island
was during the construction of the beacon from
29 May until mid September 1844 when
MacGillivray (1846) reported only irregular
isolated nestings by green turtles with numbers
increasing in September. All remaining reports
are from trips of about a month or less in duration
and at irregular intervals. The nightly density of
turtle nesting recorded on all visits to Raine
Island and other islands of the northern GBR
since November 1 974 are summarised in Tables
3-5.

The historical data and that of Tables 3-4 have
been combined to produce a summary of the
breeding season by months for Raine Island and
Moulter Cay (Table 11). While there are marked
differences in nesting density from year to year,
the general pattern is for sporadic nesting to occur
during the dry season (May - August) and
breeding turtle numbers to increase from

TABLE 4. Moulter Cay. Only Chelonia mydas was
recorded nesting at Moulter Cay. See Table 3 for
symbols and codes.

Date

Nightly count

No. ashore

Nightly count

Death rate

1 Accumulated

dead turtles

Mean nightly female
count

Mean

SD

n

Notes

5/3/75

244 tr

0

142

244 tr

_

1

2

7/12/76

(430+)

7

5

2/12/77

13 tr

0

0

13 tr

-

1

7

24/6/80

0

0

28

10

19/6/81

0

10

30/7/82

0

0

10

10/12/82

402 tr

3

402 tr

-

1

10

14/12/85

148

0

3

157

-

2

10

15/12/85

166

0

17/10/86

58

76

40

7

14

18/10/86

55

21/10/86

61

22/10/86

88

j 23/10/86

74

27/10/86

38

28/10/86

160

6/11/86

210

357

108

3

' 7/11/86

407

8/11/86

458

16/11/86

747

747

-

1

14/12/86

1669

1807

-

2

10

15/12/86

1945

4/12/88

627

3

17

627

-

1

10

13/12/92

180 tr

0

0

180 tr

-

1

10

7/12/93

5057

-

5057

-

1

10

7/12/95

2495

0

39

2495

-

1

10

27/1 1/96

4043

-

-

4043

.

1

10

7/12/97

1671

6

45

1624

.

2

10

8/12/97

1577

2

10/12/98

-

0

21

2

10

11/12/98

-

0

12/12/99

3665

31

60

3665

-

1

10

10/12/00

67

1

67

-

1

10

10/12/01

1881

-

42

2022

-

2

10

1 11/12/01

2163

3

September and reach a maximum in December
and January. Numbers decrease again into the dry
season. This also is the pattern for C. mydas
nesting elsewhere in the northern GBR: at
Bramble Cay to the north (Limpus et al., 2001)
and at No. 7 and No.8 Sandbanks to the south
(Table 5). In the southern GBR, C. mydas nesting
is seasonal, being confined to the mid summer
(October - April) period (Bustard, 1 972; Limpus,

GREEN TURTLE POPULATIONS OF RAINE ISLAND

379

TABLE 5. Other outer Barrier Reef coral cays of the far northern Great Barrier Reef. Only Chelonia mydas was
recorded at these islands. See Table 3 for symbols and codes.

Rookery

Date

Nightly

count

No.

ashore

Nightly

count

Death

rate

Accumul

ated

dead

turtles

Notes

Bramble

Cay

3/1/87

340 tr

5

4/1/87

290 tr

16/1/89

(272)

0

0

5

17/1/89

(MO)

0

Maclennan

Cay

30/11/78

(4)

0

0

7

25/6/80

0

0

0

10

19/6/81

0

0

10

30/7/82

0

0

0

10

4/12/88

(8)

0

0

10

No.8

Sandbank

28/11/76

*-65 tr

0

0

5

29/11/76

25 tr

0

1 1/12/79

1 14 tr

0

0

5

5/6/80

0

0

1

10

25/6/81

0

0

10

10/7/82

0

0

10

11/12/82

(118)

10

17/12/86

(102)

0

10

28/1 1/87

193 tr

0

10

27/1 1/88

(30)

0

0

10

24/1 1/89

234

0

0

5

27/11/91

79 tr

0

0

10

27/1 1/92

63 tr

0

0

10

3/12/95

121 tr

0

0

10

11/12/96

-

0

2

10

10/12/97

119

0

2

10

No.7

Sandbank

28/11/76

-60 tr

0

0

5

11/12/79

108# tr

0

4

5

4/6/80

0

0

3

10

1980b). The more tropical location of the Raine
Island rookery and the extremely large numbers
of breeding females at Raine Island would give a
higher chance of a few individuals nesting
outside the main breeding season and could be
the cause of the expanded breeding season
compared to elsewhere in the GBR.

There is considerable variability in the timing
of C. mydas breeding throughout the Australian
continental shelf. Along the eastern and western
Australian coasts, C. mydas nesting is principally
mid-summer. Within the southern Gulf of
Carpentaria (Bountiful, Pisonia and Rocky
Islands) and Arafura Sea (southern islands of Aru
Archipelago), C. mydas nesting occurs year
round with a midyear peak (Bustard, 1972;
Schulz, 1989; CJ. Limpus, unpub 1. data). A
similar variability in breeding season occurs for
N. depressus at its various rookeries around

Rookery

Date

Nightly

count

No.

ashore

Nightly

count

Death

rate

Accumul

ated

dead

turtles

Notes

No.7

Sandbank

17/11/81

205# tr

0

0

10

17/12/86

!_(39)

0

10

28/11/87

94 tr

0

0

10

27/11/88

(26)

0

0

10

23/11/89

1 10 tr

0

0

5

27/11/91

94 tr

1

0

10

27/1 1/92

53 tr

0

0

10

3/12/95

145 tr

0

1

10

11/12/96

-

0

6

10

10/12/97

108

0

7

10

No.6

1 Sandbank

5/6/80

0

0

10

4/6/81

0

0

10

No.l

Sandbank

14/11/84

13 (tr)

0

10

Ham Cay

12/12/79

61# tr

0

5

5/6/80

0

0

0

10

16/11/81

Many tr

0

0

10

11/12/82

Many tr

0

0

10

25/11/89

10 tr

0

0

5

Derry Cay

5/6/80

0

0

0

10

12/12/82

12 tr

0

1

10

25/11/89

-12 tr

0

0

5

Davie Islet

27/6/81

0

0

10

12/12/82

14 tr

0

0

10

23/11/89

22 tr

0

0

5

Tydeman

Cay

27/6/81

0

0

10

12/12/82

29 tr

0

0

10

18/12/86

11 tr

0

10

northern Australia (Limpus, 1971; Limpus et al.,
1983b, 1989).

This variability in timing of the nesting season
probably results from the interaction of a number
of factors which should include the temperature
environment of the rookery (Guinea, 1994), the
genetic origin of the breeding population (which,
amongst other things, should influence the
pivotal temperature for the individual
population) and the climate of the feeding
grounds providing turtles to the rookeries. The
direct cause(s) for this variability in breeding
season cannot be explained at present. It is not
simply a function of latitude, given the similarity
of the breeding seasons in northern and southern
GBR and the different breeding season of the
southern Gulf of Carpentaria rookeries at an
intermediate latitude. The issue is further
complicated by turtles from widely scattered

380

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 6. Coral cays, continental islands and mainland beaches of the inner shelf of the northern Great Barrier
Reef. Cm = Chelonia mydas; Ei = Eretmochelys imbricata ; Nd = Natator depressus; ? = species unidentified.
See Table 3 for other symbols and codes.

Rookery

Date

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

''

Notes

| Sinclair Island

11/12/76

0

0

5

Milman Island

11/12/76

(6) Ei

0

5

5/12/78

~ 1 0 tr Ei

0

7

31/7/82

-16# tr
Ei

0

10

13/6/84

~5# tr Ei

0

5

Cholmondely

Island

1/8/82

3# tr Ei

0

10

Wallace Island

7/12/76

4 tr Ei

0

- 5

09/12/76

0

0

1/8/82

1# ? 2#
tr Ei

0

0

10

Mainland coast
south of Usher
Point

10/12/76

0

0

5

Little Boydong
Island

9/12/76

0

0

5

Boydong

Island

9/12/76

(2) Ei

0

0

5

1/8/82

8# tr Ei

0

0

10

Hannibul

Island

6/3/75

1 trEi

0

2

Bird Island
(north)

5/3/75

(4) Ei

0

0

2

8/12/76

(1) Cm
(3) Ei

0

0

5

21/6/81

1 trEi

0

10

13/6/84

1# tr Ei

5

Bird Island
(south)

21/6/81

1 tr Ei

0

10

Macarthur

Island

6/3/75

1 trEi

0

2

Saunders
| Island

5/3/75

2 trEi

0

0

2

21/6/81

1 tr Ei

0

10

2/8/82

1 tr Ei

0

1 Ei

10

Ellis Island

11/11/79

0

0

5

Magra Island

22/6/81

3# tr Ei

0

10

Ashmore

Banks (north

1/12/76

2 tr Cm

1 tr Ei

0

5

western)

7/8/80

1 Ei

0

10

6/6/81

0

0

10

Ashmore

Banks (north
eastern)

7/6/81

0

0

10

Ashmore

7/6/80

1 tr/Cm

0

0

10

Banks (north
eastern)

7/6/81

0

0

10

Farmer Island

23/4/83

0

0

10

20/12/85

2 trEi

0

0

10

21/12/85

(6) Ei

2/6/86

1 trEi

0

10

28/11/87

2 trEi 2#
tr Cm

0

0

10

29/11/87

_ (5iEi_,

0

0

10

Rookery

Date

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

Notes

Sir Charles
Hardy Island
(northern)

26/10/79

3# tr ?

0

5

26/6/80

1# tr Ei

0

0

10

11/11/81

1 tr Cm
(l)Ei

0

0

10

19/12/85

1# trCm
4# tr Ei

0

0

10

5/12/87

9# tr Cm
2# tr Ei

0

10

Sir Charles
; Hardy Islands
(western)

2/12/76

1 1# tr ?

0

5

26/10/79

13# tr?

0

5

28/7/82

0

0

10

23/4/83

0

0

10

19/12/85

1# tr?

0

0

10

5/12/87

1 1# tr ?
2# trEi

0

10

Haggerstone

Island

10/11/79

0

0

5

22/6/81

0

0

10

23/6/81

0

0

15/11/81

1 tr Ei

0

0

10

28/7/82

0

0

10

30/4/83

0

0

10

5/12/87

0

0

10

Forbes Island
(north)

15/11/81

0

0

0

10

Portland Roads

30/1 1/76

0

0

5

30/4/83

0

0

10

Quoin Island

6/6/80

0

0

0

10

24/6/81

0

0

10

15/11/81

0

0

0

10

26/11/89

0

0

5

Chapman

Island

17/12/79

0

0

5

17/11/84

0

0

10

Sherrard Island

17/11/84

1# tr?

0

10

6/12/87

1# trCm
3# tr Ei

0

0

10

Night Island

25/1/79

0

0

5

9/11/81

0

0

0

10

9/11/82

0

0

10

12/6/84

0

0

5

16/11/84

0

0

10

Morris Island

6/12/87

13# tr
Cm 12#
tr Ei

0

0

10

Fife Island

28/11/76

3 trEi

0

5

9/11/82

0

0

0

10

12/6/84

0

0

0

5

6/12/87

6# tr Ei

0

0

10

| Lowrie Island

9/1 1/82

0

0

0

10

GREEN TURTLE POPULATIONS OF RAINE ISLAND

381

TABLE 6 ( Cont .)

Rookery

Date

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

Notes

Hay

Island

9/11/82

0

0

0

10

16/11/84

2# tr Ei

0

0

10

Hannah

Island

27/11/76

0

0

0

5

9/11/82

0

0

0

10

15/11/84

0

0

0

10

Pelican

Island

28/11/76

0

0

0

5

26/6/81

0

0

0

10

9/11/81

1 tr Ei 1
Nd

hatchling

0

0

10

16/11/83

1# tr Cm
2# tr Ei

0

12

7/12/87

2# tr Ei

0

0

10

Stainer

Island

1 1/6/84

0

0

0

5

15/11/84

3# tr Ei

0

0

10

7/12/87

0

0

0

10

King

Island

14/11/84

0

0

10

Clack

Island

19-21/12/

86

0

0

0

10

Stanley

Island

25/2/75

0

0

2

18/11/81

0

0

0

10

Flinders

Island

24/10/79

0

0

5

18/11/81

0

0

0

18

15/11/84

0

0

10

Pipon

Island

22/2/75

0

0

0

2

29/6/81

0

0

10

14/11/84

0

0

10

rookeries occurring together in feeding grounds
in northern Australia and adjacent eastern
Indonesia (Limpus et al., 1992). For example,
adult female C. mydas have been caught off
Coburg Peninsula in the Northern Territory that
had been tagged while nesting at Raine Island,
Lacepede Islands in Western Australia and south
eastern Java; others have been caught in the Kei
Islands of Indonesia that had been tagged while
nesting at Raine Island and at the Turtle Islands of
Sabah,

HOW LONG HAS RAINE ISLAND BEEN A
CHELONIA MYDAS ROOKERY? Prior to the
last Holocene transgression of some 10,000 yr
ago, a turtle swimming up to the reef on which
Raine Island now lies would have been
confronted with precipitous cliffs rising to
perhaps 45m or more above the then sea level

I Rookery

Date

Nightly count

No. ashore

Nightly count

Death rate

Accumulated

dead turtles

Notes

Cape

Melville

12/11/79

0

0

5

18/11/81

0

0

0

10

8/12/87

0

0

10

Stapleton

Island

29/6/80

0

0

10

13/11/84

2# tr Cm

0

0

10

Coombes

Island

14/11/84

0

0

0

10

Noble Is-
land

18/11/81

0

0

0

10

Rhodda Cay

27/6/81

0

0

10

Wilson Cay

27/6/81

0

0

10

Howick

Island

23/10/79

0

0

5

Holm Island

14/11/79

0

0

5

Barnard

Island

14/11/79

0

0

5

Eagle Islet

27/6/81

0

0

10

North

Direction

Island

28/6/81

0

0

10

South

Direction

Island

28/6/81

0

0

10

Rocky Islet

28/6/81

0

0

10

14/12/82

0

0

0

10

Three Isles

21/2/75

0

0

2

Low

Wooded Isle

3/6/80

0

0

10

Low Island

22/10/79

0

0

5

Michelmas

1 Ca y

1/6/80

0

0

10

30/6/81

0

0

10

with no beaches suitable for turtle nesting.
Indeed, there may have been no suitable beaches
for turtle nesting anywhere in the vicinity. The
present day Raine Island and the other islands of
the northern outer Great Barrier Reef have
formed since sea levels stabilised at an upper
level approximately 6,000 yr ago and slowly
receded about 1 m to their current level more than
4,000 yr ago (Hopley, 1983; Chappel, 1983).

Clam shells ( Tridacna sp. and Hippopus
hippopus) embedded in the surface of the
phosphatic limestone platform ol Raine Island
have been carbon dated to a calibrated age B.P. of
1 1 30 ±21 Oyr at the western end of the island and
as iow as 590 - 630 ± 70yr at the eastern end of the
island adjacent to the tower (Limpus, 1987;
Polach et al., 1978). Deeper layers of rock are
presumed to be older. The other sand cays of the

382

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 14. Fossil remains of an adult sized Chelonia mydas
(femur) found at the western end of Raine Island in a
bedding plane approximately 60cm below a clam
carbon dated to 1 130yr B.P.

northern outer barrier reef are presumed to be of
similar age to Raine Island.

Turtle eggs in nests fossilised in situ were
reported during the quarrying of the phosphatic
limestone platform at the eastern end of the island
for the construction of the beacon (MacGillivray,

1 846; Jukes, 1 847). This quarry site is adjacent to
the beacon and within 50m of the collection site
of the clams dated 590-630yr BP. These clutches
of turtle eggs must have been laid at least 460 yr
before their discovery in 1 844.

The weathered fossil remains of an adult-sized
C. mydas that had died plastron down and whose
skeletal remains were relatively undisturbed
prior to being covered by sand were found in a
horizontal bedding plane approximately 60cm
below the clam dated at 1 1 30yr BP. (Limpus,
1987) (Fig. 14). Fragments of turtle bone and bird
bone can be found in almost every large
phosphatic limestone rock at the margin to the
central rock platform of Raine Island.

Death of nesting turtles ashore for nesting
attempts has been observed annually on Raine
Island during the recent studies and in seasons of
high nesting density there may be >1,000
skeletons added to the islands sediments (Table
6). Most of these are scattered as individual bones
by subsequent nesting turtles. Only the skeletons
of turtles that die in the central depression of the
rock platform are likely to be left relatively
undisturbed (Fig. 15). It is also only during
seasons of high nesting density that numerous
turtles enter the central depression and die there
of heat exhaustion. Raine Island and Moulter Cay
are the only turtle rookeries of the GBR where
nesting turtles die in large numbers and where

FIG 15. Skeletal remains of an adult female Chelonia
mydas that had been disoriented and died of heat
exhaustion in the central depression of Raine Island.
This carcass, photographed in December 1987,
originated from a turtle that died during the 1984-85
breeding season. Skeletal remains in this central area of
the island have the potential for being left undisturbed
for years and subsequently buried.

turtle bones are a characteristic of the beach
sediments. This turtle mortality is a function of
the high density turtle nesting that occurs on these
islands. The first accounts of Raine Island from
1 843-4 prior to quarrying for construction of the
beacon (MacGillivray, 1846; Jukes, 1847)
described Raine Island with its turtle bones and
dead turtles in a manner consistent with its
appearance today. Numerous turtle remains and
dead and dying turtles have impressed sub-
sequent visitors to Raine Island in 1 845 (Allen &
Corris, 1977), 1874(Moseley, 1879), 1890(EIIis,
1936), 1913 (MacGillivray, 1917) and c. 1953-6
(Fig. n.

Taken together these observations indicate that
Raine Island has supported a high density C.
mydas rookery with associated frequent adult
turtle mortality for long before its discovery by
Europeans and indeed, for at least 1 1 90 ± 2 1 0 yr
B.P. (Limpus, 1987). However turtles could not
have been using the island for nesting for more
than about 4000 yr (the time that a sand island
could have existed on this reef). This is the only
marine turtle rookery in Australia with long-term
evidence of nesting prior to European colonisation.
Because of this fossil evidence for past nesting,
Raine Island has the longest continual record of
use as a marine turtle rookery anywhere in the
world (Groombridge & Luxmore, 1 989).

The origin of the turtles that colonised Raine
Island after its formation perhaps 4000 yr ago has
not been elucidated as yet by genetic studies of

GREEN TURTLE POPULATIONS OF RAINE ISLAND

383

TABLE 7. Numbers of adult female Chelonia mydas tagged and recaptured while nesting at Raine Island
1 974-2001. AE indicates turtles tagged by Applied Ecology P/L. All other tags applied by QP\VS. OR denotes
tagged turtles recorded during Operation Raleigh. * denotes migration recaptures of nesting turtles previously
tagged at foraging area. In no season was an attempt made to tag the entire nesting population. For remigrant
turtles that changed rookeries between breeding seasons, the original rookery was identified: CS = Coral Sea
Platform; MC = Moulter Cay; MM = Milman Island; MP = Moore Park; N7 = No. 7 sandbank; N8 = No. 8
Sandbank; RI = Raine Island.

Breeding

season

Month

New tags

Within

season

rookery

change

Remigrant

New turtles to
rookery

Total turtles
with tags

Same rookery

Rookery

change

Retag

1974-75

Feb-Mar

151

151

151

1975-76

Nil

1976-77

Dec

2420

15 AE

0

0

0

0

2435

2435

1977-78

Dec

53

61 AE

0

0

0

0

114

114

1978-79

Dec

490

398 AE

0

0

0

0

888

888

1979-80

Oct-Dec

1766

0

1

0

0

1766

1767

June

1

0

0

0

0

1

1

1980-81

Dec

284

0

0

0

0

284

284

June

1

0

0

0

0

1

1

1981-82

Nov- Dec

2589

0

8

1 AE

1 MC

4

2591

2603

Jul

2

0

0

0

0

2

2

1982-83

Nov-Dec

1586

0

8

0

7

1586

1601

April

6

0

0

0

0

6

6

1983-84

Apr

80

0

0

0

0

80

80

1984-85

Nov- Dec

2458

2*

0

59

0

25

2460

2544

1985-86

Nov- Dec

1853

1*

2 MC

31

1 N8

30

1857

1918

1986-87

Nov- Dec

1447

0

12 OR

80

1 N8

177

1460

1717

1987-88

Jul- Aug

4

0

0

0

0

4

4

1988-89

Nov-Dec

1859

0

63

0

29

1859

1951

1989-90

Dec

769

0

264

0

4

769

1037

1990-91

Dec

1210

0

148

1 MC

24

1211

1383

1991-92

Dec

980

0

227

0

17

980

1224

1992-93

Dec

1689

0

129

0

14

1689

1832

1993-94

Dec

1593

6*

0

152

1 N7

5

1600

1757

1994-95

Dec

991

0

31

1 N7

1

992

1024

1995-96

Dec

1485

1*

0

136

3 MC

43

1489

1668

1996-97

Nov-Dec

1664

9*

0

289

2 MC

28

1675

1992

Jan-Feb

796

1*

0

91

3N8

5

800

896

1997-98

Nov-Dec

1022

5*

0

387

1 N8

1 CS

11

1029

1427

1998-99

Nov-Dec

1599

0

156

0

31

1599

1786

1999-00

Nov-Dec

1160

17*

0

847

7 MC

3 MM
3N7

1 N8

29

1191

2067

—

2000-01

Nov-Dec

664

1*

0

72

0

19

665

756

2001-02

Dec

1145

4*

0

443

3N8

2 M

2N7

1 MM

1 MP

27

1158

1628

Total

33817

474 AE

47*

2MC

12 OR

3622

1 AE

16 MC

10 N8
7N7

4 MM

1 MP

1 CS

530

34392

38544

384

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 8. Numbers of adult female Chelonia mydas tagged and recaptured while nesting at Moulter Cay,
northern GBR, 1974-2001. See Table 7 for definition of codes.

Breeding

season

Month

New tags

Within

season

rookery

change

Remigrant

Total turtles
with tags

Same rookery

Rookery

change

Retag

New turtles to
rookery

1974-75

Mar

6

0

0

0

0

6

6

1976-77

Dec

425

5RI

0

0

0

430

430

1977-78

Dec

64 AE

0

0

0

0

10

10

1982-83

Dec

2

0

0

0

0

2

2

1984-85

Dec

1 RI

1

1

1985-86

Dec

131

11 RI

0

0

0

142

142

1986-87

Dec

174

8RI

1

2 RI

11

184

196

1988-89

Dec

98

1 RI

0

2 RI

0

101

101

1989-90

Dec

72

0

1

0

0

72

73

1990-91

Nil

1991-92

Nil

1992-93

Dec

48

0

2

1 RI

2

49

53

1993-94

Nil

1994-95

Nil

1995-96

Dec

94

2 RI

2

1 RI

1 N7

3

98

103

1996-97

Nil

1997-98

Dec

109

13 RI

1

5 RI

1 MM

0

128

"

1998-99

Dec

194

2 RI

0

1 RI

1

197

198

1999-00

Dec

132

3 RI

1

7 RI

1 MM

0

143

144

2000-01

Dec

81

0

0

1 RI

2

82

84

2001-02

Dec

272

5 RI

5

0

2

77

291

Total

1844

4 AE

50 RI

13

21 RI

1 N7

2 MM

21

1922

world wide C. mydas populations (Bowen et al.,
1989; Norman et al., 1994a,b). Given the
precipitous nature of the coastline for that part of
Australia in pre Holocene times, the present day
rookeries are not the result of a gradual shift of
beaches with rising sea levels. Entirely new beaches
became available for colonisation following the
sea level rise while others, presumably further
afield, were submerged. This emphasises the
capacity for C. mydas to colonise new rookeries
that were not natal beaches in response to slow
changes in sea level. The process by which turtles
locate and/or choose a new rookery site following
the elimination of an existing one has not be
addressed by present studies.

COURTSHIP. C. mydas courtship observed during
expeditions to Raine Island is summarised as
follows:

1 979. 29 October - 9 November: Mating pairs of C. mydas
were a comon sight in the deep water just outside the reef
edge; two pairs of mating C. mydas were captured while
stranded on the reef flat. 13-14 December: No courting

turtles were observed, although the females were still
aggregated in large numbers over the reef and beyond the
reef edge.

1980. 8-13 December: No turtle courtship observed on or
near Raine Island Reef.

1 982. 1 3-3 1 July: One mounted pair of C. mydas observed
adjacent to Raine Island Reef. 11 November - 10
December Courting C mydas were observed adjacent to
Raine Island reef on only a few occasions in mid
November.

1984. 3-26 April: No courtship observed on or near Raine
Island Reef. Vertical aerial photographs of Raine Island
from October 1984, commissioned by the Raine Island
Corporation, show hundreds of laige turtles aggregated in
the shallows of the reef flat adjacent to the island. The
aggregation in these images resembled of aggregations of
mating C. mydas observed by the authors at rookeries in the
southern Great Barrier Reef and the Wellesley Group in the
southern Gulf of Carpentaria.

1985. 9 & 16 December. During underwater observations,
adult male turtles were seldom seen (2 males out of 36 C.
mydas examined). One male had recent courtship damage
(Limpus, 1993) to tail and flippers.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

385

TABLE 9. Adult female Chelonia mydas tagged and recaptured while nesting at other outer barrier cays in the
northern GBR, 1974-2001. See Table 7 for definition of codes.

—

Within

Remigrant

Breeding

season

Month

New tags

season

rookery

change

Same rookery

Rookery

change

Retag

New turtles to
rookery

Total turtles
with tags

Maclennan Ca>

1978-79

Dec

2 2 AE

0

0

0

0

4

4

1988-89

Dec

8

0

0

0

0

8

8

Subtotal

10 2 AE

0

0

0

0

12

12

INo 7. Sandbank

1976-77

Dec

1

0

0

0

0

1

1

1986-87

Nov

74

0

0

0

0

74

74

1987-1988

Nov

62

0

0

0

0

62

62

1988-89

Nov

26

0

0

0

0

26

26

1990-91

Dec

12

0

0

0

0

12

12

1991-92

Dec

89

0

0

0

0

89

89

1992-93

Dec

53

0

0

0

0

53

53

1995-96

Dec

70

0

1

0

2

70

73

1996-97

Dec

50

0

11

3N8

0

53

64

1997-98

Dec

45

1*

0

2

0

0

46

48

Subtotal

482

1*

0

14

3N8

2

486

502

[no. 8 Sandbank

1976-77

Dec

15

0

0

0

0

15

15

1979-80

Dec

34

0

0

0

0

34

34

1981-82

Dec

1

0

0

0

0

1

1

1982-83

Dec

71

0

0

0

0

71

71

1986-87

Nov

102

0

0

0

4

102

106

1987-88

Nov

122

0

0

0

0

122

122

1988-89

Nov

30

0

0

0

0

30

30

1989-90

Dec

10

0

0

0

0

10

10

1990-91

Dec

24

0

2

0

0

24

26

1991-92

Dec

70

0

0

0

1

70

71

1992-93

Dec

60

0

3

0

0

60

63

1995-96

Dec

77

0

1

0

2

77

80

1996-97

Dec

53

0

1

1N7

0

54

55

1997-98

Dec

67

0

3

0

1

67

71

Subtotal

736

0

10

1 N7

8

737

755

Ham Cay

1981-82

Dec

1

0

0

0

0

1

>

Bramble Bay Cay

1989-90

Jan

110

0

0

1 RI

0

111

111

Total

1341

1*

0

24

1 N7

3N8

1 RI

10

1347

1381

1 986. 22 November - 1 6 December: Several courting pairs
of C. mydas were observed daily during the early part of the
study. No mating pairs were seen after 8 December. During
underwater observations on 10 and 14 December, adult
male turtles were seldom seen (5 males out of 263 C.
mydas examined).

1 987. 1 0 July - 1 9 August: Four mounted pairs of C. mydas
observed, two pair each with an attendant male. 28

November: One pair of courting C. mydas seen on No.7
Sandbank Reef. None seen on No.8 Sandbank Reef. 30
November - 5 December: Only a few isolated pairs of
courting C. mydas observed adjacent to Raine Island.
During underwater observations on 1-5 December, adult
male turtles were rarely seen (only 5 males out of 1 144 C.
mydas examined during speedboat transects).

386

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1 0. Adult female Eretmochelys imbricata tagged while nesting at coral cays and continental islands
along the inner shelf of the northern GBR, 1975- 1989. No Chelonia mydas were tagged while nesting on the
inner shelf islands during this period.

Breeding

season

Month

New tags

Within

season

rookery

change

Remigrant

New turtles to
rookery

Total turtles
with tags

Same rookery

Rookery

change

Retag

Farmer Island

1985-86

Dec

2

0

0

0

0

2

2

1987-88

Nov

2

0

0

0

0

2

2

Sir Charles Hardy Islands

1981-82

July

1

0

0

0

0

1

1

Bird Island

1974-75

Mar

3

0

0

0

0

3

3

1976-77

Dec

3

0

0

0

0

3

3

Boydong Island

1976-77

Dec

0

0

o

0

1

1

Milman Island

1976-77

Dec

6

0

0

0

0

6

6

Combined

18

0

0

0

0

0

18

18

1989. 28 November - 07 December: No courting turtles
observed on or near Raine Island Reef.

1991 . 29 November - 10 December: 2 courting pairs near
Raine Island Reef.

1992. 29 November - 10 December: 3 courting pairs near
Raine Island Reef.

1 993. 29 November - 10 December: 3 courting pairs near
Raine Island Reef.

1 994. 30 November - 9 December: 2 courting pairs near
Raine Island Reef; 12 males with courtship damage
captured during reef survey.

1 995. 5- 13 December: No courting turtles observed on or
near Raine Island Reef; 21 males recorded during reef
survey.

1997. 29 November - 6 December: 2 courting pairs
observed near Raine Island Reef; 2 males recorded during
reef survey.

1999. 28 November - 11 December: 1 courting pair
observed near Raine Island Reef; 4 males recorded during
reef survey.

2000. 28 November - 9 December: 1 courting pairs
observed near Raine Island Reef; 2 males recorded during
reef survey.

200 1 . 1 - 1 0 December: No courting turtles observed on or
near Raine Islamnd Reef; 0 males recorded during reef
survey.

Courting C. mydas were rarely observed at any
of the other rookeries of the northern outer barrier
reef after mid November. The only time when
frequent courtship was observed was in late
October - early November. This is consistent with
the pattern of courtship relative to nesting
observed in the southern GBR where C. mydas

aggregates for courtship during late September,
October and early November, after which
courting turtles are rarely seen (Limpus, 1993).

Insufficient courtship was in evidence close to
Raine Island and Moulter Cay to account for the
large nesting population. No fisherman reported
seeing aggregations of hundreds of courting pairs
near these islands, which would be expected if
the turtles courted adjacent to the rookeries.
Reports from fishermen indicated high density
courtship occurred in the shallow protected
waters north of Cape Grenville in October. The
Torres Strait islanders encounter large numbers
of courting C. mydas during September, October
and early November throughout Torres Strait.
This favoured turtle hunting time is called
‘turtlefast’, i.e. the time when the male turtle is
fastened onto the female (Johannes & Macfarlane,
1991). The years with a big turtle-fast correspond
to the high density nesting seasons and poor turtle
fast years are the years in which low density turtle
nesting occurs. There is insufficient C. mydas
nesting within Torres Strait to account for the
courtship that occurs there. Therefore it is
concluded that the majority of the courtship that
occurs within Torres Strait is by turtles that nest at
Raine Island and Moulter Cay. If this is the case
then significant levels of courtship could be
occurring up to 200km from the rookeries. This
spatial separation of the courtship area from the
immediate vicinity of the female’s rookery
characterises much of the C. mydas courtship in
the southern GBR (Limpus, 1993).

GREEN TURTLE POPULATIONS OF RAINE ISLAND

387

TABLE 1 1 . Raine Island. Summary of turtle nesting accounts by month to determine the seasonality of the
Chelonia mydas nesting. Unless otherwise stated the raw data has been obtained from the recent surveys of
nesting density - Tables 2-4.

Month

Year

Day

Observations

July

1843

29

A few turtle tracks (Jukes, 1 847)

1844

Turtles occurred at irregular intervals, generally singularly (MacGillivray, 1914)

1911

9-15

No reference to turtle nesting or to bird predation on hatchlings (MacGillivray, 1914)

1957

20-25

Average 5 turtle tracks per night (Attenborough, in Litt. 4 Apr. 1981).

1982

13-28

0-2 turtles per night.

August

1844

Turtles occurred at irregular intervals generally singularly (MacGillivray, 1914)

1871

31

No reference to turtle nesting (Moseley, 1 879)

September

1844

Beginning in September thev become more numerous (MacGillivray, 1846)

1984

1

7 tracks

October

1910

30

Great numbers of turtles are on the beach and in the shallows around the boat (MacGillivray, 1910)

1975

Early

Approximately 50 females per night

1979

30-31

381 -451 tally count

1986

17-18

123 - 137 tally count

November

1965

Was able to step or jump from turtle to turtle while a companion walked 100 paces beside him (P. Wat-
son, pers. comm. 1979).

1973

1-3

About 1 50 - 200 females per night (track count, Stoddart et al. 1981 )

1974

29

1 1 565 female on shore

1977

early

Approximately 40 - 50 females per night

1979

1-9

410 - 444 tally count

1981

12-30

906 - 2757 tally count

1982

13-30

452 - 1095 tally count

1984

21-30

5 1 79 - 7739 tally count

1985

25-30

1 19 - 308 tally count

1986

13-30

1284 - 4565 tally count

1987

30

3296 tally count

1989

29

'5919 tally count

December

1913

4-12

Reference to obvious frequent clutch destruction by nesting turtles. Turtles abundant in the shallows and
beach edge and the party^tormented at night by the nesting turtles who persist in trying to walk over or
under our sleeping shelter'. While at Moulter Cay ‘thousands of turtles’ came ashore for the night
(MacGillivray. 1917). Moulter Cay described as a ‘turtle- infested sand-bank’ (MacGillivray, 1918b).

1976

2-6

845 - 898 tally count

1977

3-7

19-53 females per night

1978

1-5

148 - 327 females per night

1979

13-14

121 1 - 1482 tally count

1980

8-12

80 - 83 females per night

1981

1-13

1 287 - 3463 tally count

1982

1-7

809 - 1091 tally count

1984

1-23

3916 - 1 1467 tally count II

1985

1-20

142 - 323 tally count

1986

1-15

2910 - 4838 tally count

1987

1-5

2312 -4585 tally count

1988

1-7

516- 1 088 tally count

1989

5-7

5925 - 65 1 6 tally count

January

1845

28

14 large turtles collected (MacKenzie, 1845). —

1953-6?

(May have been February) Estimated 10000-12000 green turtles ashore in one night (V. Vlassoff, pers.
comm. 1981)

1959

Dense turtle nesting sufficient to allow people to ’walk long distances along

tie’ (E. Gray, pers. comm. 1 983). At least 1 100 turtles counted in one walk of the island in one night (N.
Whittaker.jicrs. comm. 11 Aug. 1983).

1974

Late

Several hundred turtle tracks counted by daylight (Ben Cropp, pers. comm. 1975)

1976

29

Approximately 80 tracks (J. Kowarsky, pers. comm. 1977)

February

1959

7-14,22

‘Hundreds were coming ashore after dark’ (Warham. 1963)

March

1975

1-3

408 - 5 1 7 females per night

1981

1-9

388

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1 1 ( Corn .)

1

1 April

1980

8

1 - 2 females per night

1983

25-29

1 - 8 females per night

1984

3-26

2- 1 7 females per night

April-May

1845

28-4

Estimated harvesting rate = 2 turtles per night, calculated from Sweatmans account. (Allen & Corns, 1977)

May

No Records

June

1844

Turtles occurred at irregular intervals generally singularly (MacGillivray, 1846)

1980

9-23

Average < 1 female per night

1981

7- IS

Average < 1 female per night

Only 45 adult male C. mydas have been tagged
at Raine Island and adjacent islands (Table 12).
All of these were breeding males that were
captured in courting groups or had recent mating
scars on their flippers or tails. Adult males have
been recorded only at low density adjacent to
these outer barrier rookeries in the northern GBR.

BASKING. C. mydas at Raine Island were not
observed to haul out onto the beach or onto the
reef flat at low tide to bask during any visit. In
contrast, vertical aerial photographs of Raine
Island from October 1984 show about 1 00 adult-
sized turtles, presumably C. mydas , scattered
along the tide line of the beach. While most were

in the water, tens were above the tide wash. On
very calm days during the nesting season large
numbers of adult turtles could be seen floating
high (with dry carapaces) at the surface for
extended periods at the reef edge and over the
adjacent deep water. This behaviour is defined as
surface basking and is presumed to have a
thermoregulatory function. Aggregations of surface
basking turtles were only observed adjacent to
Raine Island Reef and Moulter Cay Reef.

Beach basking is a prominent behaviour of C.
mydas at the Wellesley Islands of the south-
eastern Gulf of Carpentaria (Limpus et al., 1 994a;
Bustard, 1972) and on the Capricorn-Bunker

TABLE 1 2. Summary of non nesting turtles tagged on Raine Island reef and other reefs of the far northern GBR.
AE = tagged by Applied Ecology P/L. This table does not include the mid year tagging of turtles at Clack Reef
that has been conducted as part of an independent study.

Size Class

Breeding

Month

Adult

Immature

Total

Season

male

female

CCL

>65cm

CCL

<65cm

for trip

Chelonia mydas

\ Raine Island Reef

1977-78

Dec

0

0

1

0

1

1978-79

Dec

12

12 AE

0

0

0

1 AE

25

1979-80

Oct-

Dec

3

0

1

0

4

1982-83

Jul*

1

0

1

1

3

1986-87

Dec

0

0

0

1

1

1987-88

Jul-

Aug

2

0

1

0

3

Nov-

Dec

3

0

5

2

10

1988-89

Dec

0

0

1

0

1

1994-95

Nov-

Dec

12

0

0

0

12

Combined

45

0

10

5

60

Pipon Reef

1974-75

Feb

o

2

0

0

^ J

Milman Island Reef

1978-79

Dec

o

0

4 AE

6 AE

10

! Maclennan Cay

1978-79

Dec

2

0

0

0

2

Size Class

Breeding

Month

Adult

Immature

Total

Season

male

female

CCL

>65cm

CCL

<65cm

for trip

Chelonia mydas

Mac Arthur Cay Reef

1985-86

Jul

o

o

o

1

.

Clack Reef

1986-87

Dec

0

0

2

3

5

1987-88

Dec

0

1

6

4

11

Corbett Reef

1988-89

Dec

25

0

0

0

25

Howick Reef

1988-89

Dec

0

0

0

4

4

Beanley Reef

1988-89

Dec

0

0

0

2

2

Bewick Reef

1988-89

Dec

0

0

0

1

1

Combined

27

3

8

4 AE

15

6 AE

63

Eretmochelys imbricata

Corbett Reef

1987-88

Dec

3

0

0

0

3

Bewick Reef

1988-89

Dec

0

0

0

1

1

Combined

3

0

0

1

4

GREEN TURTLE POPULATIONS OF RAINE ISLAND

389

FIG. 16. Nightly mortality rate of adult female Chelonia mydas
ashore for nesting on Raine Island (Table 3) expressed as a
function of the nightly nesting density.

Group cays in the southern GBR
(Barrett, 1919). Beach basking is also a
common behaviour of the Hawaiian C.
mydas population (Balazs, 1980). This
contrasts with its apparent rarity within
the northern GBR C. mydas population.

This is not just a recent phenomenon;
beach basking does not feature in any of
the historical accounts of C. mydas in the
northern GBR (see above).

MORTALITY OF BREEDING

FEMALES ON THE NESTING
BEACHES

MORTALITY RATE. The earliest
records show that regular mortality of
the female C. mydas while ashore
attempting to nest has long been a
feature of Raine Island (Jukes, 1847; Mac-
Gillivray, 1846; Moseley, 1879; Macgillivray,
1917; V. Vlassoff, pers. comm. 1981, fig. 11;
Warham, 1963). Tables 3-4 show that this adult
female mortality continues in present times.
From these tables, a comparison of daily
mortality and the number of turtles recorded
ashore was made (Fig. 16). Daily adult mortality
was dependent on the number of nesting turtles
ashore on the previous night. Turtles did not
usually die if there were less than 100 females
ashore for the night: 95% of nights with no
subsequent mortality; 5% with 1 death/day (n=43
days). Daily turtle mortality was still low when
the nightly tally counts were in the range of
100-1000 turtles: 71% of nights with no
subsequent mortality; 22% with 1 death/day; 7%
with 2-12 deaths/day (n=69 days). In contrast
when the nightly tally counts exceeded 1,000
there was a substantial increase in daily mortality
and there were few days with no turtle mortality:
7% of nights with no subsequent mortality; 2%
with 1 death/day; 91% of days with 2-75
deaths/day (n=126 days). Nightly tally counts
had to exceed 800 nesting females before the
number of dead exceeded 1/day. There was a
significant correlation between the daily mortality
and the tally count on the preceding night:

daily death rate = 0.0034 1 5*tally count - 1 .376
(r=0.526; d.f.=137. F U37 =152; p<0.0005)

Therefore, through the course of a summer
breeding season it can be expected that daily
mortality rates will increase to a maximum
during the mid nesting season and then decline to
zero as the season draw's to a close. Fig. 17 shows
the cumulative mortality during the early part of
six breeding seasons representing a wide range of

nightly nesting density: 1981/82, 1982/83,
1984/85, 1985/86, 1986/87, 1999/00. With an
assumed peak nesting not being completed until
mid January then the cumulative 1981/82 adult
mortality could have been in the 400-500 range.
A count of carcasses in July 1982 was 189 but
many carcasses known to have been present
earlier in the season had been broken up and/or
buried. In April 1983 the 83 carcasses recorded
from the preceeding breeding season was also an
underestimate because some known carcasses
had been lost in the sand, even though the
season’s nesting density was lower. An
end-of-season count of carcasses is thus likely to
give a substantial underestimate of annual
mortality if the count is not made on a daily basis,
especially in years with high density nesting. The
large counts of turtle carcasses of 281 in
November and 588 in March of the 1974/75
nesting season (Table 3) must be taken as
underestimates of the adult mortality during that
high density nesting season. The annual mortality
of nesting females on Raine Island probably is
expected to be negligible in very low density
nesting seasons like 1977/78 and 1985/86. It
could be approaching 100 in low density nesting
seasons like 1982/83 and greater than 2000 in
very high density nesting seasons like 1984/85,
1996/97 and 1999/00.

In any one season, far fewer turtles die while
ashore nesting at Moulter Cay than at Raine
Island (Fig. 1 7C, Table 4) and negligible numbers
die annually at any one of the other outer barrier
reef cays (Table 5).

Mortality of adult female C. mydas while
ashore attempting to nest at Raine Island is clearly
density dependent. This natural mortality

390

MEMOIRS OF THE QUEENSLAND MUSEUM

NOVEMBER - DECEMBER

FIG 17. Cumulative daily mortality of adult female Chelonia mydas
on Raine Island measured over the early portion of six breeding
seasons. * denotes total dead and dying recorded on arrival of the
study team at the island.

accounts for several thousand breeding
females in high density nesting years
and a few hundred or less in the low
density nesting years.

CAUSES OF MORTALITY. Table 13
summarises the causes of mortality of
the breeding females ashore on Raine
Island. During the First 22yr of the
study, 1 974/75- 1 996/97 seasons, death
from heat exhaustion of turtles that
were still ashore during the first few
hours of daylight was the principal
cause of death (70-89% of adult turtles
dying). Included in these values are
those turtles that died of heat ex-
haustion in the central depression
(2-14% of adult turtles dying). The
majority of the turtles died of heat
exhaustion on the beach platform. The
next most prevalent cause of mortality
was associated with the cliff line of the
central rock platform where 8-20% of
adult turtles died, mostly by falling
over the cliff onto their backs. Large
marine turtles cannot right themselves
when they are turned on their backs.

This regular mortality of adult turtles
that have come ashore at Raine Island
to nest resulted from a variety of
natural causes. Some turtles which
found their way above the cliff during
nesting attempts fell on their backs or
became wedged between rocks on their
return to the beach while others
became trapped in the holes beneath
the rock layer. A small (unquantified)
proportion of these turtles was killed
because their carapaces smashed down
onto their heads and fractured their
skulls when the fell head-first over the
cliff. The remainder, lying on their
backs in the sun with their pale
coloured plastrons upper most,
averaged 3.5 days to die (SD = 1.71, n
= 33, Dec 1984). Those trapped in
caves and other shaded areas
sometimes took weeks to die. The cliff
is a natural feature of the island and was not the
result of quarrying for limestone to build the
beacon or of guano mining as stated by Stannard
(1985). The turtle mortality associated with the
cliff is the result of natural processes.

The number of turtles that died by falling over
the cliff was minor compared to those that died

from heat exhaustion. Turtles that wandered
inland into or beyond the beach depression
encountered problems when endeavouring to
return to the sea. When in the bottom of this
depression, a turtle is exposed to an horizon line
at approximately 0° elevation when looking
parallel to the beach, but one at 1 3° when looking
seaward. Because marine turtles move towards

GREEN TURTLE POPULATIONS OF RAINE ISLAND

391

TABLE 13. Observed causes of adult Chelonia mydas mortality on Raine Island. Percentage of total for
observation period are given beneath the raw values. NR = not recorded. + indicates that only a portion of the
actual dead turtle for the season could be accurately counted. * denotes turtles buried alive by other nesting
turtles. Rescued turtles that would have died w ithout our intervention are shown in brackets with an R prefix and
are included in the adjacent count.

Cause of death

Study period

Heat exhaustion

Fell over

Trapped

Total

Central

depression

beach

cliff

log

Beneath

cliff

In rocks

Under

dinghy

1979: 29 Oct- 13 Dec

7(14%)

33 (67%)

7(14%)

0

2 (4%)

0

0

49

1981: 1 1 Nov-12 Dec

3 (19%)

106

(68.4%)

19(12.3%)

(R19)

6 (3.9%)
(R3)

12(7.7%)

8 (5.2%)

1 (0.6%)

155

Whole season count on 27 July

26

1982: 12 Nov-9 Dec

3(14%)

15(71%)

3 (14%)

0

0

0

0

21

Whole season count on 27 July:

5

55+

20

0

4

0

0

84+

1983-84:

Whole season count on 22 April:

9

33+

43

0

7

0

0

92+

1984: 22 Nov-23 Dec

17 (2.0%)

654

(78.0%)

135

(16.1%)

1 (0.1%)

14(1.7%)

12(1.4%)

5 (0.6%)

838

1985: 24 Nov-20 Dec

0

8 (80%)
(RD

2 (20%)

0

0

0

0

10

1986: 19 Nov-23 Dec

NR

251+2*

(79.6%)

53(16.7%)

12(3.8%)

0

318+

- J

1987: 26 Nov- 10 Dec

5(2.1%)

182 + 1*
(77.9%)

32(13.6%)

( R 1 4)

15 (6.4%)

0

0

0

235

1995: 3-13 Dec

1 (0.4%)

201 (73.4%)
(R3)

62 (23.0%)

(R36)

6 (2.2%)
(Rl)

0

270

1996: 27 Nov- 11 Dec

NR

596(89.5%)

57 (8.6%)

13 (2.0%)

0

666+

1997: 29 Nov- 10 Dec

22 (7.9%)

107(38.6%)

121 (43.7%)

27 (9.7%)

0

277

1998: 28 Nov-1 1 Dec

2 (6%)

16 (44%)
(Rl)

16 (44%)

(R2)

2 (6%)
(Rl)

0

36

1999: 28 Nov- 12 Dec

10(1.8%)

240(43.1%)

257(46.1%)

50 (9.0%)

0

557

2000: 30 Nov- 10 Dec

0

0

2

2

0

4

2001: Nov- 10 Dec

92 (66.2%)

42 (30.2%)

5 (3.6%)

0

139

low elevation, bright areas on their return to the
sea (Limpus, 1971), the profile of the Raine
Island beach platform causes the females in the
beach depression to move approximately parallel
to the water line for varying distances. Some may
wander the entire length of the island in the beach
depression before localised irregularities of
topography lead them to the outer crest of the
beach and hence to the water. The net result is to
increase the time taken to return to the water.
Turtles on the beach after daylight heat in the sun,
leading to locomotory deterioration as
hyperthermia progresses, then ultimately death
ensues. Some of the turtles crossing the cliff line
entered the central depression and some of these
became disoriented and similarly wandered back
and forth across the central depression into the
daylight hours. These too died of heat exhaustion
as the day progressed.

Nesting females remaining on the beach
platform have been observed to die as early as

0930hr and all females that had remained on the
beach after about lOOOhr, carapace uppermost,
were dead by approximately 1700hr. During
December 1989, six females that remained on the
open beach platform towards the middle of the
day were selected for study of lethal hyper-
thermia. Experience had shown that all such
turtles died within the current day. Each was
fitted with a core body temperature probe and the
temperature recorded at approximately
half-hourly intervals for several hours (Fig. 17).
The core body temperature increased from
35.8°C at 1 040hr to as high as 40.4°C in one turtle
at 1 230hr. All turtles died when their core body
temperatures exceeded 39°C. The time of death
varied from 1 230hr to 1 640hr (mode = 1445hr).
These temperatures contrast with the core body
temperatures of nesting turtles on the beach at
night, measured in eggs at the instant of
oviposition, that ranged from 26.0°C in early
November to 28.4-30.0°C in early December

392

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 14. Core body temperature of nesting
Chelonia mydas on Raine Island as indicated by the
temperature of a freshly laid egg.

Date

Temperature (°C)

Freshly laid
egg = core
body temp.

Air

Water

Sand

Night time nesting turtles (2000-0200hr

5 Dec 77

29.8

27.4

27.0

28.0

6 Dec 77

29.6

27.2

28.0

7 Dec 77

29.2

27.2

27.8

28.0

1 Dec 78

28.4

26.2

27.2

30 Nov 79

26.0

27.0

2 Nov 79

26.0

26.5

26.0

6 Nov 79

26.0

26.00

2 Dec 89

30.0

Early afternoon nesting turtle ( 1 500hr)

3 Dec 89

33.0

(Table 14). A female that came ashore and nested
in the heat of the early afternoon had a higher core
body temperature of 33°C (Table 1 4). A turtle on
the beach in the heat of the day has a dark
carapace surface exposed to the sun and this
absorbs solar radiation that can heat the turtle to
lethal temperatures. The carcasses continued to
heat post mortem as the day progressed. By
1500-1900hr the mean core body temperature,
measured during autopsy, of turtles that had died
earlier in the day was 4 1 .4°C (SD = 2.77, range =
32.2-54.4, n = 96). For the 1997/98-2000/01
seasons there was a substantial reduction in the
proportion of turtles dying from heat exhaustion
(0-47%) (Table 1 3). This change is believed to be
a direct consequence of the increased rainfall
associated with these latter seasons (Fig. 7). With
the overcast conditions and the
cooling effect of rain, turtles were less
likely to overheat and die.

Thus, those'areas of the island that
are surrounded by more elevated
habitat can become traps that result in
the turtles not moving directly
seaward but wandering for extended
distances. These disoriented turtles
are at risk of dying from heat
exhaustion if they are still on the
island during the heat of the day.

Similar disorientation and fatal
hyperthermia have been reported for
turtles stranded behind a mainland
mangrove zone by a cyclone tidal
surge (Limpus & Reed, 1985b).

Inverted turtles take longer to attain lethal
levels of hyperthermia due to the reduced heat
absorption of their pale coloured plastrons. Some
turtles that have wandered into partly shaded
areas have survived extended periods ashore
however exposed parts of their bodies have been
extensively sunburned (to the extent that scutes
and underlying tissue peeled from their bodies).
The turtles trapped in the deeper caves beneath
the phosphatic limestone platform are not
subjected to heat stress and probably die from
dehydration.

NECROPSY RESULTS. All dead turtles
examined by necropsy at Raine Island were adult
female C. mydas. All were gravid females with
enlarged mature follicles in their ovaries. All had
recently formed corpora lutea in the ovaries
consistent with having ovulated in the past 2-3
weeks. Some had oviducal eggs, all of which
were of normal ovipositional shell thickness.
Four eggs were removed from each of 7 freshly
dead turtles, preserved in isotonic formalin
within 2 hr of collection and examined for stage
of embryonic development. All were at
embryonic stage 6, (middle gastrulation) which is
the normal ovipositional stage for C. mydas
(Miller, 1985). It must be concluded that only
females ready for oviposition come ashore at
Raine Island.

The enlarged vitellogenic follicles of the
ovaries of 2 freshly dead females were measured
in early December 1988 (Fig. 19). There was a
natural grouping of large yolked follicles that
were approximately the size of the yolks of post
ovipositional eggs. In both females these were
distinct from the remaining follicles which were

FIG. 18. Core body temperatures of moribund Chelonia mydas
females that remained on the beach at Raine Island following nesting
attempts during the previous night. EST denotes Easter Standard
Time.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

393

FIG 19. Size distribution ofyolked ovarian follicles in two Chelonia
mydas females ( A & B) that died of heat exhaustion after remaining
on the beach at Raine Island following nesting attempts during the
previous night.

smaller than normal yolks. This larger size class
of yolked follicle is assumed to contain the
mature follicles from which the successive
clutches were to be produced by each female.
Both of these turtles had complete clutches in
their oviducts (clutch counts: A = 80, B = 100)
from the previous ovulation and sufficient mature
follicles in their ovaries for 4 or perhaps 5
additional clutches (mature follicle counts: A =
449, B = 420). These data suggest that by the
beginning of the nesting season or very early
within it, the female has pre-formed all the
mature follicles she will require for the remainder
of the season. If the number and/or weight of
yolked follicles >25mm in diameter at the
commencement of the migration from the
feeding grounds, during courtship or as she
arrives at the intemesting habitat adjacent to the
rookery could be quantified, it could provide a
measure of the reproductive potential of the turtle
without having to monitor it for months through
an entire nesting season. The size of mature
ovarian follicles and corpora lutea (Table 15)
now forms the basis upon which determination of
reproductive status of females captured in their
feeding grounds are made.

Additional observations of oviducal eggs and
ovaries in the necropsied turtles at Raine Island:

1984, 4-23 Dec: Shelled oviducal eggs were present in
66.2% of the dead turtles (SD = 16.21%, n = 12 daily
samples totalling 310 turtles), regardless of the cause of
death.

1 986, 22 Nov- 1 6 Dec: of 27 turtles necropsied, 2 (7%) had
laid their entire clutch and 25 (93%) were carrying oviducal
eggs.

1988. 29 Nov-8 Dec: All 4 turtles necropsied
still retained what appeared to be complete
clutches in their oviducts (clutch counts = 80,
100, 104, 113).

1 989. 29 Nov-8 Dec: of 99 turtles necropsied,
47 (47%) had laid their entire clutch, 6 (6%)
had laid only a portion of their clutch (8-33
eggs remaining in the oviducts), 46 (46%)
retained what appeared to be complete
clutches in the oviducts (mean clutch = 99.8,
SD = 13.85. range = 66 - 127, n = 45). Rain
during the first night increased nesting
success of turtles for this period, of 8 freshly
dead females examined on 8 December, 5
appeared to have begun atresia of almost all
remaining mature follicles in the ovaries,
even though each had sufficient large follicles
for several more clutches. This prompted
further examination of this issue in
subsequent seasons.

1994, 29 Nov- 10 Dec: of 39 females necropsied, mean
oviducal egg count = 98.9 eggs (SD = 21.76, range =
34- 1 60, n = 39).

1 995, 5- 1 3 Dec: of 22 females necropsied, 6 (27%) had laid
their entire clutch while the remainder still contained
oviducal eggs. At least 6 (27%) of the 22 sets of ovaries
examined contained mature follicles that had begun atresia.

1 997, 29 Nov-6 Dec: of 52 females necropsied. 8 ( 1 5%)
had laid their entire clutch while the remainder still
contained oviducal eggs. At least 38 (73%) of the 52 sets of
ovaries examined contained mature follicles that had
begun atresia.

1998, 1-9 Dec: The single female necropsied had no
oviducal eggs and both ovaries had mature sized follicles
that had begun atresia.

1999, 30 Nov-1 1 Dec: of 66 turtles necropsied, 26 (39%)
had laid their entire clutch. The remainder carried 41-134
oviducal eggs consistent with having laid no eggs or having
been disturbed by other turtles or by unsuitable nesting
habitat after laying only part of a clutch. At least 38 (58%)
of the sets of ovaries examined contained mature follicles
that had begun atresia.

200 1 , l - 1 1 Dec: of 1 6 females necropsied, 9 (56%) had laid
their entire clutch. TTie remainder earned oviducal eggs.
Thirteen (81%) of the 16 sets of ovaries examined
contained mature follicles that had begun atresia. The
remaining 3 ovaries contained less than mature-sized
follicles that had begun atresia.

These data demonstrate that while
considerable variation existed (15-56%), few C.
mydas that died at Raine Island had successfully
laid an entire clutch during the previous night.
The majority died while still carrying oviducal
eggs: some died carrying reduced numbers of
oviducal eggs, consistent with having been
disturbed from a nest before a complete clutch
was laid; and others appeared to be carrying
complete clutches.

394

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 15. Measurements of mature ovarian follicles
and corpora lutea from adult female Chelonia mydas
that died while ashore for nesting on Raine Isiand.
The corpora lutea were selected from the largest size
class, corresponding to the ovulation sites for the
follicles represented in the oviducal eggs present in
the turtles. The mature follicles were selected from
among the largest size class of yolked follicles (Fig.
1 9). Each structure was measured along its greatest
diameter.

Date

Measurement (mm)

Mean

SD

Range

n

Mature follicles

1/12/78

31.1

1.22

29.3-33.5

20

| 17/12/85

33.2

1.04

31.4-34.9

10

Corpora lutea

1 13/12/85

14.7

2.33

10.6-19.6

10

17/12/85

. l4 ->

0.94

12.4-15.6

10

Substantial proportions (27-81%) of the adult
female C. mydas attempting to breed in early
December had already begun resorption of at
least some of their mature ovarian follicles. In as
much as early December is in the first half of the
overall nesting season, these females would clearly
not be converting all mature follicles to eggs.

SIZE OF DEAD TURTLES. The size of nesting
C. mydas females that died on Raine Island is
summarised in Table 16.

RESCUING MORIBUND TURTLES. During
the research at Raine Island, the primary goal has
been to document the functioning of the nesting
population with the minimum of interference to
it. In this regard it was not a goal to rescue dying
turtles. However, in the normal course of the
work, when live turtles were encountered that
could be rescued easily, this was done. For
example in December 1984, over 100 females
were rescued. During the 1989-90 trip, 25
females were rescued. However in some years
few turtles died, as in the 1 985/86 season when
only one was able to be rescued while five others
died in the 27 nights the team was on the island
during 24 November-20 December.

The simplest turtles to rescue were those on
their backs that simply required righting. This
could be accomplished’ by one strong person. If
the cliff margin was patrolled daily at dawn, all
turtles that had fallen on their backs during the
previous night could be released before 0830hr.
Turtles out on the open sand in daylight needed to
be dragged to the beach crest. This normally
required several persons working for more than

TABLE 16. Curved carapace length of adult female
Chelonia mydas that died while ashore for nesting on
Raine Island.

Year

Month

Curved carapace length (cm)

Mean

SD

Range

n

1976-77

Dec

106.7

4.19

99.5-117.5

23

1979-80

Dec

107.5

6.10

96.5-118.0

36

1981-82

Nov-Dec

106.0

5.12

90.0-122.0

84

1987-88

Nov- Dec

104.4

5.77

94.0-114.5

54

1988-89

Nov-Dec

102.5

4.42

95.0-109.0

13

1989-90

Nov-Dec

104.4

5.32

92.5-119.5

99

1991-92

Dead

104.1

4.618

94.4-110.0

48

1992-93

Dead

101.5

98.0-105.0

2

1993-94

Dead

104.1

4.97

89.0-115.8

53

1995-95

Dead

104.7

5.00

90.9-117.0

93

1996-97

Dec

104.1

4.59

89.6-120.7

379

1996-97

Jan-Feb

103.6

4.59

90.8-117.8

415

1996-97

Dead

103.9

4.59

89.6-120.7

800

1997-98

Dead

104.7

5.39

94.0-130.1

103

1998-99

Dead

103.9

4.17

97.8-113.8

12

1999-00

Nov-Dec

105.0

4.89

91.7-119.7

366 ;

2000-01

Nov-Dec

106.0

96.6-112.9

3

2001-02

Nov-Dec

104.2

5.68

81.6-119.4

90

1 5 min per turtle, and was very strenuous work. It
was debilitating to the research team to attempt to
drag turtles across the open sand in the heat of the
day. In high density nesting seasons there were
often hundreds of turtles (sometimes over a
thousand) on the beach as late as 0700hr. Most of
these turtles successfully made it back to the
water. It was usually after 0900hr before the
turtles out on the open sand could be identified as
threatened by heat exhaustion. Even at this time
many ofthe turtles present still successfully made
their way back to the water without assistance.
However, by then it was also stressful for people
to be working over this hot sand, so few turtles
were rescued after 0900hr or before 1 50()hr.

If it were considered a priority to rescue
moribund turtles at Raine Island the following
summary points should be noted: only in high
density nesting seasons will there be numerous
nesting females dying on the island (most ofthe
mortality will occur from mid November to about
mid February); small numbers of moribund turtles
can be rescued by righting those turtles which
have fallen on their backs; most mortality occurs
on the open beach as a result of heat exhaustion,
not of turtles falling on their backs around the
cliff; to rescue a large proportion of the moribund
turtles would require a rescue team’s daily
presence on the island for months at mid season;

GREEN TURTLE POPULATIONS OF RAINE ISLAND

395

3.5 “•
3

RAINE IS.
i MOULTER CAY

2.5 -

H

Z

uJ

U

U

Ed

2 -

.5 -

0.5 "

~ r 1 '■ 1 i 1 1 ^ i 1 1 1 i 1 1 1 i"
0123456789 10

MEAN NIGHTLY TALLY COUNT (Thousands)

FIG. 20. Proportion of nesting female Chelonia mydas at Raine
Island and Moulter with recent shark bites. Data sourced from
Tables 3 & 17.

it is doubtful if most moribund turtles can be
rescued daily in high density nesting seasons
without some mechanical assistance.

SHARK PREDATION. During most summer
visits large tiger sharks (Galeocerdo cuvier) were
seen over the shallows of the reef flat and
adjacent to the edge of the reef every day. Up to
five individuals (judged by size) were seen in the
one day at the western end of the island. Tiger
sharks regularly were seen scavenging on the
carcasses of turtles that had died while ashore at
the beacon end of the island and then washed onto
the reef Hat by high tides. Less frequently they
were observed to prey on live adult turtles. For
example, in December 1986, a school of 5 tiger
sharks attacked and killed an adult female C.
mydas swimming over the reef flat adjacent to the
island. Not all turtles attacked by tiger sharks are
immediately killed. One female (X43893) had
received a bite which removed a portion of her
carapace over the right hind flipper to expose her
abdominal cavity. Her intestine was undamaged
but exposed. When she was seen nesting on 26
November 1982, the wound was healing and
judged to have occurred at about the time of
courtship. While she was nesting she had an open
hole that permitted sand entry into her abdominal
cavity. It did not appear to impede her nesting
drive. The broad snout of the attacking shark
(identified by tooth scars to the carapace)
suggested that she had been attacked by a tiger
shark. Observations such as this emphasise the
impressive capacity of the large marine turtles to
survive extensive injuries through shark attack. It
also illustrates the extent to which the female will
persist with her nesting drive in spite of
debilitation and disturbance.

Each summer a small percentage of
the nesting turtles had been recently
mauled by sharks and the resulting
wounds were still healing when the
turtles were examined while nesting
(Table 17). The degree of healing of
the majority of the wounds indicated
that they had probably occurred at, or
just prior to, the commencement of
nesting, possibly at courtship time.
While some of these injuries had been
caused by tiger sharks, others may
have been caused by the medium to
large whaler sharks ( Carcharhinus
spp.) that occur in the area.

Figure 20 illustrates the occurrence
of fresh shark bites on the nesting
turtles at Raine Island and Moulter
Cay (Table 10) with respect to the size of the
annual nesting population (December mean
nightly tally count for Raine Island has been
adopted as an index of population size; Table 3).
There was a strong negative correlation between
the size of the annual nesting population and the
proportion of turtles recently bitten. This is
further supported by the one small sample of
nesting females examined in mid year 1 987 when
the nesting density was extremely low with less
than 1 turtle/night ashore for nesting: 3 (43%) of
the 7 nesting females examined had been recently
mauled by sharks (Table 17).

The nesting turtles with fresh shark bite
wounds represent the survivors (Table 11) —
those that did not sustain a massive bite or a bite
to a critical area. The actual mortality rate caused
by shark bite of adult female Chelonia mydas at
these rookeries remains unquantified. If the
proportion of turtles surviving bites is positively
correlated with the proportion of turtles in the
population that are killed by shark attack, then
these data suggest that there is a reverse density
dependence between the risk of shark attack and
the size of the annual nesting population. There is
less risk if the turtle breeds in a high density
nesting season. However, the risk ot being
mauled by a shark is highest for turtles that nest
out side the main nesting season and for those that
nest in low density nesting seasons. In addition it
suggests that in the event of a substantial
population decline, the remaining turtles would
be at increased risk of shark attack when
aggregated for breeding, assuming that the shark
population remains stable and independent of the
size of the nesting turtle population.

396

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 17. Frequency of occurrence of recent shark
bites on nesting Chelonia mydas at rookeries in the
outer northern Great Barrier Reef.

Rookery & date

No. of turtles
examined

Turtles with recent shark bites

n | %

Raine Island

Dec 1980

284

10

3.5%

Nov-Dec 1982

1601

13

0.81%

Nov-Dec 1986

11032

22

0.20%

Jul-Aug 1987

7

3

43%

Nov-Dec 1987

2287

7

0.31%

Nov-Dec 1988

2391

32

1.34%

j Nov-Dec 1993

11887

7

0.06%

Dec 1994

1346

6

0.45%

Dec 1995

4615

5

0.11%

Nov-Dec 1997

7122

2

0.03%

Nov-Dec 1998

3284

12

0.37%

' Nov-Dec 1999

25283

10

0.04%

Dec 2000

756

8

1.06%

Nov-Dec 2001

9725

17

0.17%

Moulter Cay

Dec 1986

2581

3

0.12%

Dec 1997

1956

1

0.05%

Dec 1998

427

1

0.23%

Dec 1999

844

0

0%

Dec 2000

84

1

1.19%

Dec 2001

873

0

0%

No. 8 Sandbank

Nov 1988

30

0

0%

Dec 1995

79

4

5%

Dec 1997

183

2

1.09%

No. 7 Sandbank

Dec 1986

74

0

0%

Nov 1988

26

0

0%

Dec 1995

73

0

0%

Dec 1997

108

0

0%

FLUCTUATIONS IN ANNUAL NESTING
POPULATIONS

The most accurate method available at present
to measure the size of a turtle nesting population
requires a whole of season, total tagging census
of the rookery (Limpus et al.. 2001). At Raine
Island this would require a nightly tagging census
from early October to about April. In most years
it would require numerous staff on the island tor
more than 3 months to ensure that every turtle
ashore was accounted for without adding a major
disturbance factor to the turtles. Such an
approach has not been logistically or financially
feasible to date, and would probably cause
significant disturbance to the nesting sea birds.

Tagging at Raine Island has been restricted to
sampling a small portion of the total number of
turtles nesting during any visit.

The C. mydas nesting in eastern Australia is
confined essentially to a summer breeding season
that reaches a peak period of nesting density
during December and January (Raine Island:
Table 4; Bramble Cay: Limpus et al., 2001;
Heron Island: Moorehouse, 1933, Bustard,
1972.). At Raine Island the nesting season is
more protracted than in the southern GBR, but it
still maintains a mid summer peak of nesting
density. It has been demonstrated that the average
number of turtles ashore per night at a C. caretta
rookery within a standard mid season sampling
period can be used as an index of the total annual
nesting population (Limpus, 1985). Turtle
research at Raine Island has been timed to include
a common annual sampling period in late
November to early December. The mean nightly
tally count during the last days of November and
the first two weeks of December (Table 3) will be
used as the index of the size of the annual nesting
population for the purposes of the present study.

C. mydas nesting numbers have fluctuated
widely from year to year at Raine Island and
adjacent outer barrier reef cays (Tables 3-5). The
annual fluctuations in the mean tally count for
Raine Island in the two weeks of late
November-early December is summarised in Fig.
21. In the mid-season of the very high density
nesting years, many thousands of females come
ashore nightly at Raine Island and Moulter Cay
(Fig. 22). As many as 1 1565, 1 1467 and 14519
nesting females have been recorded on Raine
Island in single walks of the beach (1974, 1984
and 1996 respectively) (Table 3, Fig. 23). These
densities were equivalent to 6.4-8. 1 females per
metre of waterline for the island. In contrast, in a
very low density nesting year, as few as 32
females on average have come ashore nightly
(1977) (Table 3). This was equivalent to 1 turtle
ashore per 56.3m of water line. The approximate
synchrony in annual fluctuations in the size of the
C. mydas nesting population encompasses the
entire GBR. This is illustrated by the high level of
synchrony of fluctuations in total annual nesting
population at Heron Island in the southern GBR
and the early December mean tally count at Raine
Island (Fig. 21). These two rookeries have the
most comprehensive census data recorded for C.
mydas in Australia.

The historical records for Raine Island (see
above) contain some useful indications of
changing nesting population levels in past years.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

397

2000

73

O

1500 Z

c/5

H

>

1000 g

rn

O

H

500 S

FIG 2 1 . Annual variation in Chelonia mydas nesting density at
Raine Island and Heron Island. The mean tally count measured in
the first two weeks of December (solid line) is used as an index of
the size ol the total nesting population. These data are derived from
Tabic 3. The nesting season is denoted by the year it commences.
The 1974/75 data were obtained from a single nights visit to the
island on 29 November 1974.

The November 1973 estimate by Stoddart et al.
(1981) of less than 200 females beaching per
night at Raine Island suggests that 1 973/74 was a
poor nesting season. Whittaker estimated the mid
season nesting at ‘as many as 1500 turtles ...
nightly’ (breeding season not specified) when he
applied in 1956 for a permit to harvest turtles
from Raine Island. This estimate is of a similar
magnitude as occurred during the 1979, 1981,
1982 and 1992 breeding seasons. Ellis (1936)
described a scene consistent with up to a few
thousand, rather than ten thousand, nesting
females nightly at Raine Island during the 1 890 to
1892 guano mining operation. In addition, there
are several records consistent with extremely
high turtle nesting density. MacGillivray (1917)
described a scene with immense numbers of
turtles in the shallows during the late afternoon

on 4-11 December 1913 and further he
complained of being ‘tormented at night
by the nesting turtles, who would persist
in trying to walk over or under our
sleeping shelter’. On 12 December 1913
he recorded on nearby Moulter Cay
‘thousands of turtles climbing the
sandbank’ in the moonlight. Vlassoff
(pers. comm.), estimated 10-12 thousand
turtles ashore for the night during the
1955/56 season. While the accuracy of
this estimate cannot be checked, his
measure of about ten new turtles per
night falling on their backs along the cliff
line is consistent with a huge nesting
population as is his photograph showing
high density mortality in the central
depression (Fig. 1 1). Similarly Watson’s
(pers. comm.) account of a breeding
season in about 1965 when he stepped or
jumped from turtle to turtle while
Vlassoff walked 100 paces beside him in late
November is consistent with nesting densities
greater than those recorded in most recent
seasons, except perhaps 1974/75, 1984/85 and
1996-97. In 1987 and 1999. Watson’s method of
measuring turtle nesting density was repeated at
Raine Island and a stepping index recorded for
nights with a range of nesting densities (Table
18). No more than 25 paces from turtle to turtle
could be made when tally counts of up to 7,010
were recorded. From these considerations it
concluded that the 1 974/75, 1 984/85 and 1 996/97
seasons with very high nesting densities were not
unique to recent years for Raine Island. Based on
the historical data above and the recent quantified
data (Tables 3-4) it is concluded that the annual
nesting density of C. mydas at Raine Island has

FIG 22. Early morning oblique aerial photographs of Moulter Cay, 7 January 1990, showing high density nesting
Chelonia mydas.

398

MEMOIRS OF THE QUEENSLAND MUSEUM

been fluctuating widely over several orders of
magnitude for more than a century.

Limpus & Nicholls (1988, 1994, 2000)
demonstrated that the number of C. my das
nesting on the eastern Australian rookeries in a
given year is a function of the El Nino Southern
Oscillation (ENSO) climatic events. The
Southern Oscillation Index (SOI), the
standardised difference in monthly mean
atmospheric pressures between Darwin and
Tahiti, is used as a measure of the variability of
ENSO climate variation (Bureau of Meteorology
Australia, .2002). In testing for the best
correlation between breeding numbers and
ENSO events, Limpus & Nicholls (2000)
identified a significant correlation linked to the
mean SOI value approximately 1 .5yr before the
breeding season commences. This time delay
between the climate event and the onset of
breeding is determined by the duration of the
sequence of physiological processes that
culminate in egg production. A female that
commences laying eggs in November will have
migrated from her distant foraging area
approximately two months earlier. Before she
commenced her breeding migration, she will
have deposited yolk into the hundreds of mature
ovarian follicles that she will use for egg

FIG. 23. Views of Raine Island on the night of 20
December 1 984 when the tally count was 1 1 ,467 turtles.
A, 1 800 hr. There were already hundreds of females
ashore even though it was still daylight. B, 2100 hr.
There were thousands of turtles ashore, mostly
wandering and digging while searching for a suitable
undisturbed nesting site. At times it was difficult to
walk between the nesting turtles. C, 0600 hr. Most of the
nesting females had left the island but there were still
several thousand turtles ashore completing their nesting
or returning to the sea. Many of the returning turtles
were wandering along the beach depression parallel to
the water line tow ards the eastern end of the island, i.e.
towards the low elevation bright horizon.

production. This vitellogenic process
commenced in about February and is completed
by about September. Prior to commencement of
vitellogenesis, the female will have increased her
fat deposits during the previous months. This
present study extends the analysis of Limpus &
Nicholls (2000) using the more extensive eastern
Australian C. mydas census data now available
from 27 breeding seasons (Table 19). There
continues to be a significant linear correlation
between the mean May-October SOI approx-
imately 1.5yr before the breeding season
(MOSOI) and the natural log of the annual green
turtle census data (census) from representative
rookeries at each of the eastern Australian
breeding aggregations (Fig. 24):

Raine Island:

ln(census) = -0.11 78*MOSOI + 6.8949 (F U22 =
19.48, p<0.001 ; r 2 = 0.4696);

Heron Island:

ln(census) = -0.09243* MOSOI + 5.5755 (F 1 25
= 29.89 p<0.001; 1 2 = 0.5445).

It is not unreasonable that a slightly greater
proportion of the variability in the number of
breeding turtles is explained by the correlation
for the Heron Island census. The use of a
mid-season index of nesting numbers (Raine
Island census) should show more variability than

GREEN TURTLE POPULATIONS OF RAINE ISLAND

399

TABLE 18. A comparison of the stepping index and
the nightly tally count measure of Chelonia rnydas
nesting density at Raine Island.

Date

Stepping index (number of paces)

Tally count

Mean

range

n

30/11/87

3.5

3-5

4

3296

1/12/87

4.0

2-7

4

2312

2/12/87

4.2

3-6

5

4498

5/12/87

8.2

6-10

5

5011

28/11/99

4.7

4-5

3

5806

29/1 1/99

7.7

6-8

3

5834

2/12/99

20

15-25

2

7010

6/12/99

15

15

1

6745

a total season tagging census (Heron Island
census).

Limpus Si Nicholls (2000) have demonstrated
that these fluctuations in the size of the annual C.
mydas nesting populations result from variability
in the proportion of adult females present in the
distant foraging areas that prepared for breeding
in any particular year. Because ENSO is a
broad-scale regional climate event for the whole
of the Southeast Asia-Western Pacific region,
there is synchrony in fluctuations in the annual
nesting populations at all C. mydas rookeries in
the region, not only in eastern Australia but also
throughout Southeast Asia (Limpus & Nicholls,
2000; Chaloupka, 2001). Therefore, the
long-term census data for Raine Island and Heron
Island represent significant baseline measures
against which the size of nesting populations at
other C mydas rookeries in the Southeast
Asia-Western Pacific Region can be compared.

The population census data recorded at Raine
Island and Heron Island (Fig. 21) illustrate the
difficulty of quantifying the stability of a C.
mydas nesting population. In the absence of the
1 974 data, the annual nesting populations at both
these rookeries over a 15 yr period could be
interpreted as increasing. However, an inclusion
of the 1 974 data negates that interpretation. There
has been a higher frequency of high density
nesting seasons since 1984 than in the ten
preceding years. The stability of these C. mydas
populations cannot be assessed by measuring the
size of the annual nesting population over a few
years or even over a decade. It will require several
decades of detailed monitoring of the size of the
nesting population before anything more subtle
than catastrophic decline of numbers can be
reliably demonstrated.

REPRODUCTIVE DISCRETENESS OF THE
NORTHERN GREAT BARRIER REEF
CHELONIA MYDAS NESTING
AGGREGATIONS

Traditionally it has been assumed that marine
turtles return to breed at the beach of their birth.
However, there are sparse data on marine turtle
growth from birth to adulthood to demonstrate
the truth of this. There has been one good case
history that provides circumstantial evidence that
the major C. mydas rookery areas support
non-interbreeding populations. The once large C.
mydas population breeding at the Grand Cayman
rookery of the Caribbean Sea was overharvested
to approximate extinction during the 19th
century and almost no C. mydas nesting has been
recorded there in the 20th Century. However, the
large Tortuguero C. mydas rookery of Caribbean
Costa Rica (approximately 1,000 miles away
from Grand Cayman) still functions at high
nesting density (Groombridge & Luxmoore,
1989). Turtles bom at the Tortuguero rookery
have not been recruiting to breed on the Grand
Cayman beaches.

Within Australia, Bustard (1972) postulated
that the nesting populations of the separate cays
of the Capricorn-Bunker Groups, southern GBR,
were non-interbreeding groups. The manage-
ment implications of this were that each island
breeding unit could be managed independently of
the others. However, movements of breeding
turtles demonstrated by tagging-recapture
studies of courting and nesting turtles (Limpus et
al., 1984a, 1994b; Limpus, 1993) indicate that C.
mydas cannot maintain non-interbreeding
populations among a group of islands as closely
spaced as those of the Capricorn-Bunker Groups.
In addition, as a result of temperature dependent
sex determination among marine turtles and the
differences in beach temperatures, different
sexes of hatchlings can be produced from
different rookeries (Limpus et al., 1983c). This
latter study suggested that the warmer rookeries
of mainland south Queensland produce mostly
female hatchling Caretta caretta while the cooler
beaches of the adjacent coral cays of the
Capricorn-Bunker Groups produced mostly male
hatchlings. The entire assemblage of these
rookeries probably represented a management
unit comprising complementary rookeries for
Caretta caretta.

PROTEIN ELECTROPHORESIS. Using an
electrophoretic survey of presumptive protein
loci, Gyuris & Limpus (1988) demonstrated that

400

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 19. Annual census data for nesting green turtles, Chelonia my das, at Heron Island representing the
southern Great Barrier Reef nesting population and Raine Island representing the northern Great Barrier Reef
nesting population. MOSOI denotes the mean May to October SOI (Bureau of Meteorology Australia, 2002)
from approximately 1 .5yr before the breeding season; In denotes the natural logarithm. The Heron Island census
data are derived from a tagging census of the total annual nesting population. The Raine Island census data are
derived from the mean nightly tally count (number of nesting females counted in one walk of the island) for the
first two weeks of December. These data are an extension of the data summarised in Limpus & Nicholls (2000).

SOI data

Turtle Data

Year

MOSOI

Breeding season

Heron Island

Raine Island

census

In(census)

census

In(census)

1972

-13.6

1974

1121

7.02

-

-

1973

9.5

1975

21

3.04

-

-

1974

8.8

1976

355

5.87

872

6.77

1975

17.3

1977

46

3.83

32

3.47

1976

-5.4

1978

285

5.65

243

5.49

1977

-13.0

1979

513

6.24

1347

7.21

1978

4.0

1980

327

5.79

59

4.08

1979

-0.8

1981

878

6.78

2049

7.63

1980

-2.6

1982

139

4.93

925

6.83

1981

6.1

1983

125

4.83

-

-

1982

-18.8

1984

1471

7.29

6493

8.78

1983

1.6

1985

42

3.74

227

5.42

1984

-1.2

1986

505

6.22

3956

8.28

1985

-1.0

1987

947

6.85

4102

8.32

1986

-0.1

1988

111

4.71

731

6.59

1987

-15.2

1989

1009

6.92

6144

8.72

1988

11.2

1990

120

4.79

-

-

1989

6.4

1991

602

6.40

3828

8.25

1990

1.5

1992

498

6.21

1417

7.26

1991

-10.6

1993

488

6.19

8462

9.04

1992

-5.7

1994

370

5.91

500

6.21

1993

-11.7

1995

632

6.45

4647

8.44

1994

-15.0

1996

1509

7.32

12100

9.40

1995

-0.6

1997

289

5.67

4067

8.31

1996

6.28

1998

351

5.86

471

6.15

1997

-18.1

1999

1801

7.50

6693

8.81

1998

9.47

2000

26

3.26

129

4.86

1999

2.98

2001

3402

8.13

if reproductive isolation occurred among the
Caretta caretta rookeries of the southern Great
Barrier Reef then the isolation was between the
rookeries of the Swain Reefs and those of the
remainder of south Queensland. A similar study
of eastern Australian C. my das was attempted.
Non-sibling C. mydas hatchlings from Raine
Island and other eastern Australian rookeries
were collected during the 1979/80 breeding
season for a collaborative study of the population
genetics of eastern Australian C. mydas with
Commonwealth Scientific and Industrial
Research Organisation staff (Cleveland
Laboratory). The specimens were inadvertently
destroyed at the laboratory before the study was

completed and no results are available. During
the 1982/83 breeding season 21 non-sibling C.
mydas hatchlings were collected at Raine Island.
Another sample of >50 non-sibling hatchlings
collected from Raine Island in April 1984 was
lost before analysis as the result of a freezer
power failure. An electrophoretic survey of 27
presumptive protein loci was made of the
available C. mydas hatchlings from Raine Island
and a similar series collected from Heron Island
(Gyuris, 1 984). This study identified low genetic
variability within the species but the results were
insufficiently differing to reject the nul
hypothesis of there being a single breeding
population of C. mydas within the GBR. The low

GREEN TURTLE POPULATIONS OF RAINE ISLAND

401

73

2

Z

H

>

Cl

£

S

o

n

m

2

SOI (MAY-OCTOBER) 1.5 YEARS BEFORE BREEDING SEASON

FIG 24. Correlation of the natural logarithm of the annual census data
recorded for northern and southern Great Barrier Reef Chelonia
mydas populations and the mean May-Octobcr Southern Oscillation
Index (SOI) approximately 1.5 years before the breeding season.
Census data (Table 19): for Rainc Island, annual mean tally count
during the first two weeks of December; for Heron Island, annual total
tagged nesting population. See text for regression equations and
goodness of fit statistics.

levels of electrophoretic variability in sea turtles
is such that it is not sensitive enough to deliniate
reproductively isolated populations.

DNA ANALYSES. The application of mito-
chondrial DNA restriction analysis to Australian
C. mydas nesting populations at Raine Island,
Heron Island and Western Australia (Norman et
al., 1 994a) reinforced the studies of Bowen et al.
(1989, 1992) and Meylan et al. (1990) that
demonstrated that the nesting female returns to
breed in the region of her birth. Norman et al.
(1994a) further demonstrated that, while widely
separated female breeding aggregations were
genetically different, the nesting populations on
nearby islands could not be discriminated. These
results were subsequently corroborated by
mtDNA sequencing studies and microsatellite
nuclear DNA analysis by Norman et al. (1994b),
FitzSimmons et al. (1995, 1997a) and Moritz et
al. (2002). In particular, these studies have
identified that there are three separate genetic
stocks that breed in eastern Australia in the
northern (nGBR), the southern GBR (sGBR) and
the Coral Sea Platform (Fig. 13). Each of these
stocks is also genetically separate from each of
the other breeding aggregations tested from
within the Indian Ocean and Pacific Ocean
basins. Torres Strait is identified as a stock
boundary with 100% genetic isolation occurring
between C. mydas breeding at GBR rookeries
and those of the Gulf of Carpentaria. The nGBR
stock encompasses rookeries from Bramble Cay
in northeast Torres Strait south to at least No. 8

Sandbank which lies north of Princess
Charlotte Bay, w r ith the largest nesting
aggregations centred at Raine Island
and Moulter Cay. Each stock is char-
acterised by a dominant haplotype
and all show very low levels of within
population variation. The dominant
haplotypes of the nGBR and sGBR
stocks are shared at low- frequency by
the other respective stock. This over-
lap in the distribution of the dominant
nGBR and sGBR haplotypes indicates
that a very low level of interbreeding
by females could be occurring
between the two stocks. In contrast
there is no detectable female mediated
gene flow between the east Australian
and Gulf of Carpentaria-Western
Australian stocks.

FitzSimmons et al. (1997b)
compared the mtDNA haplotypes of
breeding male C. mydas at three
courtship areas (Raine Island Reef, nGBR; Heron
Island Reef, sGBR; Bountiful Island, Gulf of
Carpentaria) with mtDNA haplotypes from
nesting females at the rookeries adjacent to each
of these courtship areas. They found that
breeding males were displaying comparable
philopatry to the courtship area as the nesting
females were to their nesting beaches. A
comparison of rates of nuclear DNA gene flow
(male and female mediated gene flow) and
mtDNA gene flow (female mediated) among C.
mydas rookeries located around the Australian
coast (FitzSimmons et al., 1997a) identified that
the estimated rates of gene flow between stocks
were generally higher than expected for nuclear
DNA "relative to mtDNA. The difference was
most pronounced between turtles from the nGBR
and sGBR stocks. It was proposed that the
atypical nuclear DNA difference between the
nGBR and sGBR stocks was most likely the
result of the geography of Torres Strait that
caused a bottleneck for turtles migrating from the
west. Some breeding females from the sGBR
stock migrate from foraging areas in Arnhem
Land and Gulf of Caipentaria through Torres
Strait towards their traditional nesting area in the
southern GBR. These sGBR females could be
mated by males from the nGBR stock in the
Torres Strait courtship area as the sGBR females
pass through. Nuclear DNA gene flow could thus
occur between the stocks even though both the
males and females display comparable high

402

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 20. The geographical distribution of foraging areas identified via tag recoveries of adult female Chelonia
mydas recorded nesting at northern Great Barrier Reef rookeries. * Tag recoveries from Torres Strait could also
include migrant turtles captured at courtship or turtles captured while migrating through the area.

Foraging area
(Recapture location)

■ 1

Rookery area

Northern GBR Stock

Southern GBR

Eastern Torres
Strait

Raine Is +
Moulter Cay

No.7 4
Sandbanks

Inner shelf, nth
GBR

Total

Indonesia

2

17

19

0

i Arnhem Land & Gulf of Carpentaria

2

40

4

46

11

1 Torres Strait* (PNG+Qld)

17

215

6

1

239

15

i Eastern & northern PNG

3

14

2

1

20

3

Vanuatu

1

1

0

Fiji

0

2

New Caledonia

2

2

31

GBR: Cape York to 14°S

3

60

1

64

24

GBR: 14-24°S

1

7

1

9

298

Qld >24°S

3

3

109

New South Wales

0

.

Total

28

359

14

2

403

- 494 |

levels of philopatry to their respective breeding
areas (FitzSimmons et al., 1997b).

In summary, the assemblage of C. mydas that
breeds at the rookeries and courtship areas of the
northern GBR centred on Raine Island and
Moulter Cay and the dispersed assemblage of
these same turtles in their respective foraging
areas represents a globally significant, single,
independent management unit.

MIGRATION

Limpus et al. (1992) reported on breeding
migrations from the eastern Australian C mydas
stocks. The sample size of migration recaptures
of adult females from the northern GBR
rookeries (n=403) has approximately doubled
since that study. The current dataset is
summarised in Table 20 & Fig. 25. Conclusions
from this larger dataset are summarised.

Adult females from the nGBR stock migrate
from foraging areas in eastern Indonesia, Papua
New Guinea, Vanuatu, New Caledonia, Northern
Territory and from throughout coastal
Queensland to breed at rookeries within the
northern GBR and Torres Strait. These foraging
areas span a region 25° in latitude from southern
Irian Jaya in Indonesia to Moreton Bay in south
Queensland and 37° in longitude from Melville
Island in western Northern Territory to
southeastern New Caledonia. While the majority
of recaptures have occurred within a few hundred
kilometres of the rookery, the longest distance

between rookery and capture site was 2,773km
from Raine Island to Vanuatu.

The majority of the nGBR stock tag recoveries
have been from north and west of the nesting
beaches, from foraging areas in Torres Strait,
Gulf of Carpentaria and coastal areas of the
Arafura Sea in Irian Jaya and Northern Territory.
The north-south distribution of foraging area tag
recoveries from eastern Australia of adult
females from each of the nGBR and sGBR stocks
are summarised in Fig. 26. While there has not
been equal sampling effort at each latitude, the
nGBR stock is the dominant component of the
foraging populations within eastern Australia to
as far south as approximately latitude 13°S.
South of this, the sGBR stock is dominant. These
data indicate that adult females from stocks
breeding at opposite ends of the GBR do not
occupy identical feeding distributions.

Migration data are available for only one adult
male. This resident from Shoalwater Bay
(~22°20’S, 150°12’E) in Central Queensland was
trawl captured in Torres Strait at courtship time.

Most (76.7%, n=3 1 0) of the reported nGBR C.
mydas stock migration recaptures were of turtles
hunted for food, 3.2% (n=13) were from
incidental captures in commercial fisheries,
mostly trawling, 15.8% (n=64) were captured
during systematic sampling of feeding
populations during Queensland Turtle Research
projects and 2.2% (n=9) were from stranded dead
or rescued turtles. The remaining 2.0% (n=8) of
reports did not identify the capture method.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

403

FIG 25. Distribution of non-nesting recaptures of adult female Chelonia mydas that had been tagged at rookeries of
the northern Great Barrier Reef stock: No.7 and No.8 Sandbanks, Raine Island, Moulter Cay and Bramble Cay.

In addition to these tag recoveries there have
been other reports of tagged turtles for which the
tags and/or the data were not returned. This is
particularly so in neighbouring countries where
there may be low levels of literacy or a poor
capacity to read the English text of the tags. In
remote areas, hunters may not be able to afford
the postage to return tags. In some parts of
northern Australia groups of tags have been
found in homes of hunters, the tags having been
retained as souvenirs of the hunting. In contrast,
there is a belief among many of the turtle hunting
communities of eastern Indonesia, PNG and
some parts of northern Australia that the tag
represents ownership of the turtle and that the
hunter will ‘get into trouble’ for killing someone
else's turtle if he reports the capturing of a tagged
turtle. In the same area there also has been
negative publicity concerning turtle hunting and
tags are associated with bureaucratic interference
in the hunting practices. In these latter cases the
tags are apparently often discarded. These
reasons may account for the recent paucity of tag
returns from Indonesia even though there was a
substantial increase in C. mydas hunting for the

Bali market in the Aru and Irian Jaya areas since
about 1988 (J. Schulz and I. Suwelo, pers.
comm., Nov. 1990) until the 2001 closure of the
Bali turtle trade (W. Adnyana, pers. comm.. May
2002). This shift in hunting effort probably was in
response to the substantial decreases in C. mydas
populations in other parts of Indonesia
(Groombridge & Luxmoore, 1989).

There are some general principles for C. mydas
breeding migrations that can be drawn from these
data.

1) The individual adult female has a strong site
fidelity to her traditional nesting beach (Table 7).
However, the present study has not demonstrated
clearly the corresponding fidelity of the female to
her particular feeding area as has been
demonstrated for the females migrating to breed
at the southern GBR C. mydas rookeries (Limpus
et al., 1992). With most post-nesting migration
recaptures from the northern GBR rookeries
having been derived from turtles hunted for food,
there was a reduced potential for recording turtles
over successive legs of their migration between
foraging and nesting areas to establish foraging
area fidelity.

404

MEMOIRS OF THE QUEENSLAND MUSEUM

FREQUENCY

FIG. 26. Comparison of frequency distributions of
captures in eastern Australian foraging areas south
from the Papua New Guinea coast of Torres Strait of
adult female Chelonia mydas recorded on nesting
beaches of the northern and southern GBR stocks.
Recaptures grouped by 1° latitudinal blocks.

2) Adult females that breed at the same rookery
do not all migrate from the same foraging area
(Table 20).

3) Breeding females did not necessarily nest at
the closest rookery to their respective foraging
areas. For example, X 1 0925 was tagged at Raine
Island, stranded 1 ,057km away by cyclone Kathy
in the MacArthur River delta 3 yr later was
rescued, then' again recorded nesting at Raine
Island after a 4yr remigration interval (Limpus &
Reed, 1985b). She did not go to the Wellesley
Group rookeries of the southern Gull of
Carpentaria or the eastern Arnhem Land
rookeries that were closer to her feeding area.
This also applies for all recaptured turtles from
the Northern Territory. Similarly, not all the
females that lived on the reefs of the tar northern
GBR migrated to the adjacent Raine Island and
Moulter Cay rookeries. Some migrated past
Raine Island to breed in the southern GBR.

4) Females living in the same foraging area do not
all nest at the same rookery (Table 20).

5) Turtles in foraging areas represent populations
of mixed genetic stocks. For example, in addition
to those that breed at nGBR rookeries, female C.
mydas foraging in the Princess Charlotte Bay
area in the northern GBR have been recorded
breeding at rookeries in the Coral Sea Platform,
the southern GBR and internationally in eastern
Papua New Guinea, eastern Solomon Islands,
and northern New Caledonia. In an extreme case,
adult female C. mydas captured off Coburg in the
Northern Territory, have been recorded nesting at
Raine Island, at Western Australian rookeries and
in southeastern Java, Indonesia.

REMIGRATION

Remigration, the interval between breeding
seasons, varies between species and has posed a
problem for measurement because of the long
remigration intervals. The main problems have
resulted from tag loss and the logistics of
systematically monitoring a rookery for many
years. Early research on marine turtle
reproductive ecology did not rigorously address
tag loss. In 1974 the Queensland Turtle Research
Project was requested to take over the C mydas
field studies at Heron Island that had lapsed
following the departure of Dr H.R. Bustard from
Australia. The methodologies of the original
project were initially retained, in particular the
tagging methodology of applying a monel tag to
the LI tagging position (Bustard, 1966). Within a
few years it was apparent that there was a
significant tag loss problem within this study. In
1978 systematic studies were commenced to
identify the optimal position for tag application
and to find a tag that had low rates of loss over a
10 yr development. These studies identified the
need for a new design of turtle tag that used a
metal with very low corrosion rates, and that was
at a reasonable cost. Titanium and inconel 625
were identified as the preferred metal types.
Within the Queensland Turtle Research Project,
we were able to locate a tag manufacturer
prepared to produce turtle tags from titanium. In a
parallel study in Hawaii, G Balazs worked with
an American manufacturer to produce a turtle tag
from inconel 625. The tag trial studies (Limpus,
1992a) demonstrated that the optimal tagging
position is in the axillary area on the trailing edge
of the front flipper (L3 or R3 tagging positions).
The more distal ly the tag is applied, the more
likely it is to be lost. Monel tags have a high
probability of tag loss, approaching 100% in 10
yr. Plastic tags did not perform better than the
monel tags in the long term. Titanium tags were

GREEN TURTLE POPULATIONS OF RAINE ISLAND

405

TABLE 21. hrcquency distribution of remigration recaptures of nesting female Chelonia mydas recorded by
tagging year cohorts at Raine Island since the commencement of Raine Island Corporation funded research and
monitoring. The summer breeding season is defined by the year at the commencement of the breeding season.
Tag type definitions follow the tag design definitions of Limpus ( 1 992a): M, monel tags; T, titanium tags. Slight
variations in design of the titanium tags are denoted by T1 , T2, T3 and T4. However, the variation in the design
of the titanium tag is not included in the analysis of this study.

Census for recovery of tags

Breeding season for commencement of cohorts on

Year

1981

1982

1982

1983

1984

1985

1986

1987

1988

1989

1990

1991

1992

1993

1994

1995

1996

1997

1998

1999

2000

2001

No tagged turtles

2605

1094

504

80

2545

1919

1717

1559

1952

1037

1208

1368

1999

1593

1034

1668

2997

1434

1982

2210

840

1417

M2

M2

T1

T2

T2

T2

T2

F2

T2

T2

T2

T2

T2

T3

T3

T4

T4

T4

T4

T4

T4

T4

1981

1982

0

1983

No RIC survey

22

l

0

1984

16

8

1

0

1985

43

22

5

1

0

1986

9

5

14

2

2

0

1987

1

2

17

2

9

0

1

1988

0

0

l

1

34

23

1

0

1989

40

0

2

0

138

92

29

0

0

1990

No RIC survey

2

7

0

0

27

25

51

40

3

0

1991

2

0

0

2

37

II

29

39

86

0

0

1992

0

1

2

0

12

13

5

27

72

2

0

0

1993

0

3

3

0

38

12

12

40

53

22

6

0

0

1994

No RIC survey

0

0

0

0

8

1

l

7

3

1

6

2

0

0

1995

I

0

1

0

18

2

6

12

23

16

31

23

4

0

0

1996

2 (Dec-* Feb)

surveys

0

0

2

1

42

13

18

33

42

42

40

56

74

23

0

0

1997

0

0

s

0

33

13

9

26

45

22

44

39

84

57

13

0

0

1998

0

0

1

0

14

2

2

7

24

6

17

9

31

28

13

3

0

0

1999

0

0

0

0

57

7

21

46

56

43

61

67

114

176

74

115

32

1

0

2000

0

0

0

0

2

5

0

2

7

2

2

5

1 1

5

12

3

11

6

0

0

2001

0

0

3

3

23

3

8

17

17

16

18

25

41

31

21

59

108

53

12

0

0

Rem i gran 1 ;
Cumulative total

136

49

57

12

494

222

193

296

431

172

225

226

359

320

133

180

151

60

12

0

0

found to have at least a 50% lower rate of tag loss
within the 6 yr period that titanium tags had been
under trial. The experimental studies on tag
retention were conducted in the southern GBR
where it was logistical ly easier and less
expensive to make repetitive captures of the same
turtles over many years. As improvements in
tagging methodology have been developed they
have been applied to the nGBR studies.

REMIGRATION TAG RECOVERIES. The
number of remigration recaptures during each
study trip are summarised in Tables 7-9. There
have been 3,727 remigrant recaptures of turtles
still wearing a tag up until the end of the
2001-2002 breeding season at Raine Island and
adjacent rookeries. These are not all of the tagged
remigrants that have returned to these islands as
there has been no examination of the entire
nesting population in any one season, nor has the
entire nesting population been tagged in any one
season. The vast majority (98%, n=3,659) of
remigrant recaptures were recorded at the same
rookery at which each turtle was originally
tagged (Tables 7-9). Of the 33,234 females
originally tagged at Raine Island prior to 2001,
one has been recaptured nesting at Bramble Cay
and 21 at Moulter Cay. This very low rate of
interseason change of rookery will be partly the

result of the low sampling rate at nGBR rookeries
other than Raine Island. However, of the 2,992
females tagged on the other outer cays of the
northern Great Barrier Reef prior to 2001, only
1.2% (n=37) have been recaptured nesting at
Raine Island in a later season: 16 from Moulter
Cay, 10 from No. 8 Sandbank, 7 from No. 7
Sandbank and 4 from Milman Island. In addition,
one from No. 7 Sandbank and two from Milman
Island changed rookeries between breeding
seasons to nest at Moulter Cay; three from No.8
Sandbank changed to No.7 Sandbank and one
from No.7 Sandbank changed to No.8 Sandbank.

There has been considerable tagging of C.
mydas at other Australian rookeries in the
southern GBR, Coral Sea region and the
Wellesley Group in southeastern Gulf of
Carpentaria since 1974. Only one of the
approximately 40,000 female C. mydas , tagged
while nesting in the southern GBR prior to the
2001/2002 season, was recaptured nesting at any
northern GBR rookery (Tables 7-9), although
there have been many thousands of remigration
recaptures of southern GBR nesting females
returning to nest in the southern GBR rookeries in
later seasons (C. Limpus, unpubl. data).
Similarly, only one nesting female from the
QPWS and Environment Australia Studies at the
Coral Sea Platform rookeries has been recaptured

406

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 27. Cumulative recapture rate of remigrant adult female Chelonia mydas at northern GBR rookeries
recorded by the yearly tagging cohorts at Raine Island. Data are presented for the yearly tagging cohorts for
which titanium tags were applied from 1984 until 1996.

breeding at Raine Island. None of the females
tagged while nesting at the northern GBR
rookeries have been recaptured nesting at the
more intensively studied southern GBR
rookeries. None of the more than 1 ,000 nesting C.
mydas tagged nesting at Bramble Cay by Applied
Ecology Pty Ltd in 1974 to 1980 has been
recaptured nesting at Raine Island. No female C.
mydas tagged nesting at an eastern Australian
rookery has been recaptured nesting at a rookery
outside of eastern Australia or vice versa.

Collectively, these data demonstrate that an
adult female C. mydas exhibits a high level of
fidelity to a particular nesting beach and can
return to that same nesting beach even after
migrating large distances to home feeding areas
in the intervening years (Limpus et al., 1992).
These observed rates of movements of breeding
females among rookeries are reinforced by the
results of DNA genetic analyses of C. mydas
breeding populations of the Indo-Pacific Region
(Moritz et al., 2002) that demonstrate a genetic
interchange among adjacent rookeries such as
those of the northern GBR and Torres Strait. At
the same time, there is very little genetic
interchange between the rookery regions of the
north and south of the GBR (Norman et al.,
1994a) or between rookeries of the nGBR stock
and those much further away (Norman et al.,
1994a; Moritz et al., 2002). The remigration tag

recovery data are consistent with the genetic data
in identifying groupings of C. mydas rookeries
that support an interbreeding population
separated from more distant breeding
aggregations with which they do not interbreed.

RECAPTURE RATE OF REMIGRANTS. The
remigration recaptures at Raine Island, un-
corrected for tag loss, are summarised according
to the breeding season in which they were tagged
in Table 21 along with a summary of the tagging
methodology used with each tagging year class.
Less than 1% recapture rates resulted from
tagging with monel tags applied in the LI or R1
positions. The change to tagging in the L3
position improved the recovery of monel tagged
turtles by a factor of approximately 5 to 1 0 times.
The change to L3 applied titanium tags has more
than doubled the rate of recovery over that of L3
applied monel tags (Table 22). Currently, the
sampling of the Raine Island nesting population
for approximately 2 weeks in early December
each year is yielding a recovery of titanium
tagged remigrants equivalent to 12-22% of the
turtles originally tagged in the cohort (Fig. 27).
These recovery rates continue to improve as
additional sampling continues. In 2001 there
were still recoveries being made from all year
classes of titanium tagged turtles including 23
remigrants from the 1984/1985 season (17 yr
since first tagged) (Table 21). There were even

GREEN TURTLE POPULATIONS OF RAINE ISLAND

407

TABLE 22. Frequency distribution of remigration
from various year classes pooled by tag design. Ml
and M2 denote different designs of monel tags
(National Band and Tag Co. #49 and #19
respectively); T1 and T2 denote titanium tag designs
as described by Limpus (1992a).

Breeding season

1979/1980 +
1980/1981

1981/1982 +
1982/1983

1982/1983 +
1983/1984

No. tagged in cohort

2053

3696

584

Tag design

Ml

M2

T1 &T2

Tag position

L3

L3

L3

Remigration inter-
val

Frequency

Frequency

Frequency

1 yr

0

0

0

2yr

0

1

2

3yr

4

30

7

4 yr

9

38

16

5 yr

23

48

18

6 yr

2

10

3

7 yr

5

3

3

2

3

9 yr

1

Total

46

133

49

Proportion

recaptured

2.2%

3.6%

8.4%

three turtles recorded for the first time since they
were tagged in 1982/1983 when titanium tags
were first deployed. These extremely long
intervals between captures probably represent
multiple remigration intervals, given incomplete
annual censusing. The tag recovery rate would
improve if the sampling period at the island each
summer was increased in duration.

In the December 1986 sample there were 2.2
times as many tag-scarred remigrants as there
were remigrants with tags. During the December
1988 sampling period (6 yr alter introduction of
titanium tags) when many hundreds of nesting
females were examined for tags each night,
tagged remigrants were recorded at the rate of
2.0% of the nesting females examined (SD =
0.888, range = 0-2.55%, n = 9 nights).
Remigrants that had lost their tags were
recognised at an equal rate (mean = 2.0%, SD =
1.49, range = 0-3.95%, n = 9 nights). Most of
these tag-scarred turtles are probably from the
monel tagging years. By the December 2001
sampling (19 yr after introduction of titanium
tags), tagged remigrants were recorded at the rate
of 5.07% of the nesting females examined (SD =
1.996, range = 3.012-10.030%, n - 10 nights).
Remigrants with lost tags were recorded at a rate
that was an order of magnitude lower (mean =

TABLE 23. Regression equations (in the form y = ax +
b) for transformation from curved carapace length
(CCL. cm) to other carapace measurements (cm) for
breeding Chelonia mydas at Raine Island. * = not
significant.

X

Y 1 a

b | r 2

n

P

Female

CCL

Curved

carapace width

0.776

14.469

0.769

504

<0.001

CCL

Straight
carapace
length (SCL)

0.983

3.356

0.997

54

<0.001

CCL

Carr's straight
carapace
length (CSCL)

0.787

16.243

0.963

29

<0.001

SCL

Straight
carapace width
(SCW)

0.494

26.733

0.575

54

0.02>p

>0.01

Male

CCL

Curved

carapace width

0.735

16.272

0.703

25

0.0 l>p
>0.005

CCL

Straight

carapace

length

0.771

17.613

0.967

25

<0.001

CCL

Carr’s straight
carapace
length

0.799

15.516

0.937

14

<0.001

SCL

Straight
carapace width

0.540

20.681

0.569

24

0.2>p

>0.1*

0.79%, SD = 0.419, range = 0.149-1.399%, n =
10 nights). Large numbers of nesting turtles at
Raine Island need to be examined on a nightly
basis to ensure good sample sizes for remigration
tag recoveries.

REMIGRATION INTERVAL. The frequency
distribution of recaptures, uncorrected for tag loss,
of remigrant C. mydas at Raine Island by tagging
cohort and year of recapture are summarised for all
years since the commencement ot the Raine Island
Corporation funded research in 1981 (Table 21).
The usual method for presenting remigration data
summarises the remigration interval recorded at a
rookery within the year of recapture (Limpus et al.,
1994b). While this method has value when
considering events associated with the immediate
breeding season, it confounds nesting season and
tagging cohort effects. To enhance understanding
of population dynamics, an analysis ol
remigration interval by tagging cohorts is more
appropriate. Because the consecutive annual
tagging cohorts have differing durations over
which remigrants are recorded, an analysis of the
raw data from each cohort (Table 21) would
introduce a bias from the longer studied cohorts.
The following considerations have been taken into
account in choosing an interval for comparison

408

MEMOIRS OF THE QUEENSLAND MUSEUM

5 6 7 it 9 III II 12 I J 14 15

REMIGRATION INTERVAL (yr)

REMIGRATION INTERVAL (yr)

REMIGRATION INTERVAL (yr)

1988 TAGGING COHORT (n-1952)

REMIGRATION RECAPTURES-42I

5 6 7 8 9 10 II 12 IJ 14 15 I*

REMIGRATION INTERVAL (yr)

* 12

K 7

1989 TAGGING COHORT (I.-I0J7)

REMIGRATION RECAPTURES- 1 72

► 7 a 9 IU II 12 IJ 14 15 16 17

REMICRATION INTLRV'AL (yr)

FIG 28. Frequency distribution for remigration interval recorded from yearly cohorts of adult female Chelonia
mydas tagged nesting at Raine Island since the introduction of large scale use of titanium tags, 1 984- 1 997 (Table
21). * denotes an incomplete sampling year when no Raine Island Corporation survey trip occurred.

between cohorts. Bustard ( 1 976) recorded no 2yr
or 3yr remigration intervals during a major part of
his nine consecutive years of total tagging census,
commencing in. 1964/ 1965, of the Heron Island C.
mydas population (single monel tagging in LI
position). The Applied Ecology project at Bramble
Cay recorded no remigration recaptures from four
consecutive years of total tagging census of the C.
mydas population during 1976/1977-1979/1980
(double monel tagging in LI and R1 positions.
Limpus et al., 2001 ). Based on long term titanium
tagging, Limpus et al. (1994a) recorded that the
majority of the C. mydas remigrants at Heron
Island returned after 5-8 yr intervals. These studies
indicate that the eastern Australian C. mydas
stocks may not be breeding on the usually
hypothesised 2-3 yr remigration interval for the
species (Hirth, 1997). Because of the incomplete

annual sampling of the nesting population at Raine
Island, the longer recorded remigration intervals
may be confounded by recaptures of turtles that
had actually remigrated in intervening years but
had not been recorded. The majority of the re-
migration intervals for the Raine Island C. mydas
population were within 4-6yr (Table 21). Most
tagging cohorts show a reduction in remigration
recapture rate at about eight years (Fig. 27). There-
fore the results from the 1 0 consecutive years of
large scale titanium tagging at Raine Island during
1984-1993 will be analysed for remigration
interval using only recaptures at less than nine
years following first tagging or the last recorded
breeding for a recaptured turtle. It is expected that
restricting the analysis of the first eight years of
remigration data for each cohort will minimise the
number of multiple cycles that are mistakenly

GREEN TURTLE POPULATIONS OF RAINE ISLAND

409

REMIGRATION INTERVAL (YR)

RIMICRATION INTERVAL (vr)

REMIGRATION INTERVAL (yr)

1995 TAGGING COHORT (n-1668)

REMICRATION RECAPTURES-180

iii'ii I l I I l —

2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17
REMIGRATION INTERVAL (yr)

2 2

- 1996 TACCINC COHORT (n-2997)

REMICRATION RECAPTURES-152

^1 — i — , , . — . — , — . —

M

» 6 7 I 9 10 II 12 1J 14 15 16 17

REMIGRATION INTERVAL (vr)

FIG 28 ( Cont .)

- 1997 TAGGING COHORT (n-1434)

REMICRATION RECAPTURES-60

N

I — r-^P^L — i — i — i — i — i — i — i — ' —

T 1 1 1

5 6 7 8 9 10 II 12 13 14 15 16 17

REMICRATION INTERVAL (yr)

included in the analysis, while capturing the
majority of the remigrations for the cohort.

It should be noted that the quantification of
remigration interval at Raine Island has been
partly compromised by the lack of continuity of
sampling effort. The standard December
sampling trips were not run in some years: 1983,
1990, 1994. The impact of reduced numbers of

recaptures resulting from not checking large
numbers of turtles for tags in these years can be
tracked through the sequential remigration
frequencies for each cohort (Fig. 28). Each cohort
was impacted at a different remigration interval.
For the combined 1 0 years of data from 1 984 to
1993, some of these resulting between-year
differences are expected to be smoothed.

410

MEMOIRS OF THE QUEENSLAND MUSEUM

1 2 3 4 5 6 7 8

REM IGRATION INTERVAL (yr)

FIG 29. Frequency distribution of emigration interval
for adult female Chelonia mydas recorded at northern
GBR rookeries pooled for ten annual tagging cohorts.
1984-1993. To minimise the confounding effect of
varying durations of study period for the cohorts,
only remigration recaptures spanning <9 yr have
been considered. To minimise the confounding effect
of tag design, only titanium tagged cohorts have been
included.

At Raine Island, short remigration intervals of
one and two years were extremely rare (0.33%
from a sample of 2,094 turtles) (Fig. 29). Three
year remigration intervals accounted for only
10.7% of recaptures. The modal remigration
interval for the Raine Island C. mydas population
was 4-5yr (Fig. 29). There are significant
differences in annual remigration intervals
among the annual tagged cohort from 1984
to 1993 (one way ANOVA: F9,2084 = 42.02;
p<0.001). No explanation is offered for the
abrupt increase in mean remigration interval
between the 1988 and 1989 cohorts (Fig. 30).
While some of the differences may be
attributable to the lack of continuity of sampling
(Fig. 28), many of the differences appear to result
from stochastic variability in the modal return
interval among cohorts. When a strong positive
SOI results in very depressed nesting numbers in
a particular season, it limits the number of
remigrants available for capture in that season.
For example, during the extremely low density
2000/200 1 nesting season, the remigration group
corresponding to that in 2000 was very depleted
within each cohort’s recaptures (Fig. 28).
However, a different remigration interval was
impacted for each tagging cohort. In contrast,
with high density nesting seasons such as
1999/2000, the remigration groups
corresponding to those in 1999 were atypically
high for their respective tagging cohorts (Fig.
28). It is apparent that there is no fixed
remigration interval that applies for this stock.
Remigration interval is variable in a complex
response to regional climate fluctuations.

ANNUAL TAGGING COHORTS

FIG 30. Variability in mean remigration interval
measured for 1 0 consecutive annual tagging cohorts
of adult female Chelonia mydas nesting at Raine
Island: 1984- 1993.

To further investigate remigration variability
from the perspective of the breeding season of
recapture, the remigration data (Table 21) was
reanalysed with respect to the year of recapture
(breeding season) within the 11 seasons
1991-2001 (Fig. 31). Again, the analysis was
restricted to the remigration intervals <9 yr.
Remigration intervals analysed from this period
encompass returns from the years since the
introduction of titanium tagging of the Raine
Island nesting population in 1983. A reasonable
comparison cannot be easily made with data
originating from the earlier years when turtles
were tagged with monel tags, given the high rate
of tag loss for monel tags ( Limpus, 1 992a). There
were significant differences in remigration
interval by nesting season of recapture for the
years 1991-2001 (1 way ANOVA: F 10 , 2 i 46 =
15.65; p<0.001). In addition there was a
significant linear correlation between the mean
remigration interval and the year:

Remigration interval = 0.08364*year- 161.5452
(F|. 9 = 8.30, 0.0 1 <p<0.025. r 2 = 0.48, DF = 9).

BREEDING SEASON OF RECAPTURE

FIG. 31. Variability in mean remigration interval
measured within 1 1 consecutive breeding seasons for
adult female Chelonia mydas nesting at Raine Island:
1991 -2001.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

411

200

CURVED CARAPACE LENGTH (cm)

FIG 32. Frequency distribution of size of nesting
female Chelonia ntydas recorded at Raine Island and
Moulter Cay during the 1 999-2000 breeding season.

Mean remigration interval has increased over
at least the past decade (Fig. 3 1 ).

No significant linear correlation was found
between the mean annual remigration interval
measured within the breeding season of recapture
and the mean May-October SOI measured 1.5 yr
before the breeding season (F, t9 = 0.007, p>0.25).

SIZE OF BREEDING ADULTS

FEMALES. A comparison of several methods of
measuring turtle size is shown (Table 23).
Because CCL has been the standard length
measurement of marine turtles used in Australia
(Bustard, 1972; Limpus & Walters, 1980;
Limpus, 1980b; Limpus et ah, 1983a,b) and
South Africa (Hughes, 1974a,b) and because of
the high correlation between CCL and the two
other length measurements in common use
(Carr's straight carapace length: Carr & Ogren,
1960; Straight Carapace length: Hughes,
1974a,b), CCL will continue as the standard
length measurement for this study. Should the
need arise for comparison with overseas studies.

BKEF.DINC SEASON

FIG. 33. Changing size (CCL) of nesting female Chel-
onia mydas recorded nesting at Raine Island and
adjacent rookeries in the northern GBR during 26
years of monitoring. * = remigrants; square = primary
taggings; dot = all turtles.

CCL can be transformed to other required lengths
using the equations of Table 23.

The carapace measurements of nesting females
are summarised in Tables 24-26 and Fig. 32.
Averaged over the 25 seasons of sampling in
November - December at Raine Island, the mean
female CCL = 105.97cm (SD = 5.142, range =
86.0 - 1 30. 1 , n = 20947. Table 24). Averaged over
6 of these seasons, mean female CCW = 98. 1cm
(SD = 5.04, range = 82.3 - 1 1 5.5, n = 2237, Table
28). Mean straight carapace measurements are
summarised for comparison with other studies in
Table 28.

There were significant differences in mean
CCL among annual November-December
samples (Table 24, Fig. 33). There has been a
significant downward trend in mean carapace
length of the nesting female C. mydas at Raine
Island and Moulter Cay during mid summer over
26 breeding seasons, 1976-2001 (Fl,23 = 28.66,
p<0.0005; r = 0.555, df = 23, 0.00 1 <P<0.0025).
Within this trend, whenever there were very high
density nesting seasons proximal to low density
nesting seasons, there were fluctuations in mean
CCL. On average smaller turtles bred in the high
density nesting seasons and larger ones in the low
density seasons. Parmenter (1980b) recorded
significant differences in mean CCL between
nesting seasons (1976-1 979) for Bramble Cay C.
mydas and attributed the differences to only the
larger size turtles being able to complete
physiological preparation for nesting in the low
density years. We have shown the size of the
annual nesting C. mydas population is a function
of the mean May-October Southern Oscillation
Index ~1.5yr before the commencement of the
breeding season. Therefore we tested for a

* 109

S 108
2 107
£

^ 105

< 104

< 103

a >02
3

> 101
ec

3 100

U

FIG 34. Examination of a possible regional climatic
regulation of the size of adult female Chelonia mydas
nesting at Raine Island. The mean May-October SOI
measured ~1.5yr before the breeding season (Table
19) was used as the measure of climate variability.
Turtle size = mean carapace length measured in
December (Table 24).

412

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2 4. Annual variation in curved carapace length (CCL) of nesting female Chelonia mydas at Raine Island
and adjacent Moulter Cay ##MC) during the mid summer nesting season. Primary denotes turtles tagged for the
first time in that season. Remigrant denotes turtles recorded breeding in a previous season.

Year

Rookery

All Turtles

Primary

Remigrant

Curved carapace length (cm)

Curved carapace length (cm)

Curved carapace length (cm)

Mean

(Range)

SD

n

Mean
( Ran S e >

SD

n

Mean

(Range)

SD

n

1976

L(Pe C )

RI

107.1

(93.5-124.0)

5.24

266

1977

(Dec)

RI

108.9

(99.5-120.5)

4.39

125

1978

L(. Dcc .)

RI

107.0

(90.5-125.0)

5.62

331

1979

(Dec)

RI

108.7

(96.5-125.5)

4.86

390

1980

(Dec)

RI

109.1

(96.0-124.0)

5.18

201

1981

L(Pec)

RI

107.0

(95.0-121.5)

4.96

538

1982

LiPec)

RI

108.0

(91.0-123.0)

5.11

504

1 1983-

1984

(Dec)

RI

106.4

(93.0-119.0)

4.81

489

1985

(Dec)

RI

107.5

(92.0-124.0)

5.41

400

107.2

(92.0-124.0)

5.47

303

110.0

(100.0-117.0)

4.07

30

1 1986

! (Dec)

RI

106.4

(93.0-125.0)

4.82

771

106.3

(93.5-125.0)

4.83

445

107.0

(95.0-121.5)

5.17

75

1987

(Dec)

RI

106.0

(87.0-123.5)

5.34

494

105.7

(87.0-118.5)

5.36

403

107.4

(98.5-116.5)

4.43

40

' 1988

1— <Pgg)..

RI

106.7

(93.0-122.0)

4.85

534

106.6

(93.0-122.0)

4.73

476

108.0

(95.5-120.0)

5.09

54

1989

(Dec)

RI

106.00

(89.0-120.0)

4.92

488

105.2

(89.0-120.0)

5.30

227

106.7

(97.0-120.0)

4.48

261

1990

(Dec)

RI

106.72

(90.0-126.0)

5.23

526

106.32

(90.0-126.0)

5.37

380

107.76

(98.5-118.5)

4.72

146

1990

(Dec)

MC

107.61

(95.4-120.5)

5.49

46

1991

( Dec >

RI

105.5

(90.9-119.4)

4.86

660

105.0

(90.9-119.4)

5.00

450

106.8

(91.2-117.3)

4.31

210

1992

(Dec)

RI

105.91

(89.5-123.0)

5.14

673

105.65

(89.5-123.0)

5.24

565

107.31

(98.5-118.0)

4.39

108

1992

(Dec)

MC

105.66

(93.0-118.5)

5.62

48

1993

(Dec)

RI

105.31

(87.8-122.0)

5.36

1360

105.14

(87.8-122.0)

5.37

1172

106.36

(92.8-119.4)

5.13

188

1994

(Dec)

RI

105.30

(86.0-123.5)

5.49

520

105.30

(86.0-123.5)

5.56

423

108.4

(97.5-120.5)

5.48

20

1995

(Dec)

RI

105.41

(88.8-123.7)

5.28

1733

105.30

(88.8-123.7)

5.26

1468

106.49

(94.6-120.5)

5.14

141

1995

(Dec)

MC

105.40

(86.6-123.7)

5.21

109

1996

(Dec)

RI

104.70

(86.5-127.5)

4.88

1945

104.47

(86.5-127.5)

4.90

1466

106.34

(92.5-118.4)

4.90

287

1996

(Jan)

RI

104.12

(90.2-121.0)

4.65

1180

104.15

(90.2-121.0)

4.73

822

105.61

(94.3-117.5)

4.44

88

1996

(Pooled)

RI

104.48

(86.5-127.5)

4.80

3125

104.36

(86.5-127.5)

4.84

2288

106.17

(92.5-118.4)

4.80

375

1997

(Dec)

RI

105.35

(87.2-130.1)

4.99

1861

104.82

(87.2-130.1)

4.99

1487

106.70

(90.6-121.3)

4.83

374

1997

1 (Dec)

MC

104.43

(92.0-119.1)

4.97

164

GREEN TURTLE POPULATIONS OF RAINE ISLAND

413

TABLE 24 (Cont.)

All Turtles

Primary

Remigrant

Year

Rookery

Curv ed carapace lengt

h (cm)

Curved carapace length (cm)

Curved carapace length (cm)

Mean

(Range)

SD

n

Mean

(Range)

SD

n

Mean

(Range)

SD

N

1998

(Dec)

R1

106.22

(91.0-126.5)

4.85

1715

106.08

(91.9-126.5)

4.80

1521

107.38

(91.0-119.7)

5.01

154

1998

(Dec)

MC

105.86

(94.2-117.4)

4.55

193

105.87

(94.2-117.4)

4.57

191

1999

(Dec)

RI

105.59

(88.4-126.1)

5.01

2071

105.13

(88.4-126.1)

5.04

1160

106.6

(92.3-125.5)

4.87

766

1999

(Dec)

MC

105.27

(92.3-118.4)

4.79

168

105.12

(95.7-118.4)

4.84

131

2000
j (Dec)

RI

106.93

(91.5-121.3)

5.39

735

106.87

(91.5-121.3)

5.43

661

106.95

(99.0-118.8)

5.02

—

57

2000

(Dec)

MC

106.50

(96.6-116.1)

3.99

85

106.59

(97.3-116.1)

3.81

81

2001

(Dec)

RI

105.58

(88.6-121.7)

4.92

1617

105.46

(91.6-120.7)

4.89

1145

106.00

(88.6-121.7)

4.98

361

2001

(Dec)

MC

105.00

(86.0-123.3)

5.04

281

105.02

(86.0-123.3)

4.97

270

relationship between the mean May-October SOI
(Table 1 9) measured -1.5 yr before the C. my das
nesting season and mean summer CCL measured
in December (Table 24). However, the mean
summer CCL of the nesting females did not show
a significant correlation with the mean SOI trom
- 1.5 vr before the breeding season (r = 0. 1 44, DF
= 22; F,,22 = 3.71; 0.1>p>0.05) (Fig. 34).

In contrast with the Raine Island results,
Limpus et al. (1984a) found no between-season
variation in the size of nesting C. mydas at Heron
Island over eight breeding seasons, 1974-1981.
Similarly there was no significant difference in
CCL among annual samples from No.7 and No.8
Sandbanks (Table 26), possibly because of the
small annual samples relative to the variance
within the samples. No significant difference in
CCL was found between the Raine Island sample
and the various northern GBR island samples
within any one year.

For all years except 2000 during the 17 yr
period, 1985 to 2001, remigrant nesting females
were significantly larger on average than females
that were tagged for the first time (Table 24). The
first-time-tagged group of turtles included new
recruits to the nesting population as well as turtles
that had nested in previous seasons but which had
not been tagged. Caretta caretta (Limpus, 1991 )
and N. depressus (Parmenter & Limpus, 1995)
nesting in the southern GBR and E. imbricata
(Miller et al., 2000) nesting in the northern GBR
display a similar characteristic of new recruits to
the breeding population being smaller on average
than remigrant females.

There were significant differences in mean
CCL among annual November-December
samples for both remigrant and primary tagged
females at Raine Island across the 1 7 yr period,
1985-2001 (Fig. 33; Table 24) (Primary tagged
turtles: F\ 6A3m = 15.71, p<0.0001. Remigrant
turtles: F , 63<2 55 = 2.75, p<0.001). There was no
significant downward trend in the annual mean
size of the primary tagged females across this
period (F U5 = 2.36, 0. 1 <p<0.25; r 2 = 0. 1 36, df =
15, p>0.25). Although the correlation was poor,
there was a significant downward trend in the
annual mean size of the remigrant turtles within
the same period (Remigrant turtles: F 1 , 1 5 = 7.08,
0.01<p<0.025;r2 = 0.321 df= 15 , 0.1<p<0.25).

There was no significant log-log correlation
between adult female C. mydas CCL and weight
for the small sample weighed after completion of

TABLE 25. Annual variation in curved carapace
length (CCL) of nesting female Chelonia mydas at
Raine Island outside the mid summer nesting season.

Year.

month

Curved carapace length (cm)

Mean (Range)

SD

n

1975 Feb

106.3 (98.0-117.0)

4.66

64

1980 Jun

110.0(110.0)

-

1

1981 Jun

100.0(100.0)

-

1

1982 Jul

104.3 (100.0-108.5)

-

2

1983 Apr

99.9 (94.5-104.5)

3.90

4

1984 Apr

106.1 (97.0-118.0)

4.59

77

1987 Jul

108.5(97.5-114.5)

6.55

4

2001 Feb

104.9 (92.9-114.4)

4.09

99

combined

105.6 (92.9-118.0)

4.55

252

414

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 26. Annual variation in curved carapace
length of nesting female Chelonia mvdas at No. 7 and
No. 8 Sandbanks during the mid summer nesting
season.

Year

Rookery

Curved carapace length (cm)

Mean (Range)

SD

n

1976

N8

105.6(98.0-112.0)

4.15

14

1979

N8

107.1 (101.0-118.0)

4.69

16

1986

N7&N8

105.1 (93.0-121.0)

4.42

119

1987

N7&N8

106.3 (93.0-121.0)

5.30

169

1988

N8

107.6 (95.0-117.0)

5.07

47

1989

N8

105.9 (99.5-119.0)

5.45

10

1990

N7

106.8(91.6-113.5)

6.14

12

1990

N8

107.5 (99.5-118.0)

4.61

V ]

1991

N7

104.9 (94.0-114.7)

4.63

85

1991

N8

104.6 (93.0-118.0)

6.03

56

1992

N7

106.9 (98.5-118.5)

4.16

53

1992

N8

106.6(94.5-115.5)

4.91

63

1995

N7

106.4 (91.8-121.6)

5.32

72

1995

N8

105.6 (97.1-117.4)

4.71

80

1996

N7

105.1 (92.2-117.9)

5.75

70

1996

N8

103.8 (92.7-116.3)

4.65

54

1997

N7

105.3 (87.2-114.0)

5.29

52

1997

N8

104.7(91.2-119.5)

5.11

185

combined

N7&N8

105.6 (87.2-121.6)

5.11

1194

laying: logiowt =1.28 x logioCCL - 0.491 (n =
24, r = 0.606, 0. 1 >p>0.05. wt in kg and CCL inc
m). The average female weighed 1 26.1 5kg (SD =
13.913, range = 1 0 1 - 1 54, n = 24; CCL of weighed
sample ranged 95-1 14cm).

MALES. A series of male C. mydas captured
while courting, or in association with courting
groups on Raine Island reef (Table 28) had a
pooled mean CCL = 99.4cm (SD = 4.89; range =
90.5-1 14.5cm; n = 37). A set of various carapace
measurements using the same methodologies as
used for the females is summarised in Table 28.
The breeding males, on average, were very much
smaller than the nesting females.

There was a significant log-log correlation
between CCL and weight of breeding males at
courtship: logiowt = 1 .65 x log I0 CCL - 1 .273 (n =
24, r 2 = 0.756, 0.002<p<0.005. Wt in kg and CCL
in cm). The average male weighed 97.63kg (SD =
9.528, range = 83- 1 22.5, n = 24; CCL of weighed
sample ranged 90.5 - 112cm).

SIZE COMPARISONS. The size of the nesting
female C. mydas is relatively uniform throughout
the GBR rookeries with a CCL = 107cm (Heron
Island: Limpus et al. 1984a; Bramble Cay:
Parmenter 1 979, Limpus et al., 200 1 ). The Raine
Island female C. mydas are of intermediate size

TABLE 27. Annual variation in curved carapace width
of nesting female Chelonia mydas at Raine Island and
Moulter Cay during the mid summer breeding
season. One way ANOVA: F 7 , 2 229 = 4.35; P<0.001,
significant.

Year

Curved carapace width (cm)

Mean (Range)

SD

n

1974/75

98.1 (86.0-111.0)

4.92

64

1976/77

97.5 (83.0-113.0)

5.26

263

1977/78

98.9(88.5-111.5)

4.48

125

1979/80

98.7 (86.0-115.5)

5.09

390

1980/81

98.8 (86.5-115.0)

5.37

201

1981/82

97.3 (84.0-108.5)

4.81

538

1982/83

98.2 (82.5-114.0)

5.07

504

for the species world wide, being approximately
the same size as the females from the Tortuguero
rookery in Caribbean Costa Rica and Aldabra
Island in the Indian Ocean (Hirth, 1980). Much
smaller sized females nest in Hawaii, the Arabian
Peninsula and the eastern Pacific Ocean while
considerably larger sized nesting females occur
at Europa Island, Surinam and Ascension Island
(Hirth, 1980; Miller, 1989).

The mean CCL of the dead females recorded on
Raine Island over 1 8 breeding seasons (Table 1 6)
was smaller than the mean CCL of the nesting
female sample recorded in the respective
breeding seasons (Table 24). Why the death of the
nesting female should be size related is not clear.
To address this question, detailed autopsies of
turtles dying from a variety of causes are needed
to identify the extent to which the mortality is
related to the turtle being small, irrespective of its
past breeding history, or to newly recruited turtles
to the breeding population. This bias in the death
of smaller turtles is contrary to the bias that would
be needed for the death of nesting turtles on these
islands to be the primary cause of the long term
reduction in CCL of turtles breeding at Raine
Island and Moulter Cay.

Size is not a reliable measure of maturity or age
in adult marine turtles (Limpus & Reed, 1 985a, b;
Limpus et al., 1994c; Limpus & Chaloupka,
1997) and not all turtles reach sexual maturity at
the same size (Limpus, 1991). Therefore the
question of why there are inter-rookery and
inter-seasonal differences in CCL of nesting
females probably cannot be resolved by studies at
the rookeries. The size of the breeding female
should be a function of a number of factors,
including its genetic stock, the climate of its
particular feeding area, its diet and its breeding
age. Potentially all these factors can now be

GREEN TURTLE POPULATIONS OF RAINE ISLAND

415

TABLE 28. Carapace measurements of adult male and
female Chelonia mydas breeding at Raine Island and
adjacent islands.

Measurement (cm)

Mean (Range) SD | n

Female

Straight carapace
length: 1978/79

100.3

(89.0-107.6)

4.37

54

Straight carapace
width: 1978/79

76.3 (68.6-90.0)

3.76

54

Carr’s straight cara-
pace length: 1978/79

99.7

(93.0-109.3)

3.69

29

Male

CCL: 1978/79

98.5

(90.5-112.0)

4.36

28

CCL: 1979/80

103.7

(103.0-104.5)

0.76

3

CCL: 1982/83

960 (-)

-

1

CCL: 1987/88

102.4

(90.5-114.5)

6.51

5

Curved carapace
width: 1978/79

88.8 (82.5-97.0)

4.50

28

Straight carapace
length: 1978/79

93.1

(86.9-102.5)

3.37

25

Straight carapace
width: 1978/79

71.0(66.1-78.2)

3.34

24

Carr’s straight cara-
i pace length: 1978/79

93.0 (88.4-98.4)

2.74

14

addressed in studies of C. mydas in its feeding
areas. The north-south distribution of the eastern
Australian feeding areas and the capacity to
identify the genetic stock of individual turtles
make the eastern Australian turtles ideally
located for further studies on what determines the
size composition of the nesting population.

EGG PRODUCTION

NESTING SUCCESS. Observations of
individual turtles ashore through the night have
shown that a female does not always successfully
lay her eggs on each nesting crawl. During 22
November - 16 December 1986, 26 nesting
females were followed for their entire time
ashore. Of these, only four were observed to lay
eggs (nesting success = 0.15).

It was concluded from the results of autopsies
of turtles dying while ashore at Raine Island that
only females attempting to nest come ashore.
When a turtle comes ashore and fails to lay her
eggs it should result primarily from negative
impacts on the turtle while it is ashore. Two
principal sets of factors that contribute to whether
or not a turtle will successfully lay during a
nesting crawl on these remote northern GBR
beaches are identified.

FIG 35. Adult female Chelonia mydas wandering into
each other as they searched for nesting sites at Raine
Island, 19 December 1984. The nett result of these
interactions is that turtles spread more widely across
the beach and spend more time ashore for nesting than
would occur in a low density nesting situation.

1 ) Environmental disturbance. The sand of Raine
Island beach was very dry, and without grass/tree
roots to bind it together. The coralline sand
particles did not adhere readily. The dryness and
looseness of the sand generally resulted in the
digging of very deep body pits with the
associated likelihood of collapse of the body pit
wall into the egg chamber. Egg chambers were
repeatedly collapsing as the females attempted to
dig their nests. Turtles with damaged hind
flippers had even more problems in digging if this
hampered their ability to lift sand from the egg
chamber. Bustard & Greenham ( 1 968) discussed
this problem for the Heron Island C. mydas
nesting population. Another potential cause of
disturbance to the nesting turtle on these islands
was buried rocks, timber and turtle bone. Large
obstructions could prevent the turtle digging an
egg chamber, but this type of problem occurred
less frequently. It mostly confronted those turtles
that attempted to nest on top of the phosphatic
limestone platform.

2) Turtle interactions. Most turtles, when
undisturbed, made their first nesting attempt for
the night near the crest of the outer beach rim.
When turtles were beaching at a density that
resulted in approximately overlapping turtles
around the perimeter of the beach in the early part
of the night, i.e. 1 turtle per 2m of beach line,
there was a high probability of a turtle en-
countering another turtle already ashore (Fig.
34). Therefore, with higher density nesting
(mean nightly tally count exceeding ~~900),
turtles that beached early in the night and become
established in nest digging were often disturbed

416

MEMOIRS OF THE QUEENSLAND MUSEUM

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•

•

0.6 -

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0.4 -

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0.2 -

■

■

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■

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■ SECTOR A

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• SECTOR B

i 1 ■ ■

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MEAN NIGHTLY TALLY COUNT (THOUSANDS)

0.25

0.2

0.15

s

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0.05

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0 2 4 6 8 10 12 14

MEAN NIGHTLY TALLY COUNT (THOUSANDS)

FIG 36. Variation in annual nesting success (A), rate of
clutch disturbance by nesting turtles (B) and clutch
equivalent egg mortality (CEEM) (C) for Chelonia
mydas breeding at Raine Island with respect to nesting
density. See Table 29 for seasonal data.

by later arrivals. This usually resulted in the one
or both turtles moving on to seek an alternative
nest site further in from the water line.
Disturbance that caused a turtle to abandon a site
ranged from one turtle bumping another to a
turtle that was digging a nest throwing sand into
the face of another. A person walking near or
gently touching a turtle wandering on the beach
platform on a high density turtle night appeared
to make no additional impact on the nesting
success of the turtle which was already reacting
to regular ‘disturbance".

It was not uncommon for turtles to be ashore
for over 6hr, alternately wandering and digging
without successfully constructing a nest before
returning to the sea. Without a completed nest,
the turtle would not lay. Difficulties in preparing

a nest and completing a laying will increase the
time a turtle spends on the beach and hence
increase the probability of still being ashore after
daylight. This in turn increases the probability of
heat induced mortality. Hence at Raine Island,
death on the nesting beach is density dependent
on the size of the nightly nesting numbers.

The nesting success recorded in 1986 above
contrasted with the nesting success recorded for
individual turtles during low density nesting in
April 1984 at Raine Island. Of 165 recorded
beachings, 109 (66%) resulted in laying, 41
(25%) resulted in the turtle leaving the beach
without laying and 15 (14%) were not assessed
for nesting success. During this visit, it rained on
7 of the 23 days/nights and the sand was
noticeably moist and egg chambers appeared to
be dug easily without the walls collapsing. The
combination of low nesting density (daily
number of beachings =3-17) (Table 3 ) and moist
sand appears to have contributed to the high
nesting success (= 0.73 for the 150 beachings
assessed for nesting success) at this time. These
data illustrate the variability that can occur in
nesting success for turtles attempting to nest in
coralline sand. Similar intra-seasonal variability
in nightly nesting success has been described C.
mydas attempting to nest in similar coralline sand
at from Bramble Cay (Limpus et al., 2001 ).

For a more rapid assessment of nesting success,
sampling sites A to E (Fig. 2) were selected and
where possible, nesting success was quantified
on opposite sides of Raine Island on the same
night. Nightly nesting success recorded in these
sectors for the November- December sampling
periods is summarised in Table 29. The mean
nightly nesting success data showed a significant
correlation with the mean tally count (MTC) for
the respective sampling period (Fig. 35A):
nesting success = -0.000042 x MTC + 0.441 1
(F|j 5 = 19.69; p<0.001; significant, r 2 = 0.565,
DF = 15.). Nesting success, measured at a
standard sampling period across several breeding
seasons, is therefore negatively correlated with
the density of the nesting turtles on the island.
However, factors such as how recently it rained
and the amount of rainfall can vary on a daily
basis within and between seasons and will
influence nesting success by changing the
adhesion of the sand particles.

When a turtle docs not lay her eggs on one night
she attempts to nest again on the same or on one
of the following nights (Miller, 1985). This can
be repeated a number of times until the turtle

GREEN TURTLE POPULATIONS OF RAINE ISLAND

417

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£

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7 • ■ ■

C- ■ ■ ■ | a ■

H 100 - _ ■ "

Z " ■

O 80 -
U

5 60 ^

H

jj 40 :

< 20 ^
w

S 0 H i * 1 f * > ' 1 » 1 '

-20 -10 0 10 20
SOI (MAY-OCTOBER) 1.5 YEARS BEFORE BREEDING SEASON

FIG 37. Comparison of the mean November- December
clutch count for adult female Chelonia mydas nesting
at Raine Island (Table 29) with the mean May-October
Southern Oscillation Index (SOI) approximately 1 .5 yr
before the breeding season (Table 19).

eventually lays her clutch. When a factor such as
nesting density contributes to lowered nesting
success, it results in the female having to make
more nesting crawls per clutch.

Within the one nesting season, because nesting
density will be low at the beginning and end of the
season and maximal in December and January,
nesting success must vary throughout the season.
In the absence of rain, the trend should be for
nesting success to be minimal in December and
January.

RENESTING INTERVAL. Renesting interval is
measured as the time from successfully laying a
clutch to the return of the turtle to the beach to
attempt to lay her next clutch. During most visits
to Raine Island, the high density nesting
precluded spending the time with each turtle to
establish its nesting success for the night. High
nesting density also made it difficult to examine
every turtle ashore for the night to check it for
tags and hence establish its identity. Therefore,
on most visits it was not possible to reliably
measure renesting interval at Raine Island and
Moulter Cay.

During the April 1984 visit to Raine Island,
nesting density was low enough for the nesting
success of almost every turtle for the night to be
assessed. Because of this, it was possible to
record the timing of subsequent returns of
females following a successful nesting. The
mean renesting interval was 12.4 d (SD = 1.02,
range = 10-14, n = 16). This is very similar to the
mean renesting interval of 12.0 d (SE = 0.04,
range = 9 - 2 1 , n = 979) and 1 2.4 d (SD = 1 .67,
range = 9- 1 9 d, n = 278 1 ) measured over an entire
nesting season at Bramble Cay during the
1978/79 and 1979/80 breeding seasons.

respectively (Parmenter, 1979; Limpus et al.,
2001). These values recorded within the nGBR
C. mydas stock are considerably shorter than the
renesting interval for the species at Heron Island
in the southern GBR (renesting interval: 13.52 d
in 1974/75; 14.08 d in 1980/81. Limpus, 1980b;
Limpus et al., 1 984a). These differences are more
likely to be the result of different water
temperatures in the interesting habitat during the
oviducal phase of embryogenesis and egg shell
formation (Miller, 1985) rather than a genetic
difference between the stocks.

NESTING BEACH FIDELITY. Once nightly
tagging of nesting turtles at Raine Island
commenced on each sampling trip there were
numerous recaptures of these tagged turtles on
subsequent nights at Raine Island. For example in
November 1 984, 77 recaptures of recently tagged
turtles were made over the 2nd to 10th night of
tagging on the island. These were of turtles that
had not completed a successful nesting on the
previous beaching. When Moulter Cay was
visited after a period of intense tagging at Raine
Island there were small numbers of recaptures of
nesting females that had been tagged while
attempting to nest at Raine Island within the
previous few nights (50 recaptures) (Table 8).
Two similar recaptures of turtles changing
islands have been made on Raine Island when
intense tagging on Moulter Cay was followed by
work on Raine Island (Table 7). There have been
no within season changes of rookery recorded
from the studies at MacLennan Cay, No. 7 and
No. 8 Sandbanks. There has been no record ol
intra-seasonal C. mydas interchange between the
northern and southern GBR rookery areas.

These observations are consistent with those
from more detailed long term recapture studies of
C. mydas at other rookeries (Heron Island:
Bustard, 1972, Limpus et al., 1984a: Sarawak
Turtle Islands: Hendrickson, 1958; Hawaii:
Balazs, 1980; Tortuguero: Carr et al., 1978). C.
mydas displays a high level ol fidelity to the
individual nesting beach with only a small
percentage of the females interchanging between
nearby rookeries w'ithin the same nesting season.

EGGS. C. mydas of the nGBR stock lay the
typical white, approximately spherical, solt
shelled eggs for the species. The mean clutch
counts recorded across the breeding seasons at
Raine Island and No. 7 and No. 8 Sandbanks are
summarised in Table 30. There was a significant
difference among the annual samples ol clutch
counts. The combined seasons mean clutch count

418

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 29. Nesting success and clutch disturbance measurements for Chelonia mydas at Raine Island.
Measurements made in sampling sites A and B (Fig. 2).

Turtles entering sector

Number of existing
clutches disturbed

Nesting

success

C lutch

Date

Sector

From beach

From sides

clutches laid

disturbance

rate

X

Y

Z

C

TJX

C/Z

12/12/81

A

222

-

75

25

0.34

0.33

13/12/81

A

170-200

-

57

35

-0.31

0.61

1981

Combined values

-0.32

0.46

29/12/82

A

52

-

20

2

0.38

0.10

3/12/82

A

93

-

34

5

0.37

0.15

6/12/82

A

93

-

25

5

0.27

0.20

1982

Combined values

-0.33

0.15

28/11/85

A

20

0

3

0

0.15

0

1/12/85

A

16

1

2

0

0.13

0

9/12/85

A

27

4

8

1

0.30

0.13

16/12/85

1985

A

Combined values

0.26

0.12

28/11/85

B

9

4

4

0

0.44

0

3/12/85

B

20

2

7

1

0.35

0.14

13/12/85

B

13

3

2

1

0.15

0.50

16/12/85

B

22

3

11

0

0.50

0

1985

B

Combined values

Combined values

0.38

0.08

28/12/86

A

313

-

37

20

0.12

0.54

4/12/86

A

449

-

50

17

0.11

0.34

8/12/86

A

520

-

47

28

0.11

0.60

12/12/86

A

369

-

47

21

0.13

0.45

1986

A

Combined values

Combined values

0.11

0.48

26/11/86

B

216

-

24

23

0.11

0.11

2/12/86

B

275

-

35

37

0.13

1.06

6/12/86

B

334

-

58

44

0.17

0.80

10/12/86

B

241

-

34

20

0.14

0.59

1986

B

Combined values

Combined values

0.17

0.82

3/12/87

A

330

-

35

30

0.11

0.86

1/12/87

B

73

-

24

7

0.33

0.29

6/12/88

A

78

10

39

5

0.50

0.13

7/12/88

A

74

17

31

0

0.42

o

8/12/88

A

76

21

50

9

0.66

0.18

1988

A

Combined values

Combined values

0.53

0.12

6/12/88

B

33

8

24

5

0.73

0.21

7/12/88

B

33

10

15

6

0.46

0.40

8/12/88

B

80

10

29

5

0.36

0.17

1988

B

Combined values

0.47

0.24

6/12/91

A

157

65

11

9

0.07

0.82

8/12/91

A

117

63

22

10

0.19

0.45

1991

A

Combined values

0.12

0.58

6/12/91

B

125

74

23

2

0.18

0.09

8/12/91

B

77

65

14

6

0.18

0.43

1991

B

Combined values

0.18

0.22

6/12/92

A

122

12

24

3

0.20

0.13

8/12/92

A

134

38

21

5

0.16

0.24

1992

A

Combined values

0.18

0.18

6/12/92

B

35

27

20

5

0.57

0.19

8/12/92

B

44

32

42

0.95

0.06

GREEN TURTLE POPULATIONS OF RAINE ISLAND

419

TABLE 29 ( Cont .)

Date

Sector

Turtles entering sector

Number of existing
clutches disturbed

Nesting

success

Clutch

disturbance

rate

From beach

From sides

clutches laid

X

Y

Z

C

TJX

C/Z

1992

B

Combined values

0.78

0.11

4/12/93

A

131

43

25

9

0.19

0.36

6/12/93

A

87

124

31

10

0.36

0.32

1993

A

Combined values

0.26

0.34

4/12/93

B

77

30

14

2

0.18

0.1

6/12/93

B

119

37

14

11

0.12

0.79

1993

B

Combined values

0.14

0.46

6/12/94

A

13

3

9

0

0.69

0

8/12/94

A

36

3

20

1

0.56

0.05

1994

A

Combined values

0.59

0.03

6/12/94

B

86

16

32

8

0.37

0.25

8/12/94

B

27

14

5

1

0.19

0.2

1994

B

Combined values

0.33

0.24

6/12/95

A

413

100

57

12

0.13

0.21

12/12/95

A

323

126

18

5

0.06

0.29

1995

A

Combined values

0.10

0.23

6/12/95

B

99

113

52

12

0.53

0.23

12/12/95

B

88

64

35

8

0.40

0.13

1995

B

Combined values

0.47

0.23

2/12/96

A

259

146

1

11

0.004

11.0

4/12/97

A

495

86

20

7

0.03

0.35

4/12/97

B

465

34

28

8

0.06

0.29

30/11/98

A

29

7

11

0

0.38

0

6/12/98

A

26

4

8

0

0.31

0

1998

A

Combined values

0.33

0

6/12/98

B

60

5

47

5

0.72

0.11

30/11/98

C

29

7

11

0

0.31

0

7/12/98

D

30

5

11

1

0.31

0.09

7/12/98

E

47

2

28

0

0.57

0

1/12/99

B

212

59

53

17

0.22

0.32

8/12/99

B

272

117

90

10

0.34

0.11

1999

B

Combined values

0.30

0.19

1/12/99

D

589

34

93

6

Incomplete

sampling

0.06

8/12/99

D

328

183

59

7

0.18

0.12

1999

D

Combined values

0.18

0.09

1/12/00

A

11

0

6

0

0.55

0

4/12/00

A

5

1

2

0

0.40

0

2000

A

Combined values

0.50

0

1/12/00

B

12

1

10

0

0.83

0

4/12/00

B

6

1

6

0

1.00

0

2000

B

Combined values

0.89

0

4/12/01

A

115

34

34

6

0.23

0.18

4/12/01

C

199

49

29

11

0.12

0.38

of 104.3 for the November- December nesting
period at Raine Island was very similar to the
mean clutch count of 103.8 measured for the two
samples from No. 7 and No. 8 Sandbanks. There

was no significant correlation between annual
November-December clutch count and the
southern oscillation index measured
approximately 1 .5 yr before the nesting season

420

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 30. Comparison of annual clutch counts of
Chelonia mydas from Raine Island and No. 7 and No.
8 Sandbanks.

Year

Clutch count

Mean | SD Range N

Raine Island: November- December

1976/77

102.7

19.02

63-147

48

1977/78

103.1

21.34

65-166

38

1978/79

105.1

19.31

69-143

59

1979/80

104.3

24.85

59-158

24

1981/82

111.3

17.25

79-152

20

1982/83

103.9

12.32

79-123

22

1984/85

117.8

22.13

62-175

36

1985/86

106.2

19.51

76-163

38

1986/87

93.9

14.50

66-130

39

1991/92

99.1

21.37

62-142

45

1992/93

101.8

14.0

73-130

54

1994/95

111.2

19.65

75-139

40

1995/96

100.7

23.6

47-124

9

1997/98

118

-

118

1

1998/99

91.1

18.92

65-142

20

2000/01

99.0

18

72-126

8

Combined

103.88

19.83

47-175

501

One way ANOVA: F\<m$ = 3.60; p<0.00l; significant

Raine Island: April

1983/84

(Apr)

83.3

17.82

62-111

7

No.7 and No. 8 Sandbanks: December

1988/89

116.0

26.55

88-147

4

1991/92

97.4

19.19

68-118

6

Combined

104.84

25.52

68-147

10

(Fj.h = 3.43; 0.2>p>0. 1 . r 2 = 0.196, DF = 14)
(Fig. 36). However, the clutch count from the
extreme El Nino and La Nina years are very
poorly represented in the samples (Fig. 36). This
relationship warrants further investigation.

No attempt has been made to record successive
clutches from. the same turtle within a breeding
season at Raine Island. The smaller clutches laid
in April (Table 30) are consistent with the decline
in clutch count that Parmenter (1979) recorded
with the last clutch for the season with each turtle
at Bramble Cay. The mid season clutch count
(Table 30) and egg diameters from Raine Island
(Table 32) is very similar to the December clutch
counts and egg diameters from Bramble Cay
(Parmenter, 1979) and the mean clutch count of
1 02.2 measured across the entire breeding season
at Bramble Cay in 1 979/80 (Limpus et ah, 200 1 ).
Clutch counts from Heron Island (Bustard, 1972;
Limpus, 1980b; Limpus et al., 1984a) are higher
than the clutch counts recorded in most years at
Bramble Cay and Raine Island. When compared

FIG 38. Portion of an ovary from an breeding female
Chelonia mydas that died while ashore for nesting on
Raine Island, 9 December 2002. In addition to the
normal 25-32mm diameter mature follicles (MF), this
turtle had many large 25-32mm diameter atretic
follicles (AF).

worldwide, C. mydas clutch counts and egg
diameters from the various rookeries scattered
within the GBR arc relatively uniform and occur
close to the middle of the range for the species
(Hirth, 1980; Miller, 1989).

Counting the number of eggs in a clutch at
Raine Island is complicated by the disturbance of
nesting turtles by other turtles, A turtle that has
been disturbed before she has laid any eggs from
the current clutch can be expected to return to lay
those eggs later on the same night or on a
subsequent night without reducing the number of
eggs in her clutch (Miller, 1985; Limpus, 1985).
However, if a turtle has laid part of her clutch and
is disturbed while laying, she can be expected to
cease laying the remainder of the clutch and
move off the nest with the following
consequences (Miller, 1985; Limpus, 1985). If
she has laid less than approximately half a clutch,
she can be expected to return to lay the remainder
of the clutch either on the same night or within the
next few nights. In this case she will have
deposited two ‘clutches’, each with less than less
than the actual clutch count. Should she have laid
more than half a clutch before she was disturbed,
she is unlikely to return to lay the remainder of
that clutch on the beach. She will normally return
in about two weeks to lay a complete clutch.
Some turtles can be seen laying small numbers
eggs as they return across the beach to the sea at
Raine Island. On occasions, turtles can be seen
underwater and resting under ledges along the
edge of Raine Island Reef with small clusters of

GREEN TURTLE POPULATIONS OF RAINE ISLAND

421

TABLE 31 . Frequency of occurrence of yolkless and multiyolked eggs in Chelonia mydas clutches at Raine
Island. Unless otherwise stated, measurements were recorded in November - December.

Year

Yolkless egj

5 s per clutch

Multiyolked eggs per clutch

Mean SD

Range

n

Mean

SD

Range

n

1976/77

0.05

0.224

0-1

20

0

0

20

1977/78

0.13

0.404

0-2

40

.

1978/79

0.10

0.399

0-2

60

0

0

60

1979/80

0.04

0.204

0-1

24

0

0

24

1981/82

0.18

0.529

0-2

62

_

1982/83

0.09

0.288

0-1

23

1983/84 Apr

0.14

0.350

0-1

7

0

0

7

1984/85

0.11

0.393

0-2

36

0.17

0.697

0-4

36

1985/86

0.11

0.383

0-2

39

0.03

0.160

0-1

39

Combined

0.12

0.399

0-2

349

0.03

0.290

0-4

224

One way
| ANOVA

F 9.339 = 0.37; p>0.25; not significant

F 6.217 = 1.69; 0.1>p>0.25; not significant

eggs under the tail. In both these cases, it is
presumed that these are females that have been
disturbed after they have laid more than half a
clutch and were voiding the remainder of that
clutch. For these turtles, the ‘clutch' laid on the
beach is less than a complete clutch and the
voided eggs represent a loss of eggs to the
rookery. Thus in high density nesting seasons,
there will be a number of apparent clutches

TABLE 32. Seasonal variation in Chelonia mydas egg
diameters at Raine Island and No.8 Sandbank. 10
eggs measured per clutch during November -
December.

^ Eg g diameter

Mean | SD [ Ran ge | N

Raine Island

1976/77

4.38

0.081

4.15-4.56

160

1977/78

4.44

0.163

4.10-5.38

400

1978/79

4.39

0.138

4.01-4.70

460

1981/82

4.37

0.129

3.93-4.67

200

1982/83

4.42

0.114

4.18-4.60

200

1983/84 (Apr)

4.33

0.179

3.86-4.61

70

1984/85

4.40

0.132

3.93-4.85

390

1985/86

4.36

0.119

4.03-4.58

240

1986/87

4.34

0.127

3.86-4.78

390

1988/89

4.33

0.126

4.11-4.58

540

1991/92

4.34

0.117

4.02-4.59

450

1993/94

4.25

1.87

3.66-5.09

950

1994/95

4.30

0.131

3.84-4.67

44

No. 7 and No.

8 Sandbanks

1988/89

4.31

0.161

4.11-4.50

40

1991/92

4.31

0.105

4.19-4.42

60

All combined

4.34

0.860

3.66-5.38

4594

I On e way AN O VA: F | 4 . 4S 79 = 1 56; 0. l>p>0.05; no t signi ficant

counted that will be less than a complete clutch.
The impact of this turtle-turtle disturbance in
altering the average number of eggs in a clutch
remains unquantified. Of greater consequence in
high density nesting seasons should be the
depletion of the stored fat reserves if the turtle
makes numerous unsuccessful nesting crawls
over an extended series of nights. The turtles seen
at necropsy that have large numbers of mature
sized, atretic follicles are believed to be turtles
from this latter category that have commenced
mobilisation of the nutrient and energy reserves
of some mature follicles to maintain their nesting
activity and possibly support their homeward
migration. For these turtles, the consequence of
repetitive disturbance would include a reduction
in the number of clutches she can lay for the
season. While the impact also remains
unquantified, there is a density dependent effect
in high density nesting seasons that should result
in a reduction in clutch count and a reduction in
the number of clutches laid for the season.

Yolkless eggs and multiyolked eggs were
uncommon and there was no significant
difference in the occurrence of yolkless eggs or
multiyolked eggs per clutch among the seasons
(Table 3 1 ). The low frequency of yolkless eggs
per clutch is typical of the species from the non
Arabian Peninsula rookeries (Miller, 1989).

There were significant differences in the mean
egg diameter between clutches laid by different
females within the one breeding season (one way
ANOVA, p<0.001 in each season). The seasonal
samples of clutch counts are summarised in Table
32. There were no significant differences in egg
diameters among nesting seasons or between

422

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 33. Depth from beach surface to top and bottom of Chelonia mydas clutches at Raine Island, measured
from the beach surface at the completion of laying.

Year

Nest depth to top of eggs (cm)

Nest depth to bottom of eggs (cm)

Mean

SD

Range

N

Mean

SD

Range

N

1977/78

70.0

_

70

1

85.7

9.81

74-98

3

1978/79

52.0

11.40

36-77

20

75.7

12.36

60-100

17

1979/80

51.8

13.53

33-88

24

91.1

19.73

64-152

16

1981/82

_

82.5

15.73

55-96

60

—

1982/83

_

78.2

7.78

60-95

1992/93

.

79.5

9.64

66-90

11

1994/95

81.0

18.12

65-109

5

Combined

52.3

12.61

33-88

45

81.63

14.73

55-152

138

One way
ANOVA

F 2 ,42 = 1 .01 ;p>0.25; not significant

F$j 3 i = 1.99; 0.1>p>0.05; not significant

Raine Island and No. 8 Sandbank. The mean size
of the eggs laid by nGBR stock C mydas (Table
32) are very similar is size to those laid by sGBR
stock females (Limpus et al., 1984a). However,
in the extremes, the smallest nGBR C. mydas
eggs were smaller than eastern Australian Caretta
caretta eggs (Limpus et al., 1984a) and the largest
eggs were comparable in size to eastern
Australian N. depressus eggs (Limpus, 1971).

Given the similarity of the recorded data from
Raine Island with that from Bramble Cay and the
difficulty of repetitively sampling the same
female at Raine Island to'determine valid whole
season values for most clutch related parameters
and for renesting interval, it is proposed that the
Bramble Cay clutch data and renesting interval
should be used to describe these parameters for
Raine Island turtles.

NEST DEPTH. Depths of eggs within the sand at
the completion of laying are summarised in Table
33. No significant inter-seasonal difference in
nest depth was detected (Table 33). The mean
depth to the top of a clutch was 52.3cm and the
mean depth to the bottom of a clutch was 8 1 .6cm.
These nest depths are considerably deeper than
nests constructed by sGBR stock females
(Limpus et al., 1984a).

EGG MORTALITY CAUSED BY NESTING
TURTLES. Some nesting turtles dig into existing
clutches (Fig. 37) as they prepare their own nests.
It is unusual for the entire existing clutch to be
disturbed, either by being dug out or rolling down
into the new egg chamber. When incubating
turtle eggs are moved there is a high probability
of mortality unless care is taken in their handling
(Limpus et al., 1979; Parmenter, 1980b). Eggs
dug up by nesting turtles are jarred violently and
even if they are not broken in the process, they are

most likely killed as they are thrown out. In
addition, eggs thrown to the surface and
surviving the jarring are most likely to desiccate
in the hot dry sand by day and die. thus the eggs
thrown out of existing clutches by nesting turtles
are regarded as totally killed. When these eggs
are eaten by birds it will be treated as scavenging
rather than predation, since birds are not the
primary cause of the mortality. At Raine Island
and Moulter Cay, buff-breasted landrails, rufous
night herons, silver gulls and all the species of
terrestrial crabs regularly scavenged on turtle
eggs brought to the surface by nesting turtles.
Egg mortality caused by nesting turtles was
measured in a two step process that quanti lied the
rate of disturbance of clutches by nesting turtles
and quantified egg mortality per disturbed clutch.
Clutch Disturbance by Nesting Turtles. Two
counts of clutch disturbance by nesting turtles
have been measured over the entire beach at
Raine Island during periods of low density
nesting. 1980/81, 8-12 December: In this very
low density nesting season only 1 clutch was

FIG 39. Loss of eggs as a nesting turtle digs into an
existing clutch at Raine Island, 19 December 1984.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

423

TABLE 34. Number of eggs destroyed per clutch when
nesting Chelonia mydas dug into existing clutches at
Raine Island. Samples were taken from throughout
the nesting habitat of the island.

Egg mortality per disturbed clutch

Mean | SD 1 Range I N

Raine Island

1981: 12 Nov-15 Dec

18.0

13.69

2-54

33

1982: 11 Nov- 10 Dec

21.6

15.53

6-62

21

1984: 21 Nov-23 Dec

23.3

13.15

5-61

49

1985:29 Nov- 19 Dec

14.0

13.24

1-71

44

1986: 22 Nov- 16 Dec

32.9

26.62

4-125

70

1987: 29 Nov-5 Dec

28.0

12.78

12-59

11

1988: 29 Nov-8 Dec

16.9

5.65

2-59

48

1994: 30 Nov- 10 Dec

33.5

18.2

1-82

4

1996: 1-12 Dec

10.8

8.8

2-40

80

1997:29 Nov- 19 Dec

18.2

12.95

1-62

74

1998: 30 Nov-7 Dec

8.18

7.15

1-25

11

1999: 28 Nov- 11 Dec

15.3

11.11

1-50

241

2001: 1-10 Dec

19.77

13.81

2-77

128

Combined

18.28

14.881

1-125

814

One way ANOVA: F l2Ml = 1 1.94; pcQ.001; significant

disturbed by nesting turtles in 5 successive nights
of observation. With approximately 8 1 nesting
females ashore per night on the whole island, if
nesting success is assumed to be at least 50% then
this would give an estimated nightly clutch
disturbance by nesting turtles for the entire island
= 0.005 (Table 35). 1 982/83, April: In five nights,
a total of 16 beachings occurred (Table 3) but no
incubating clutches were disturbed by the nesting
turtles. This is the only measure of clutch
disturbance from late in a nesting season.

For a more rapid assessment during higher
density nesting seasons, clutch disturbance was
quanti fied in the same sampling sites, A to E (Fig.
2), on Raine Island as were used for measuring
nesting success. Nightly clutch disturbance by
the nesting turtles during November-December
within these sites is summarised in Table 29.
These clutch disturbance data, combined with
those from following individual females, showed
a significant correlation with the mean tally count
(MTC) for the respective sampling period (Fig.
35B): clutch disturbance = 0.000548 x MTC +
0.9 1 1 8 (Fu 6 = 1 5.90; p<0.00 1 ; significant, r 2 =
0.4984, DF = 16.). Clutch disturbance by nesting
turtles, measured in a standard sampling period
across several breeding seasons, is therefore
positively correlated with the density of the
nesting turtles on the island. Since the density of
clutches in the beach will increase through the
nesting season until substantial numbers begin

TABLE 35. Turtle induced egg mortality - a summary
of the annual egg mortality per disturbed clutch
(EMpDC), rate of clutch disturbance (CD) and clutch
count recorded for Chelonia mydas at Raine Island
(From Tables 34, 29, 30 respectively.). The clutch
equivalent egg mortality per clutch laid (CEEM) was
calculated as: CEEM = (EmpDC x CD)/clutch. *
denotes an estimated value.

Sampling

period

EMpDC

CD

Clutch

count

Clutch equiva-
lent of eggs
killed per clutch
iaid

November-December

1980

-

0.005

-

0.001*

1981

18.0

0.46

111.3

0.074

1982

21.6

0.15

103.9

0.031

1984

23.3

0.80*

117.8

0.158*

1985

14.0

0.10

106.2

0.013

1986

32.9

0.65

93.9

0.228

1987

28.0

0.58

94.6

0.172

1988

16.9

0.18

102.2

0.030

1991

_

0.40

99.1

-

1992

-

0.15

101.8

.

1993

.

0.40

-

-

1994

33.5

0.14

111.2

0.042

1995

_

0.23

100.7

-

1996

10.80

11.0

-

-

1997

18.2

0.32

118.0

0.049

1998

8.18

0.04

91.1

0.040

1999

15.30

0.14

.

-

2000

_

0

99.0

0

2001

19.77

0.28

.

-

April

1983

-

0

-

0 11

hatching, clutch disturbance should also be a
function of when in the nesting season it is
measured. Thus the above mid season
measurements cannot provide a total seasonal
summary of clutch disturbance by nesting turtles.

Egg Mortality Per Disturbed Clutch. The number
of eggs dug from existing clutches when a nesting
turtle disturbed them was counted for a series of
clutches in most breeding seasons (Table 34).
There were significant inter-seasonal differences
in egg mortality per disturbed clutch (one way
ANOVA: F 1 2,80 1 = 11.94; p<0.001; significant)
but there was no significant correlation detected
between egg mortality per disturbed clutch and
the mean tally count of the sampling period (F,, n
= 0.305; p>0.25. r 2 = 0.027, DF = 1 1 ). The mean
egg mortality per disturbed clutch, pooled across
the nesting seasons was = 18.3 (Table 34).

The rate of egg loss per clutch laid during each
sampling period is expressed as a clutch

424

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 36. Straight carapace length and weight of hatchling Chelonia mydas from Raine Island, h denotes
hatchlings, c denotes clutches.

Breeding

Season

Straight carapace length (cm)

Weight (g)

Sample

Mean

SD

range

Mean

SD

range

1974/75: Feb

4.91

0.191

4.17-5.22

23.9

1.92

17.5-27.5

69 h, 7 c

1978/79: Dec

4.80

0.099

4.70-5.00

22.9

0.88

21.5-24.5

8 h, 1 cl

1979/80: Jun

4.83

1.978

3.84-5.17

22.6

1.6

18.0-25.0

57 h, 6 c

1982/83: Apr

4.83

0.160

4.31-5.14

24.5

1.95

20.0-28.5

100 h. 10 c

1983/84: Apr

4.87

0.183

4.25-5.43

23.8

1.91

18.5-29.5

460 h, 46 c

Combined

4.86

0.588

3.84-5.43

23.8

1.92

17.5-29.5

694 h, 70 c 1

equivalent egg mortality (CEEM) and is
tabulated in Table 35. When the annual CEEM
values for the December sampling period are
correlated against the respective mean tally
counts (MTC. Table 35), there was a very
obvious positive correlation of egg mortality
with nesting density (Fig. 35A): CEEM =
0.000036MTC x 0.002125 (F L8 = 14.88;
0.0025>p>0.00 1 . r 2 =0.6504; DF = 8). The clutch
equivalent egg mortality varied from a low value
of approximately zero in December 2000 and
0.005 in December 1980 to 0.23 during dense
nesting in December 1986. There is also the
potential for substantial intra-seasonal
variability, particularly in the early and late
breeding seasons relative to mid season. For
example, CEEM was approached towards the
end of the 1 982/83 nesting season. The data (Fig.
32) suggest that there will be a definable
mathematical relationship between CEEM and
the mean tally count that should have potential in
modelling the seasonal egg mortality caused by
nesting turtles. However, to achieve this it will be
necessary to systematically sample from
additional medium to very high density nesting
seasons to establish the relationship with any
precision. Additionally it will be necessary to
determine the variability of CEEM throughout
several breeding seasons with differing nesting
densities and relate the within season model to
the standard sampling period data that can be
gathered over a larger series on breeding seasons.

Bustard & Tognetti (1969) proposed that C.
mydas clutch destruction at Heron Island was
density dependent on the size of the nesting
population. However, they did not precisely
define their methodology. Bustard & Mathers
(1975, unpubl., reported in Bustard, 1976)
indicated that there was substantial destruction of
Caretta caretta clutches by nesting C. mydas at
Heron Island. Studies at Heron Island (Limpus
unpubl. data) have been unable to reproduce

comparable data sets to those in the above studies.
The data as presented by Bustard & Tognetti
(1969) suggests that nesting densities such as
those at Raine Island and Moulter Cay are un-
sustainable. It would be appropriate at this time to
investigate new r models to describe the possible
role density dependent egg destruction by nesting
turtles in regulating the population size.

While egg destruction is an obvious feature
observed on nights of dense turtle nesting at
Raine Island, it represents an egg loss equivalent
to only a small portion of the nightly egg
production — certainly in the early to middle part
of the nesting season. There is a large net gain in
eggs to the beach each night during the nesting
season. This turtle induced egg mortality in low
to medium nesting density years at Raine Island
is smaller than the 33-44% mortality of Bramble
Cay C. mydas eggs which are lost through
erosion each summer (Parmenter, 1980b; Limpus
et al., 200 1 ). In Surinam, Schulz ( 1 975) estimated
that C. mydas egg mortality from flooding and
erosion of nests was 30%, 38% and 44% in each
of the 1971, 1972 and 1973 breeding seasons
respectively. High egg mortality would appear to
be a natural feature of at least several widely
scattered C. mydas rookeries which are
functioning at very different nesting densities.

HATCHLINGS

Hatchling C. mydas were only examined in
detail from Raine Island. These hatchlings were
typical for the species with dark brown to black
dorsal colours and white ventrally. Dorsally the
trailing margin of the flippers and the carapace
margin were edged in white.

HATCHLING SIZE. There was no significant
difference in hatchling straight carapace length
(SCL) among the annual samples of freshly
emerged hatchlings (Mean hatchling SCL =
4.86cm) (Table 36). There were significant
differences in hatchling weight among the annual

GREEN TURTLE POPULATIONS OF RAINE ISLAND

425

TABLE 37. Frequency distribution scute pattern on
hatchling Chelonia mydas from Raine Island. * All
costal and vertebral' variability and most of the
remaining variability occurred in one clutch. # 3
hatchlings from one clutch sample each with 2 small
scutes between the pair of prefrontals = inter-
prefrontal scutes. One hatchling from another clutch
with an intervertebral scute between 1st and 2nd
vertebral scutes. ** preoccular scales were not
counted with all clutches.

1978

Dec

1979/

80

1983

April

1984

April

Total

Clutches examined

1

6

10*

46

63

Scute

Scute

count

Frequency

Nuchal

1

8

57

100

457

622

2

3

3

Vertebral

5

8

52

97

441

598

6

3

3

14

20

7

2

4

6

8

1

1

Post-

vertebral

1/1

8

57

98

450

613

1/2

2

4

6

2/2

6

6

Costal

4/4

8

53

97

450

608

4/5

1

2

3

5/4

2

4

6

5/5

2

1

4

7

5/6

1

1

Marginal

10/11

1

1

11/10

1

1

11/11

8

57

99

453

617

12/12

5

5

13/13

1

1

Postoccular

1/1

1

1

3/3

5

1

14

20

3/4

1

3

2

6

12

4/3

2

1

10

13

4/4

6

35

80

336

457

4/5

3

4

40

47

5/4

4

9

35

48

5/5

1

5

2

18

26

6/5

1

1

Preocular**

0/0

.

57

100

157

Prefrontal

2

8

57

100#

459

624

1

1

1

Post-

parietal

1

2

2

2

8

54

14

372

448

3

3

16

71

90

4

69

14

83

5

1

1

2

Infra-

marginal

3/3

2

2

3/4

1

3

4

3/5

1

1

4/3

2

5

7

4/4

8

54

97

435

594

4/5

1

1

2

5/4

1

9

10

5/5

5

5

samples (Mean hatchling weight = 23.80g)
(Table 36). The size of the hatchlings varied
significantly between clutches within each breed-
ing season (Table 36). The inter clutch differences
in hatchling weights may reflect varying degrees
of desiccation of the hatchlings as they dig their
way from the nest to the beach surface.

For a sample of 1 00 hatchlings from ten clutches,
there was a significant log-log correlation
between straight carapace length and weight:
logiowt = 1 .446 logioCCL - 1 .046 (wt in g, CCL in
cm; r 2 = 0.607, n = 1 00 hatchling, p<0.00 1 ).

These hatchlings were very similar in size to
emerged C. mydas hatchlings from Bramble Cay
in 1978/79 (mean SCL = 4.85cm; mean wt =
23.23g; Parmenter, 1979) and from Heron Island
in 1974/75 (mean SCL = 4.97cm, mean wt =
24.83g; Limpus, 1980b) and in 1980/81 (mean
SCL = 4.89cm, mean wt = 24.93g; Limpus et al.,
1984a). C. mydas hatchlings from both the
northern and southern GBR are similar in size
and collectively are near the middle of the size
range for hatchlings worldwide (Hirth, 1980).

HATCHLING SCALATION. The variability in
hatchling scute pattern is summarised in Table
37. The modal scute pattern of the Raine Island C.
mydas hatchlings was typical of the species: 1
nuchal, 5 vertebrals, 1/1 post vertebrals, 4/4
costals, 11/11 marginals, 4/4 postocculars, 0/0
preocculars, 2 prefrontals, 2 post parietal, 4/4
inframarginals (Pritchard, 1979).

INCUBATION AND EMERGENCE SUCCESS.
In assessing hatching and emergence success at
Raine Island, no attempt was made to quantify
the proportion of clutches that produced zero
emergence of hatchlings to the beach surface.
There is, therefore, no measure of total clutch
failure for this rookery. Total clutch failure can
result from erosion and flooding by high tides,
from problems within the egg that are derived
from the female herself and possibly from
microbial infections. At Raine Island there is an
additional problem of flooding by a rise in the
water table below the island following rain, with
very high tides or from storm wash over of the
beach. For example: in April 1984, following
heavy rain, the water table on the beach was
above the sand surface in the lower parts of the
beach depression (between the beach crest and
the cliff) with pools of water lying scattered in the
lower areas of the beach. Nests along this
depression that contained eggs yet to hatch would
have been flooded for several days. When turtle
eggs are immersed in water for more than a few

426

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 38. Hatching and emergence success of
Chelonia mydas clutches that produced hatchlings to
the beach surface at Raine Island.

1979 June

1983 April

1984 April

No. of
clutches

37

16

162

Hatchlings

Emerged
from nest

2814

74.37%

1278

82.77%

13115

78.35%

Live in
nest

7

0.18%

16

1 .04%

78

0.47%

Dead in
nest

13

0.34%

11

0.71%

81

0.48%

Eggs

Unhatched

256

6.77%

142

9.20%

1328

7.93%

Undevel-

oped

669

17.68%

94

6.09%

1942

11.60%

Predated
by crabs

25

0.66%

3

0.19%

195

1.16%

Total

3784

1544

16739

minutes they drown. Fig. 40 illustrates the
flooding of eggs by an elevated water table in
February 1997. That February visit to Raine
Island followed one of the largest recorded
December nesting populations, so it was
expected that vast numbers of hatchlings would
be crossing the beach daily. Instead, only tens to
low hundreds of C. mydas hatchlings were
crossing the beach nightly. In response to this
unexpected low hatchling production, 93 holes
were dug to nest depth at random locations within
each habitat type and each 100m sector around
the beach platform. Dead turtle eggs in situ in
nests were encountered in every hole. No live
eggs were found in any hole. It is highly likely
that only a trivial proportion of the eggs laid on
Raine Island during the massed nesting season of
1996/97 produced hatchlings.

Since 1996, flooding of much of the nesting
habitat during the mid nesting season has been a
regular event. Subsequent extensive flooding of
the nesting habitat, as evidenced by the water
table being visible in the bottom ot body pits,
occurred through at least early to mid December
of 1 998, 1 999 (I Figs 9, 1 0), 2000 and 200 1 . Eggs
that were laid up to 8 weeks prior to these
flooding events are expected to have drowned
and failed to hatch. Only 1 997 had a relatively dry
early nesting season (Fig. 7) and should not have
had the clutches subjected to this broad scale
flooding.

Hatchling emergence in February-March
which should originate from the peak period of
nesting has only been evaluated in February
1997. Most hatching and emergence success data

FIG 40. Chelonia mydas laying eggs into a water-filled
egg chamber on the cliff top at the western end of
Raine Island, early February 1997.

have been recorded from four samples of nestings
from the latter part of nesting seasons. The
hatching and emergence success are summarised
in Table 39. There was a low proportion of eggs
that showed no obvious embryonic development
(6-18% of eggs laid) and a high proportion of
hatchlings successfully emerged onto the beach
surface (74-83% of eggs laid). Predation of eggs
within the nests was low (0.19-1.1 6%) and could
be attributed to predation by the ghost crab
Ceratophthalmus cordimana , the only ghost crab,
living in the supratidal habitats. These values
apply only to clutches that produced hatchlings to
the beach surface.

On four occasions, emerged clutches were dug
to determine the proportion of clutches that had
been laid into an existing clutch:

1979, 13-14 December: of 38 emerged clutches dug, 1 was
a clutch on top of another and both these clutches had a
high hatching success. Only one of the 38 clutches
contained yolkless eggs.

1980, 8-24 June: of 78 emerged clutches dug, 3 were
clutches that had been laid into an existing clutch and in
each case both the clutches had a high hatching success.

1983, 24-29 April: of 18 emerged clutches dug, 2 were
clutches that had been laid into an existing clutch and in
each case both the clutches had a high hatching success.

1984, 3-25 April: of 162 emerged clutches dug, no clutch
had been laid into an existing clutch.

These observations indicate that the eggs remain-
ing in disturbed clutches can have a comparable
hatching success to those in undisturbed clutches.

Emerging hatchlings dug to the surface from
nests that had a mean depth of 80.6cm (SD =
9.073, range = 44-102, n = 200). This nest depth

GREEN TURTLE POPULATIONS OF RAINE ISLAND

427

TABLE 39. Hatching and emergence success data
(Table 38) from Chelonia mydas clutches at Raine
Island reanalysed on a clutch by clutch basis.

1 Mean SD Range

"71

1979: June (37 clutches)

—

Clutch (counted at
emergence)

102.27

22.345

68-168

37

Emerged hatchlings
per clutch

76.05

30.305

1-149

37

Live hatchlings in

nest

0.19

0.462

0-2

37

Dead hatchlings per

nest

0.35

0.538

0-2

37

Unhatched eggs per
clutch

6.92

9.867

0-41

37

Undeveloped eggs per
clutch

18.08

15.004

1-55

37

Crab predated eggs
| per clutch

0.68

2.028

0-10

37

Hatching success (%)

74.45

23.137

1.17-99.0

37

Hatchling emergence
success (%)

73.90

23.184

1.17-99.0

37

Nest depth (cm),
bottom

76.64

7.407

65-96

33

i 1983: April (16 clutches)

Clutch (counted at
emergence)

96.50

25.458

46-139

16

Emerged hatchlings
per clutch

79.88

25.458

46-139

16

Live hatchlings in
nest

1.00

1.317

0-3

16

Dead hatchlings per
nest

0.69

1.352

0-5

16

Unhatched eggs per
clutch

8.88

8.484

0-31

16

Undeveloped eggs per
clutch

5.88

7.822

0-32

16

Crab predated eggs
per clutch

0.19

0.544

0-2

16

Hatching success (%)

85.52

13.139

45.68-100.0

16

Hatchling emergence
success (%)

83.92

13.352

44.82-100.0

16

Nest depth (cm), bot-
tom

79.73

6.638

65-90

15

1984: April (167 clutches)

Clutch (counted at
emergence)

103.33

19.568

37-155

162

Emerged hatchlings
per clutch

80.96

21.600

18-139

162

Live hatchlings in
nest

0.48

1.035

0-8

162

Dead hatchlings per

nest

0.50

2.622

0-31

162

Unhatched eggs per
clutch

8.20

7.846

0-53

162

Undeveloped eggs per
clutch

11.99

12.882

0-91

162

Crab predated eggs
per clutch

1.20

2.735

0-18

162

Hatching success (%)

79.58

14.958

20.43-100.0

162

Hatchling emergence
success (%)

78.62

15.205

19.35-100.0

162

Nest depth (cm),
bottom

81.55

9.400

44-102

152

was not significantly different to the nest depth
recorded at laying (t>0.2) (Table 33).

The high emergence success of non disturbed
and non inundated C. mydas eggs incubate at
Raine Island (Table 39) indicate that they do not
suffer from the very high level of microbial
induced mortality associated with the high
density Lepidochelys olivacea rookeries of
Pacific Costa Rica (Cornelius 1986). The limited
data suggest that the hatchling emergence from
undisturbed nests will be very similar to that of
the low density C. mydas rookeries elsewhere in
the GBR, at Bramble Cay in 1978/79 (72.1%, n
=31; Parmenter, 1979) and at Heron Island in
1 966/67 and 1967/68 (87.9%, n = 26; 84.9%, n =
40 respectively; Bustard, 1976). Similarly the
hatching success of the undisturbed Raine Island
clutches was comparable to the hatching success
recorded for Heron Island C. mydas clutches in
1980/81 (mean = 90.2%, n = 60; Limpus et al.,
1983c).

HATCHLING PREDATION. 1975, 28
February -4 March: A typical day’s observation of
hatchling emergence and predation was as
follows:

4 March. 1500hr almost no hatchlings were crossing
beach. 1 5 1 5hr, immediately following the mid afternoon
heavy min shower, large numbers of hatchlings were
observed crossing the beach and entering the flat calm sea.
Bird predation was intense at this time. Sharks and large
fish were very obvious as they broke the smooth sea
surface while feeding on the hatchlings streaming away
from the island. The sharks were mostly Carcharhinus spp.
(black-tipped whalers and grey whalers) the largest of
which were in excess of 2m in length. The fish were mostly
trevally, Catwix sp. By 1630hr there was again intense
sunlight and few hatchlings were crossing the beach. From
1730hr onwards, even though there was only scattered
cloud, abundant hatchlings were crossing the beach. Vast
numbers of hatchlings were crossing the beach tliroughout
the night into the hours approaching dawn. Approximately
no hatchlings were crossing the beach during the mid
morning hours the following day.

Predation by birds: Rufous night herons were scattered
around the beach late in each afternoon and were observed
regularly to pick up C. mydas hatchlings. Some were eaten
on the beach but others were carried away. Regurgitated C.
mydas hatchlings were seen at rufous night heron nests
containing unfledged chicks. Crested terns w ere observed
on four instances to pick C. mydas hatchlings from the
water and on three instances to take hatchlings from the
beach. In each case the hatchling was subsequently
dropped and no further attempt to recover the hatchling
was made by the bird. The regurgitated remains of C.
mydas hatchlings were adjacent to numerous crested tern
nests on Moulter Cay (5 February). None of the other
species of birds (King, 1986) that were numerous around
the island during these mass daylight emergences of

428

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 40. Observed predation of Chelonia rnydas
hatchlings during emergence from the nest and the
beach crossing phase at Raine Island during 4-7 April
1 984. * denotes avian predators did not approach the
nest, probably because of its close proximity to the
bird hide and most hatchlings reached sea. The
numbers of hatchlings that successfully reached the
water are shown in parenthesis.

Date

Clutches examined

No. of
clutches

Predation on hatchlings
crossing beach

Not

quantified

100%

0%

Part of
clutch

4th

15

10

1 (104)

3(87)

1* __

5th

10

10

0

0

6th

25

21

0

3(75)

1*

7th

19

8

0

9(84)

i -

Total

69

49

1 (104)

15(246)

- « 1

hatchlings were observed to show any interest in the
hatchlings. Even the larger species of sea birds present
(brown, red- footed and masked boobies, lesser and greater
frigatebirds, red-tailed tropiebirds) were not predators of
the hatchlings.

Attraction of hatchlings to lights: When the deck lights of
boat anchored adjacent to the western end of Raine Island
reef w ere left on after dark, there was a dense carpet of
hatchlings across the smooth water surface in the
illuminated area within minutes. The hatchlings did not
swim from the illuminated area but swam back and forth
within it. Laige numbers of fish and shark could be seen
preying on the hatchlings. The light acted as a trap, holding
the hatchlings in a dense concentration, and made them
easy prey for the fish. When the lights were switched off the
hatchlings dispersed aw'ay from the island out into the open
sea. A Cancharhinus sp. (-150cm long) caught on the
western reef edge at 2 1 OOhr had a stomach content of 1 7 C.
mydas hatchlings, ranging from well digested to still alive.

1978, December A small crocodile, Crocodylus porosus ,
was captured on the reef flat (Limpus, 1980c). This
crocodile was using the island as a basking site by day and
appeared to be foraging along the beach margin at night. Its
stomach contained no food.

1980, 8-13 December: Ghost crabs, Ocypode ceratop-
thalmus and O. conlimana were abundant along the water
line at night. Both were observed preying on the hatchling
Chelonia nivdas. Several of the small grey terrestrial
hermit crab Xoenobita sp.. were present but they were not
observed to prey on turtle hatchlings.

1984, April: Rufous night herons were observed daily and
nightly on the beach throughout the visit.

These were observed aggregating around nests
and preying on the hatchlings as they emerged
from the nests (Fig. 41). Example predation case
histories:

1) At 1745hr on 7 April, 113 rufous night herons were
observed in the vicinity of one nest with 58 of them

FIG 41. Rufous night heron, Nycticorax caledonicus,
preying on a Chelonia mydas hatchling at Raine
Island, April 1983.

crow ded at the nest site. These birds were observed to eat
67 C. mydas hatchlings from this nest. The hatchlings were
taken as they emerged from the nest to the beach surf ace or,
for hatchling just below the sand surface and wriggling to
disturb the beach surface, the herons stabbed down to grasp
and pull the hatchling from the nest. The herons carried the
turtle hatchlings from the nest site and away from other
birds before sw allowing them whole.

2) On 11 April, observations over the southeastern beach
commenced at 1 600hr. A group of 1 5 rufous night herons
was present . Other birds in the vicinity included common
noddy, lesser frigatebird, silver gull, ruddy tumstone,
Australian ibis, crested tem. I620hr 25 rufous night herons
present; 1 650hr 9 rufous night heron present. 1710-171 5hr,
C. mydas clutch emerged with 50 hatchlings taken by
rufous night herons and 1 by a crested tem. Herons were
coming and going during the hatchling emergence and 39
w'ere still present after the hatchlings ceased emerging.

HATCHLING PREDATION RATES DURING
THE BEACH CROSSING. Hatchling predation
was quantified at two periods of different
densities of rufous night heron at Raine Island.

During April 1984, thousands of rufous night
herons were breeding at the island with chicks in
the nests. The number of clutches of C. mydas
hatchlings emerging per 24hr period was counted
on 13 days during 3-25 April 1984. The mean
daily number of clutches of hatchlings emerging
was 20.5 (SD = 5. 18, range = 14-32, n= 13). Over
four days and nights, 4-7 April, predation of C.
mydas hatchlings during the beach crossing up to
their entry to the sea was quantified for as many
clutches as possible. The results are summarised
in Table 40. From this sample, 350 hatchlings
reached the sea from 69 clutches (assuming
clutch count = 102, emergence success = 78.4%)
(Table 39). This is equivalent to a hatchling
survivorship while crossing the beach to the point
of entry to the sea = 6.7%. Almost all of the
associated predation was by rufous night herons.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

429

TABLE 41. Field data based on flipper tag recognition of female turtles
within the intemesting habitat tagged while nesting at Raine Island. These
data were used to calculate Petersen estimates with Bailey’s correction of
the number of breeding Chelonia mydas present (N) in the intemesting
habitat of Raine Island Reef during December in successive breeding
seasons. M = total number of marked turtles in the population; m = number
of marked turtles recaptured in the adjacent waters out of a sample of n
turtles examined for marks.

Date

Transect

type

Marking

Days since
marked

M

m

n

Estimate

N±SE

4/12/77

snorkel

Monel tag, LI

1

37

1

12

241 ± 128

6/12/77

snorkel

Monel tag, LI

1-2

49

1

16

417 ±226

4/12/78

speedboat

Monel tag. LI

1-3

496

8

79

4,409 ±1,313

1 4/12/81

speedboat

Monel tag, L3

1-27

2,421

34

197

13,696 ±2,071 !

2/12/84

snorkel

Titanium tag, L3

1-11

1,765

1

83

74, 130 ±42,286

3/12/84

snorkel

Titanium tag, L3

1-12

2,034

7

127

32,544 ± 12,732

8/12/84

speedboat

Titanium tag, L3

1-17

2,104

7

91

24.196 ±7,665

17/12/84

snorkel

Titanium tag, L3

1-26

2,362

2

44

35,430 ± 17,114

19/12/84

snorkel

Titanium tag, L3

1-28

2,475

6

97

34,650 ± 11,805

9/12/85

snorkel

Titanium tag. L3

1,385

4

10

3.047 ±919

[ 16/12/85

SCUBA

Titanium tag, L3

1,765

4

10

3,883 ±1,171 1

During June 1980 there
were only hundreds of rufous
night herons on Raine Island
and none were breeding. Few
birds aggregated on the beach
to prey on the hatchling turtles.

Estimates o f predation rates were
made by direct observation
and by examination of tracks
on the beach for hatchlings
emerged from 1 1 clutches: no
predation for 5 clutches; crabs
took 1 hatchling per clutch from
each of 5 clutches ( Ocypode
cordimana identified at two of
the hatchlings): an unidentified
bird took 1 hatchling from 1
clutch. From these 1 1 clutches
(assuming clutch count = 102,
emergence success = 74%)

(Tabic 39), the 6 hatchlings
preyed upon during the beach
crossing represent a hatchling
survivorship during the beach crossing = 99.3%.

The rufous night heron breeds on Raine Island
in large numbers during the main period of
hatchling emergence. It may require some 20-30
clutches of hatchlings to satiate these birds each
evening/night. When there are hundreds or
thousands of turtle clutches emerging nightly, the
birds will make little impact on the hatchling
numbers. However, when there are few clutches
emerging, most of the hatchlings may be eaten
before they reach the water. Outside the rufous
night heron breeding season these birds are minor
predators of the turtle hatchlings. This season-
ality of intense predation by this nocturnal bird
species on turtle hatchlings is a similar to that
observ ed with the rufous night heron predation
on N. depressus hatchlings on the beach at Crab
Island (Limpus et al., 1983b).

These observations suggest that the most pro-
ductive seasons with respect to hatchling survival
during the beach crossing to the sea will be the
dense nesting seasons. No attempt has been made
to quantify predation of hatchlings after they
entered the sea or to determine where they swim
after they disperse from Raine Island Reef.

ESTIMATION OF THE ANNUAL RAINE
ISLAND NESTING POPULATION SIZE

At present, the only precise method for
measuring the total annual nesting population at a
turtle rookery is via a total tagging census. This
requires every female using the rookery to be

tagged or checked for existing tags on every night
of the five months or more of the nesting season.
The remoteness of Raine Island makes it
logistically and financially difficult to monitor an
entire breeding season. In addition, the usually
large size of the nesting population also makes it
very difficult to tag every turtle present on any
one night at mid season.

From repetitive total tagging censuses of the
Heron Island Chelonia mydas rookery (Limpus,
unpubl. data) and the Mon Repos Caretta caretta
rookery (Limpus, 1 985) it can be established that
the nesting density at a standard sampling time
within the mid nesting season can be used as an
index of the size of the annual nesting population.
However, it would be an advantage it the absolute
size of the nesting population could be estimated
from population sample data recorded at the
island.

PETERSEN ESTIMATE. To reduce the problem
of widening confidence limits that result from
multiplying several factors, each with their own
95% confidence limits, a tagging - recapture
method was used to relate the number of nesting
turtles ashore on the island directly to the number
of turtles swimming in the intemesting habitat
surrounding the island (see Methods). This
method provides a single step calculation for
estimating the number of female turtles within
the nesting habitat.

The flipper tagging and transect data used to
calculate the Petersen estimates with Bailey’s

430

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 42. Data recorded during estimation of the size of the annual Raine Island Chelonia mydas nesting
population. The turtles were marked with paint on the carapace. Different colours or markings were applied on
different nights within a breeding season. The number of adult female Chelonia mydas in the waters adjacent to
Raine Island was estimated using a Petersen estimate, with Bailey’s correction.

Painting on beach

Counts by speedboat transects

Counts of adult female C. mydas

Counts of non ‘adult females’

Date

No.

painted

CM)

Paint code

Date

Paint code

Total

Short-

tailed

Adult-sized

(n)

Painted

(m)

C. mydas

C. mydas
Population
estimate

Long-tailed

adult

Immature

sized

E.

imbricata

: 1987: coloured oil-based paint

Speedboat transects

30/11/87

441

Yellow

1/12/87

Yellow

217

10

0

0

0

8,740 ± 2,458

1/12/87

241

Blue

2/12/87

Yellow

55

1

0

1

0

12,348 ±7,001

2/12/87

838

Red

2/12/87

Blue

2

4,499 ±2, 188

3/12/87

542

Blue-green

3/12/87

Yellow

184

3

2

1

0

20,396 ± 9,022

3/12/87

Blue

0

.

3/12/87

Red

5

25,838 ± 9,606

4/12/87

Yellow

327

3

3

2

0

36,162 ± 16,073

4/12/87

Blue

0

.

4/12/87

Red

5

45,811 ± 17,156

4/12/87

Blue-green

0

-

5/12/87

Yellow

356

3

3

1

0

39,359 ± 17,503

5/12/87

Blue

0

-

5/12/87

Red

12

23,013 ±6,037

5/12/87

Blue-green

0

-

SCUBA

1/12/87

Yellow

376

7

0

0

0

20,782 ± 6,853

2/12/87

Yellow

56

2

0

0

0

8,379 ±4,078

2/12/87

Blue

0

-

3/12/87

Yellow

239

2

0

6

0

35,280 ± 17529

3/12/87

blue

0

-

3/12/87

red

6

28,731 ± 10,009

1988: white oil-based j

)aint

Speedboat transects

30/11/88

402

Stripe

1/12/88

Stripe

402

19

9

1

0

1,970 ±384

1/12/88

237

Cross

2/12/88

Stripe

147

8

8

1

6,611 ±2,026

6/12/88

289

V

2/12/88

Cross

0

.

1

3/12/88

Stripe

161

8

0

0

0

7,236 ±2,224

3/12/88

Cross

2

12.798 ±6,339

7/12/88

Stripe

130

7

0

0

0

6,582 ±2, 126

7/12/88

Cross

3

7,762 ±3,4 18

7/12/88

V

5

6,310 ±2,330

SCUBA

1/12/88

Stripe

19

4

0

0

0

1,608 ±569

3/12/88

Stripe

46

3

0

0

0

4,724 ±2,021

3/12/88

Cross

2

3,713 ± 1,796

7/12/88

Stripe

23

1

0

6

0

4,824 ± 2,667

7/12/88

Cross

1

2,844 ± 1,572

7/12/88

V

0

-

1991-2002: Transects standardised to boat transects

6/12/91

950

Stripe

7/12/91

Stripe

435

30

0

0

0

13,361 ±2,276

8/12/91

600

V

9/12/91

Stripe

543

18

0

2

0

27,200 ± 5,975

9/12/91

V

2

108,800 ± 54,250

GREEN TURTLE POPULATIONS OF RAINE ISLAND

431

TABLE 42 ( Cont .)

Painting on beach

Counts by speedboat transects

C. mydas
Population
estimate

Counts of adult female C. mydas

Counts of non ‘adult females’

Date

No.

painted

(M)

Paint code

Date

Paint code

Total

Short-

tailed

Adult-sized

(n)

Painted

(m)

C. mydas

E.

imbricata

Long-tailed

adult

Immature

sized

1991-2002: Transects standardised to boat transects (cont.)

1992/93

Sea too rough to make counts

1993/94

Sea too rough to make counts

1 1994/95

Sea too rough to make counts

II 1995-2002: Paint standardised to Airpavc™

6/12/95

1,010

Lengthwise

7/12/95

Lengthwise

1,039

24

21

0

1

42,016 ±8,140 |

1/12/96

1,033

Lengthwise

2/12/96

Lengthwise

194

6

-

.

-

28,776 ± 9,990

7/12/96

1,526

Crosswise

8/12/96

Crosswise

489

28

-

.

_

25,784 ± 4,566

1/12/97

553

Lengthwise

2/12/97

Lengthwise

145

3

2

3

0

21,290 ±9,397

1998/99

Sea too roug

h to make counts

8/12/99

1,239

Lengthwise

9/12/99

Lengthwise

3*261

50

4

13

0

79,247 ± 10,903

10/12/99

1,147

Crosswise

11/12/99

crosswise

3,812

55

0

8

0

78,098 ± 10,268

4/12/00

86

Lengthwise

5/12/00

Lengthwise

39

2

2

0

0

1,147 ± 551

7/12/00

96

Crosswise

6/12/00

Lengthwise

30

2

1

0

0

889 ± 422

7/12/00

Lengthwise

54

5

0

1

0

788 ±281

8/12/00

Lengthwise

40

3

0

0

0

882 ± 374

8/12/00

Crosswise

0

-

6/12/01

1,188

Lengthwise

7/12/01

Lcngthise

1,024

8

1

3

0

135,300 ±
42,597

8/12/01

1,163

Crosswise

9/12/01

Crosswise

1,024

16

1

3

0

70,122 ± 16,390

correction of the number of adult female C.
my das in the waters surrounding Raine Island
during early December, 1977-1985, are
summarised in Table 41. While there is some
variability in the results from within a single
nesting season, the 95% confidence limits
usually overlapped.

When the method was first used, it relied on the
turtles being tagged distally on the trailing edge
of the front flipper (LI tagging position) with
monel tags. These distally applied tags were
highly visible (flashing in the sun light) on the
swimming turtle in the clear shallow waters
surrounding Raine Island. When the tagging
position was changed from LI to L3, the tag
became more difficult to distinguish. The tag
became even more difficult to detect on a
swimming turtle when the tag design changed, to
improve tag retention, from the highly polished
monel tag to the more dull lustred titanium tag.
Under the latter circumstances many more turtles
were being excluded from the underwater counts
of tagged/untagged females because of
uncertainty of recognising the presence/absence
of a tag, especially during speedboat transects.
This resulted in more time having to be spent
counting turtles with divers/snorkelers in the

water rather than using speedboat transects. With
the former came the increased frequency of
encountering big tiger sharks.

In 1987 a system of paint tagging the nesting
females was introduced (Figs 42-43) to increase
the visibility of a tagged turtle in the water. The
paint marks on the turtle carapaces held well for
about 4 days following application. Some turtles
were encountered back on the island after more
than 4 days with most of the paint mark
obliterated. Within the intemesting habitat, most
of the paint loss was caused by abrasion as turtles
pushed into crevices under coral. The paint
peeled off more readily if the carapace was not
dry when painted or if there was an algal coat on
the carapace. The different paint colours (Table
42) were not equally recognisable in the water.
For example, no green painted turtles were
‘recaptured’ swimming over the reef while all
other colours were recognisable. This lack of
recaptures was not because the green paint marks
had been shed by the turtles. Green painted turtles
were frequently encountered ashore on the island
on subsequent nights. Blue paint was only
marginally more visible than the green paint.
From 1998 onward, the paint tagging has been
standardised to white paint. White paint was very

432

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 42. Paint tagging a nesting turtle at Raine Island,
December 2000 as part of the Petersen Estimate of the
size of the total nesting population.

visible on the backs of the turtles in the water and
contrasting paint mark designs were easily
distinguished up to 30m underwater, even when
the turtle was barely distinguishable at that
distance. Since 1995, the paint tag has been the
more durable outdoor paint designed for road
marking. As it became apparent that very large
sample sizes had to be marked and examined to
provide estimates with tight confidence limits,
the underwater transects swum with snorkel or
SCUBA were phased out in favour of the speed
boat transects. This also provided for reduced
risk to the team given that large tiger sharks were
regularly encountered during swim transects at
Raine Island Reef.

A search of Raine Island Reef waters and the
water of adjacent reefs during December 1988
demonstrated that turtles nesting at Raine Island
were mostly concentrated in the waters
immediately surrounding Raine Island and were
not scattered widely throughout the region during
the intemesting intervals (Table 43).

Within any one sampling period, the
population estimates obtained using carapacial
painting to tag the nesting females were variable.
When Targe numbers of nesting females were
painted ashore at Raine Island and a very large
sample (thousands) of the turtles examined in the
intemesting habitat on Raine Island Reef, there
was usually reasonable agreement among the
estimates of the intemesting population size
(Table 42). However, the results are probably
influenced by sea surface conditions.

V.

.r.

A

t

'Vi 7 ™ w —

/ *

/

..y

FIG 43. A paint tagged female (longitudinal stripe) in
the intemesting habitat of the reef margin to Raine
Island Reef, December 2002. (Photo W. MacPharlane)

Unfortunately, the weather is not always
compatible with these types of studies and in at
least 4 years the counts on the reef flat could not
be undertaken because of the weather.

The relationship between the mean nightly
tally count and the estimated size of the
intemesting population in the adjacent waters is
illustrated in Fig. 44. For most years, the Petersen
estimate has correlated well with the mean tally
count. This has been particularly so for mean
tally counts up to about 4,000 females per walk of
the beach. At this mean tally count level, there is
an estimated intemesting population adjacent to
the Raine Island rookery of approximately
25,000 breeding female C. mydas. The
relationship is poor when there are very dense
nesting populations. In particular, in the 1999

TABLE 43. Numbers of turtles recorded during
speedboat transects over reefs adjacent to Raine
Island. 4-6 December 1988. Only Chelonia mydas
were identified during these transects. No painted
turtles from Raine Island or from Moulter Cay were
observed during these transects.

Date

Short-tailed

Long- tailed

Reef location
(transect time)

Adult sized

immature

4/12/88

76

0

0

Moulter Cay
(30 min)

5/12/88

1

0

0

11-093

0

5

0

11-095

1

0

0

11-096

0

0

0

11-097

0

0

0

McGillivray Cay
(20 min)

6/12/88

2

0

0

Jukes Reef

GREEN TURTLE POPULATIONS OF RAINE ISLAND

433

80

a

z

<

C/3

3

o

n

H

U3

H

P 40

C/3

H
Z

70

60

50

30 -

H

<

-

20

10

• MONEL TAG, Ll
o MONEL TAG, U

♦ TITANIUM TAG, L3

■ PAINT & SPEEDBOAT
a PAINT & SNORKEL

0 2 4 6 8 10 12 14

MEAN NIGHTLY TALLY COUNT (THOUSANDS)

FIG 44. Comparison of the Mean nightly tally count with the estimated
number of adult females in the interesting habitat of Raine Island.
The Petersen estimate was used to estimate the size of the interesting
population. See text for description of the tagging and transect count
methods.

season, the estimate of 78,672 ± 10,586 was not
expected from the mean tally count of 6,693.

However, this estimate would appear to be con-
sistent with the exceptionally dense population of
C. myclcis photographed in the Raine Island
intemesting habitat in late 1999 (Chadwick &

Doubilet, 2000). In the exceptionally dense
nesting seasons, there could be approximately
80,000 females in the Raine Island intemesting
habitat in early December.

Estimating the size of the total annual nesting
population for Raine Island via a single Petersen
estimate calculation using data derived from
marking on the nesting beach and ‘recapture* in
the intemesting habitat in the days immediately
following tagging is cost effective. The method
warrants further investigation to improve its rigor
in the very high density nesting seasons.

Carapacial painting is a workable method for
tagging the turtles, provided the paint marks are
prominent and of a suitable colour. Selected colours
and/or paint marks can be used to distinguish
different tagging cohorts. To maximise the
precision of the estimate, it is essential to
maximise the proportion of the total population
of intemesting turtles tagged in any one nesting
sample and to maximise the number of turtles
examined in the aquatic recapture phase. In this
regard, further investigation of the painting
technique would be appropriate to develop a
method for paint tagging many thousands of
nesting females in a single night. At the same

time the paint-retention time needs
investigation to improve the
capacity for pooling painted cohorts
from different nights.

Within any one nesting season
there will be different numbers of
active nesting females present at
different times of the season, with
small numbers present in early and
late season and maximum numbers
present in mid season. From total
tagging census studies at Bramble
Cay in 1 979/80, approximately 85%
of the total annual nesting
population visited the island during
the last two weeks of December
(Limpus et al., 2001). This should
equate to the total population in the
internesting habitat in early
December. In addition, there is a
strong correlation between the
nightly number of nesting C. mydas
using Moulter Cay (MCTALLY)
(Table 4) with the nightly tally count
Raine Island (R1TALLY) (Table 3) on the same or
adjacent night: MCTALLY = 0.3928 x
RITALLY + 95 (F U4 = 47.6; p<0.005. r = 0.773,
DF = 14.). By combining these two values it is
possible to obtain an estimate ol the size ot the
total annual nesting C. mydas at Raine Island,
Moulter Cay and immediately adjacent islands:

= (Raine Is December estimate + Moulter Cay
December estimate) / 0.85
= (RI December estimate + 0.3928 X Raine Is
December estimate + 95) / 0.85
= 1.6386 x Raine Is December estimate -M l 2
Thus the estimated total annual nesting
population for Raine Island and Moulter Cay in
the very high nesting density seasons would
approximate 131,000 females. The total nesting
population in the extremely dense nesting
seasons for the entire nGBR stock will be slightly
higher if a few thousand additional females
nesting at Bramble Cay, Murray Islands, No. 7
and No. 8 Sandbanks, Milman Island and the
other C. mydas rookeries of the northern GBR
and Torres Strait are added. A more typical dense
nesting population with a mean tally count of
about 4,000 females at Raine Island, may be in
the order of 41,000 females breeding annually.

The nGBR stock, for which Raine Island -
Moulter Cay are the primary breeding sites,
supports the largest nesting aggregation of C.
mydas remaining anywhere in the world (King,
1982; Groombridge & Luxmoore, 1989).

434

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 44. Frequency distribution of the presence and absence of corpora
albicantia on the ovaries of breeding adult female Chelonia mydas
examined at Raine Island. The proportion of females with no corpora
albicantia provides an estimate of the rate of recruit of new females into the
breeding population (± 95% confidence limits).

Breeding

season

Method of
assessment

Nesting females examined

Recruitment

Total

With

albicantia

Without

albicantia

Estimated

value

95%

confidence

limits

1989, Dec

Necropsy

98

85

13

13.3%

7.2-21.6%

1990, Dec

Necropsy

Laparoscopy

8

52

59

1

1.7%

0.04-8.9%

1991, Dec

Necropsy

Laparoscopy

29

50

77

2

2.5%

0.3-8. 8%

1992, Dec

Necropsy

Laparoscopy

2

97

93

6

6.1%

2.3-12.7%

1993, Dec

Necropsy

33

33

0

0%

-

1994, Dec

Laparoscopy

51

44

7

13.7%

5.7-26.2%

1995, Dec

Gonads sampled by non-standard method. Data deleted from analysis.

1996, Dec

Necropsy

35

34

1

2.9%

9.9-14.8%

1997, Feb

Necropsy

100

100

0

0%

0.0-3. 6%

1997, Dec

Necropsy

52

48

4

7.7%

2.1-18.5% ;

1998, Dec

Necropsy

1

1

0

-

' 1999, Dec

Necropsy

65

53

12

18.5%

9.9-30.0%

2000, Dec

No freshly dead turtles available during study.

2001, Dec

l!= =====k

Necropsy 16

14

2

12.5%

1.6-38.8%

ESTIMATION OF ADULT RECRUITMENT
RATE

Ovaries of samples of breeding female C. mydas
at Raine Island were examined for the presence or
absence of corpora albicantia (Fig. 45). Corpora
albicantia are ovarian scars from ovulations in
previous years. Hence, females without corpora
albicantia are in their first breeding season i.e. new
recruits to the breeding population. The proportion
of breeding females without corpora albicantia
provides a measure of the recruitment rate of new
adult females into the breeding population (Table
44). Commencing in 1 989, an attempt was made to
quantify the annual recruitment of new turtles into
the Raine Island breeding population.

The annual recruitment rate of new adults into
breeding population is low: mean = 7. 17% (SD =
6.4 1 , range = 0-1 8.5%, n = 1 1 ) (Table 44, Fig. 46).

These measurements are based on two under-
lying assumptions. 1 ) the rate measured on dead
females at Raine Island is similar to the rate for the
entire population, i.e. the nesting turtles that died
on the island were a random sample of the entire
nesting population with respect to recruitment. 2)
that the recruitment rate measured in early
December is representative of the entire breeding

season. This assumption needs
testing with C. mydas given
that new recruit C. caretta and
N. depressus tend to arrive later
in the nesting season at south
Queensland rookeries (C.J.
Limpus and C.J. Parmenter,
respectively, unpubl. data).
This assumption was tested
during the 1996 breeding
season only and similar results
were obtained in early
December and early February
(Table 44). Although the
results from laparoscopy and
necropsy provide no indication
of a bias from the dead turtles
(Table 44), this assumption
warrants further testing.

There are few studies of
marine turtles against which to
judge the significance of these
recruitment values. This recruit-
ment rate for the Raine Island
C. mydas is considerably lower
than that estimated from
tagging studies with Caretta
caretta (30-50%, Natal, South
Africa, Hughes, 1989; 30-40%, Georgia, USA,
Richardson & Richardson, 1982; 44.2% ± 5.7%,
Queensland, Australia, Limpus, 1985). However,
all of these values for C. caretta have been
measured in depleted populations. In contrast, the
annual recruitment rate estimated from tagging

FIG 45. An ovary with corpora albicantia. This ovary
was from an adult female Chelonia mydas that died
while ashore attempting to nest at Raine Island,
December 2002. MF = mature ovarian follicle; AF =
atretic follicle; PVF = pre-vitellogenic follicle; CA =
corpus albicantium; CL = corpus luteum.

GREEN TURTLE POPULATIONS OF RAINE ISLAND

435

~ 100
w 90

5 80
2

H 70
Z

S 60
2

t 50
g 40
3 30

< 20

1 10
< 0

1985 1990 1995 2000

BREEDING SEASON

FIG. 46. Annual variability in recruitment rate of new females into the
Chelonia mydas breeding population at Raine Island.

studies with the stable population of the eastern
Australian N. depressus stockbreeding at Mon
Repos, Peak Island and Wild Duck Island was in
the range of 10-20% (Parmenter & Limpus, 1 995;
Limpus et al., 2002). The mean annual recruitment
rate estimated from gonad examination using
laparoscopy with the declining E . imbricata
population the breeds in the northern GBR during
1991-2000 was 15.5% (SD = 6.65, range = 8.3 -
27.3%, n = 8) (Miller et al., 2000). Thus, the
recruitment rate of new adults into the nGBR C.
mydas stock is only slightly lower than the recruit-
ment rate estimated for the stable population of A.
depressus and the declining E. imbricata
population that breed in eastern Australia.

ACKNOWLEDGEMENTS

This report is dedicated to Dr G Saunders and
Dr H. Lavery who recognised the biological
importance of Raine Island and whose support
and encouragement enabled the project to
commence. This research was conducted as part
of the Queensland Turtle Research Project of the
Queensland National Parks and Wildlife Service.
It has been funded in part by grants from the
Raine Island Corporation, the Australian
Department of Science and the Australian
National Parks and Wildlife Service. Ben Cropp
provided the opportunity for one of us (CJL) to
visit the island in March 1975. The visit in 1976
was funded in part by the ABC Natural History
Film Unit for a filming project with QPWS staff.
During 1976 -1978, a parallel turtle study was
conducted at Raine Island by staff of the Torres
Strait Turtle Farm Project of Applied Ecology Pty
Ltd under the leadership of one of us (CJP).
Results of this study have been incorporated in
the present study.

Annette Fleay, Valonna Baker,
David Butler, Margaret Card,
Michael Cassematy, Robyn
Delaney, Carl, French, Matt Freher,
Emma Gyuris, Mark Hamann. Brad
Jones, Steven Neil, Mark Read,
Darryl Reimer, Linda Rhineholdt
and numerous volunteers collected
data under the primitive field
conditions on the islands. Brian
King provided valuable team
coordination for many study trips to
Raine Island and worked in
collaboration with us in document-
ing the significance of turtle nesting
on the islands of the northern Great
Barrier Reef.

Ken and Betty Butler, through their personal
commitment to conservation in Queensland
which led to the establishment of the Raine Island
Corporation, have made a lasting contribution
conservation within the Great Barrier Reef and to
marine turtle conservation within the Austral-
asian region. This assistance and support is
gratefully acknowledged.

CONCLUSION

Raine Island has been a nesting site for green
turtles for over 1.000 years. The nesting
population of green turtles has been known to
Europeans for approximately 175 years during
which a navigation beacon tower was built,
guano was mined and turtles were harvested.
Green turtles nesting at Raine Island are part of a
larger genetic stock that utilizes many islands in
the northern Great Barrier Reef (nGBR) and
eastern Torres Strait. This population is one of the
largest remaining stocks for the species on a
global scale. The nesting green turtles of the
nGBR are morphologically similar to other
genetic stocks of eastern Australia and the Gulf of
Carpentaria, as well as elsewhere in the world.
The population is chatacterised by a very long
remigration interval with a mean of 4-5 years.
The number of turtles using Raine Island varies
by orders of magnitude among seasons. Fluctu-
ations in numbers of nesting turtles are correlated
with alterations in the ENSO measured 18
months preceding. Intra-annual variation also
occurs; the highest density nesting occurs during
the mid summer months and the lowest density
nesting occurs during the winter months.

Adult mortality on the nesting beach and egg
mortality were positively correlated, while
predation by sharks on the nesting turtles and

436

MEMOIRS OF THE QUEENSLAND MUSEUM

nesting success were negatively correlated with
an index of the size of the annual nesting
population. In high density nesting seasons
following repetitive disturbance females reduce
the clutch production as a result of resorption of
ovarian follicles. Death of nesting turtles is a
natural phenomenon and occurs primarily from
heat exhaustion on the beach by day following
extended unsuccessful nesting attempts.

There has been a progressive decline in the size
of nesting females over the decades of the study.
This decline was associated with a decline in size
of remigrant turtles. There was no decline in the
size of turtles being tagged for the first time. The
decrease in size of the females occurred in con-
junction with a progressive increase in remigration
interval. In recent times, the nesting population is
also characterised by a very low recruitment rate.
These demographic characteristics may indicate
that this large population is in the early stages of
decline as a result of loss of adult turtles. There is
wide spread hunting of turtles from this
population that preferentially targets the adult
and near adult size (Schulz, 1984; Kwan, 1991;
Johannes & MacFarlane, 1991) that could
account for these changes. In addition, if the
recent phenomenon of very low hatchling
production resulting from flooding of the Raine
Island nesting habitat continues, then significant
declines can be expected in the numbers of young
C. mydas in the dispersed foraging areas. Taken
together, there is a high probability that this
population will decline in the foreseeable future.
It is recommended, that monitoring of the impact
of human induced mortality on the population
throughout it dispersed foraging areas be
increased and that steps be taken to reduce human
mortality to a sustainable level.

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AOUPINIA , A REMARKABLE NEW GENUS OF ADELIINI FROM NEW CALEDONIA

(COLEOPTERA: TENEBRIONIDAE)

ERIC G. MATTHEWS

Matthews, E.G 2003 06 30: Aoupinia , a remarkable new genus of Adeliini from New
Caledonia (Coleoptera: Tenebrionidae). Memoirs of the Queensland Museum 49(1):
441-445. Brisbane. ISSN 0079-8835.

Aoupinia pseudohelea, new genus and species, is described from Aoupinie Special Fauna
Reserve, New Caledonia, where it occurs in rainforest leaf litter. Although strongly
resembling the unrelated Australian tenebrionine Helea Latreille, Aoupinia belongs to the
lagriine tribe Adeliini, a Gondwanan group which has diversified extensively in New
Caledonia. There is an analysis showing that the new taxon belongs to a group of ten
Australian and New Caledonian genera termed the northern squalid group, but that it is of
uncertain relationship vis-a-vis particular genera within this group. □ Coleoptera,
Tenebrionidae , Adeliini , New Caledonia, new genus.

Eric Matthews, South Australian Museum, North Terrace, Adelaide 5000, Australia (e-mail:
matthews.eric@saugov.sa.gov.au); 27 August 2002.

New Caledonia has a diverse representation of
the Gondwanan tribe Adeliini, which also occurs
in Australia, New r Zealand and Chile (Matthews,
1998). Kaszab (1982), in his revision of the
Tenebrionidae of the island, described and keyed
eight genera and 50 species of Adeliini, and 20
additional species were described after two
subsequent visits there by G.B. Monteith of the
Queensland Museum (Kaszab, 1986). While it
was evident from the small number of known
specimens of some taxa that the tenebrionid
fauna of New Caledonia was still imperfectly
known, it nevertheless came as a surprise to
discover there a taxon of Adeliini which was
completely different from any other member of
the tribe, one which in fact bears a striking
external resemblance to the unrelated Australian
genus Helea Latreille.

A single specimen of this species was found in
late 2000 in the Reserve Speciale de Faune de
1’ Aoupinie by G.B. Monteith. This reserve
straddles the 1000m Massif Central which forms
the east/west divide in the centre of the island and
is traversed by a road which crosses between
Poya and Ponerihouen servicing a radio repeater
station on the summit. In the following year three
more specimens were found in the same place by
the same collector.

SYSTEMATICS
Aoupinia gen. nov.

TYPE SPECIES: Aoupinia pseudohelea sp. nov.

DESCRIPTION. Oblong, castaneous, alutaceous,
both dorsal and ventral surfaces squalid, covered

with a thin layer of fine soil retained by numerous
small, curved, thickened setae disposed over
entire body and legs, not tuberculate.

Head. Clypeus with anterior edge convex,
straight in middle. Clypeo-frontal suture deeply
impressed. Frontal grooves present, distinctly
impressed. Eyes strongly transverse, surrounded
by a groove. Antennae not quite reaching base of
prothorax, antennomeres (Fig. 3A) 1-7 more or
less oblong or a little obconic, with sparse long
setae, 3 as long as 4 and 5 combined, 7 1 .5 x as
long as 6, 8-11 moniliform, paler and slightly
wider than preceding segments, densely covered
with very short setae as well as a few longer
bristles. Mentum trapezoidal, depressed on basal
and middle areas, a little raised anteriorly in
middle. Terminal maxillary palpomere enlarged
and securiform.

Prothorax. Pronotum with sides expanded into
wide flanges which arch fonvard over head. Hind
edge with 4 projections partly overlapping elytral
bases. Prostemum long before coxae, its anterior
edge raised into a prominent convex collar (Fig.
2, c). Prosternal process flat, moderately wide,
slightly expanded posteriorly with almost
straight hind edge.

Pterothorax. Elytral disc smooth, not tubercu-
late. A wide reflexed flange emerging along
entire length of elytron just outside 8th stria. No
epipleuron evident underneath, outside edge of
elytron with narrow beading. Wings absent.
Meso- and metaventrites as in Fig. 2.

Legs. Slender, femora and tibiae subparallel,
densely covered with small curved setae holding
a layer of soil, apical spurs very small.

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MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. \. Aoupinia pseudohelea, 6.

Tarsomeres not expanded, obconic or monili-
form, densely and finely setose, penultimate
tarsomere with outer face moderately oblique and
only slightly prolonged into a small rounded
lobe. Basal metatarsomere subequal in length to
apical one.

Abdomen. Intercoxal process of first ventrite
broadly rounded (Fig. 2). Tergite 7 without
stridulatory files or thickened basal portion.
Segment-7 glands absent. Very long extrusible
glands emerging between segments 8 and 9.

Ovipositor (Fig. 3C) with elongated and fused
coxites 3 and 4, gonostyles terminal. Vagina
without sclerites, with diverticulum (Fig. 3C) a
cluster of a few spermathecal tubules (Fig. 3C) at
base of accessory gland (ag).

Aoupinia pseudohclea sp. nov.

(Figs 1-3)

MATERIAL. HOLOTYPE: Museum National d'Histoire
Naturelle, Paris: <3, New Caledonia, Aoupinie top camp,
23 November 2001, GB. Monteith, QM Berlesatc 1045,

NEW GENUS OF ADELIINI

443

FIG. 2. Aoupinia pseudohelea , outline of underside, c,
collar of prostemum. Scale = 2mm.

21 0 11 ? S I65°18’E, Rainforest 850m, Sieved litter.
PARATY PES: All in Queensland Museum: 1 9 : Same
data as holotype (QMT93428); 1 6 : Aoupinie, 20 Nov
2000, GB. Monteith, QM Berlesate 1035, 21°H’S
165°19'E, Rainforest, 850m, Sieved litter (QMT93427); 1
9 : Aoupinie top camp, 2-3 Nov 2001 , GB. Monteith, QM
Berlesate 1060, 21°H*S 165°18'E, Rainforest, 850m,
Sieved litter (QMT93426).

DESCRIPTION. Total length 9.8-1 1.0mm,
maximum width across elytra 5. 0-5. 7mm.

Head. As for genus.

Prothorax. Disc of pronotum smooth and
somewhat uneven, a little depressed in middle,
with trace of median impressed line at base only,
anterior edge concave, posterior edge straight in
middle, each side expanded into a basally very
wide upcurved flange with scalloped edges
which narrows anteriorly to produce an extension
arching forward over head, not quite meeting
extension of other side (Fig. 1). Edge provided
with a fringe of numerous closely-spaced short,
curved and thickened setae. Hind edge of flange
deeply arcuately excised outwardly with
produced hind angle, inwardly with a strong

FIG 3. Aoupinia pseudohelea. A, left antenna from
below; B, aedeagus in ventral view; C, ovipositor and
female tract in ventral view, a, fused alae of aedeagus,
ag, spermathecal accessory gland; c3+4, fused
coxites 3 and 4; d, diverticulum; o, common oviduct;
s, spermathecal tubules. Scales = 1 mm.

projection directed backward toward base of
elytron, upper surface of flange feebly rugose and
punctate.

Pterothorax. Discal surface feebly convex, inter-
vals flat, striae consisting of closely spaced
moderately impressed punctures, 8th stria at base
of flange with larger and deeper punctures.
Flanges" wide, about 1/3 as wide as elytral disc,
their edges even and provided with serried row of
hooked setae, dorsally with numerous large
shallow punctures not aligned into striae.

Legs. As for genus.

444

MEMOIRS OF THE QUEENSLAND MUSEUM

Majority rule

100

52

76

Isopteron A
Adelodemus A
Bellendenum A
Pseudobyrsax NC
Ozotypoides NC
Monteithium A
AoupiniaNC
Adelozotypus NC
Montaguea NC
Bolusculus A
Neoadelium NC

FIG. 4. 50% majority rule consensus of 33 trees tor ten
related squalid genera of Adeliini with Neoadelium as
the outgroup, using PAUP 3.1. All characters
unordered and unweighted. Tree length 54, Cl 0.533.
A = Australia; NC = New Caledonia.

Abdomen. Aedeagus (Fig. 3B) with sides of basal
piece inflected but not quite meeting, alae (Fig. 3B)
membranous, fused, no basal flange. Otherwise
as for genus.

HABITAT. Aoupinia was extracted by Bcrlese
funnel from sieved leaf litter collected in
undisturbed rainforest at 850m altitude at the
point on the summit of the range where the east
and west access roads meet. It is curious that no
other rainforest-inhabiting Adeliini were found
in this area, apart from a specimen of an
undescribed species of Ozotypoides Kaszab
obtained by pyrethrum fogging of a tree trunk
(G.B. Monteith, pers. comm.). Like all the squalid
Adeliini Aoupinia is covered in life with a layer of
fine clay which presumably serves to make the
beetle inconspicuous, and which in this case
bridges the gap between the two wings of the
pronotal flanges.

RELATIONSHIPS OF AOUPINIA

In spite of its close resemblance to Helea ,
which belongs to the Tenebrioninae, Aoupinia is
shown to be in the Adeliini ( Lagriinae) by the pair
of long eversible defensive gland sacs emerging
between segments 8 and 9 of the abdomen. All
other Tenebrionidae with defensive glands have
them between segments 7 and 8 (and much
shorter). Also, the ovipositor and female tract
configuration (Fig. 3C), with slender coxites,
terminal gonostyles, and multiple spermathecal
tubules, is typical of Adeliini and unknown in
Tenebrioninae.

Within Adeliini Aoupinia is strikingly different
in appearance from other genera because of its

TABLE 1 . List of characters and their states.

1. Clypeus anterior edge: 0, straight; 1, cmarginate.

2. Clypeo-frontal suture: 0, shallow; 1, deep.

3. Frontal grooves: 0. absent; l, present.

4. Groove around eye: 0, absent; 1, present.

5. Eye shape: 0, transverse; 1, round.

6. Antennomere 7: 0, subparallel, a little longer than wide; 1,
subparallel, 1.5- 1.7 X as long as wide; 2, eupuliform.

7. Tomentose (‘club’ ) segments of antennae: 0, none; 1 , three;
2, four.

8. Pronotal lateral outgrowths: 0, absent; 1, present.

9. Interlocking of fore and hind body: 0, absent; 1, present.

10. Prostemal collar: 0, absent; 1, present.

11. Prosternal process shape: 0, simple; 1, nodose; 2,
bifurcate.

12. Swollen metastemum: 0, absent; 1, present.

13. Elytral flange or carina along stria 8:0, absent; 1, present.

14. Elytral intervals: 0, smooth; 1, tuberculatc.

15. Epipleuron width: 0, widening anteriorly; 1, narrow
throughout or not evident.

16. Penultimate tarsomere: 0, feebly or not lobed; 1, strongly
lobed, symmetrical; 2, strongly lobed, asymmetrical.

1 7. Basal metatarsomere length: 0, shorter than or subcqual to
claw segment; 1 , longer than claw segment.

18. Intcrcoxal process of first abdominal ventrite: 0, arcuate;
1 , truncate.

19. Stridulatory files of tergitc 7: 0, present; 1, absent.

20. Vaginal sclerites: 0, present; 1, absent.

2 1 . Sides of aedeagal basal piece: 0, not meeting; 1 , meeting.

22. Alae of aedeagus: 0, present, separate; 1 , present, fused; 2,
absent.

wide pronotal and elytral flanges with the former
nearly meeting above the head, and its relatively
large size. Nevertheless it is clear that it belongs
to the group of northern rain forest genera
informally called the Pseudobyrsax group by
Matthews (1998), which includes all the New
Caledonian squalid genera plus Monteithium
Matthews of the Queensland wet tropics. This
group is characterised by a squalid
integumentary surface (one which holds a layer
of soil) which is also tuberculate in all members
except Aoupinia , antennae with the apical three
or four segments contrastingly tomentose (Fig.
3 A ), anterior edge of the prostemum often raised
into a collar (Fig. 2), epipleuron narrow
throughout, presence of a diverticulum on the
vagina (Fig. 3C), and absence of sexual
dimorphism. It is a characteristic of some

NEW GENUS OF ADELIINI

445

TABLE 2. Character state distribution for the northern
squalid group with Neoadelium as outgroup. A =
Australia, NC = New Caledonia.

II

1 2

12345 67890 12345 67890 12

Neoadelium NC

00100 00000 00001 10011 12

Isopteron A

11010 00000 20000 00100 12

Bolusculus A

00100 00100 00111 00111 12

Adelodemus A

11100 00101 2101011001 02

Pseudobyrsax NC

01010 02101 20011 10111 02

Adelozotypus NC

1

1111001011 00011 00111 12

Montaguea NC

0111001011 20011 00111 12

Ozotypoides NC

01010 12000 20011 10111 01

Bellendenum A

00100 00100 11010 21010 00

Monteithium A

00101 22101 21011 10110 01

Aoupinia NC

OHIO 12111 00111 00011 01

members of the group to interlock the fore and
hind bodies by means of an interdigitation of
integumentary projections and cavities, a feature
which is maximally developed in Montaguea
Kaszab. In Aoupinia there are similarly two
backward inner projections on the prothorax
(Fig. 1) which can be braced against the
depressed part of the elytral bases.

In a previous analysis (Matthews, 1 998) it was
found that the intuitive Pseudobyrsax group is
not clearly differentiated from the similarly
squalid Isopteron group of four Australian
genera, although the two together appear to be
monophyletic and may be termed ‘the northern
squalid group' (there is an unrelated southern
squalid group involving taxa from New Zealand,
Chile and southeastern Australia). The northern
squalid group is a syntaxon of five New Cal-
edonian and five Australian genera, which is
confined to the rainforests of northeastern
Queensland and New Caledonia (with the single
exception of the much more widespread
Australian xerophilic Isopteron Hope). In New
Caledonia the squalid genera have a very
polarised distribution: of the five genera present,
all occur in the northern half of the island while
only one ( Pseudobyrsax Kaszab) occurs in the
southern half.

Here the 10 genera of the northern squalid
group are briefly analysed again using PAUP3. 1 ,
with Aoupinia and Ozotypoides Kaszab added to

the previously included eight in Matthews ( 1 998),
and some additional characters introduced. The
outgroup is the non-squalid, or carabiform, New
Caledonian Neoadelium Carter which appears as
a sister group in Matthews ( 1 998, fig. 1 75). Table
1 lists the 22 characters involved and their states,
and Table 2 shows the taxon character matrix.
Autapomorphies (character states 5.1, 6.2, and
16.2) are included in the matrix even though
uninformative in order to present a complete
picture of character state distribution.

Intuitively it appears that Aoupinia is related to
the Australian Monteithium , as suggested by the
sharing of elaborate pronotal outgrowths and four
tomentose antennal club segments, but in fact
there are more numerous differences between the
two as may be seen in the table. The resultant
analysis (Fig. 4) leaves both these genera
ungrouped either with each other or with any
other single genus, while most of the remainder
form a series of terminal pairs which are either
wholly Australian or wholly New Caledonian. In
other words, there is no suggestion of a closer
relationship between any given Australian and
New Caledonian taxa beyond that of their
common membership in the northern squalid
group as a whole.

ACKNOWLEDGEMENTS

I am grateful to Geoff Monteith of the Queens-
land Museum for giving me the opportunity to
describe this extraordinary insect, latest in a long
series of his discoveries in tropical rainforest. I
also wish to thank Geoff Thompson of the
Queensland Museum for the fine illustration of
the whole beetle (Fig. 1), and the curators of the
following collections for the loan of comparative
material of New Caledonian Adeliini: Hungarian
Natural History Museum (O. Merkl), New
Zealand Arthropod Collection (R. A. Leschen),
and Queensland Museum (G.B. Monteith).

LITERATURE CITED

KASZAB, Z. 1982. Die Tenebrioniden Neukaledonien
und der Loyaute-Inseln. Folia Entomologica
Hungarica 43: 1-294.

1 986. Tenebrioniden (Coleoptera) aus Neukaledonien.
Annales Historico-Naturales Musei Nationalis
Hungarici 78: 151-175.

MATTHEWS, E.G 1998. Classification, phylogeny and
biogcography of the genera of Adeliini
(Coleoptera: Tenebrionidae). Invertebrate
Taxonomy 12: 685-824.

ANEW CAVERNICOLOUS SHRIMP IN PYCNISIA BRUCE, 1992 (CRUSTACEA:
DECAPODA: CARIDEA: ATYIDAE) FROM NORTHWESTERN QUEENSLAND

HIROSHI SUZUKI AND PETER J.F. DAVIE

Suzuki, H. & Davie, P.J.F. 2003 06 30: A new cavemicolous shrimp in Pycnisia Bruce, 1 992
(Crustacea: Dccapoda: Caridea: Atyidae) from northwestern Queensland. Memoirs of the
Queensland Museum 49(1): 447-451. Brisbane. ISSN 0079-8835.

A new cavemicolous shrimp, Pycnisia hunyip , is described from Forbes Inferno Cave,
Riversleigh, Lawn Hill National Park, northwestern Queensland. Pycnisia is differentiated
from all other atyid genera by its robust pereiopods 3-4. Pycnisia hunyip is distinguished
from the only known congener, Pycnisia raptor Bruce, by the shape of the rostrum and
telson. This is the seventh species of troglophilic atyid from Australia. □ Pycnisia, Atyidae,
Caridea, Queensland, new species, troglobite.

Hiroshi Suzuki, Aquatic Resource Science, Faculty of Fisheries, Kagoshima University,
4-50-20 Shimoarata, Kagoshima 890-0056, Japan ( e-mail : suzuki@
fish.kagoshima-u.ac.jp) ; Peter J.F. Davie, Queensland Centre for Biodiversity’, Queensland
Museum, PO Box 3300, South Brisbane 4101, Australia; 15 April 2003.

The first Australian subterranean shrimp was
described in 1960, and another five species have
been described in indigenous Sty’giocatis Holthuis,
1960, Parisia Holthuis, 1956, Pycneus Holthuis,
1986, and Pycnisia Bruce, 1992. All have so far
been found only in wells or caves in northwestern
Australia. These troglophilic shrimps have
reduced eyes without pigments, and no carapace
spines (except for an antennal spine in
Stygiocaris). The previously monotypic Pycnisia
was established for P raptor because of its
unique remarkably robust pereiopods. Pycnisia
hunyip sp. nov. is the seventh species of
troglophilic atyid from Australia.

Reproductive isolation leading to incipient
speciation can be expected to occur amongst
freshwater crustaceans occurring in widely
isolated freshwater cave systems. During surveys
of the Forbes Inferno Cave, Riversleigh, Lawn Hill
National Park, northwestern Queensland,
conducted in 1993 and 1994, unusual specimens
of a Pycnisia species were found. The shape of
the rostrum, and structure of the telson of these
shrimps indicate that they represent a new
species that is herein described and illustrated.

The types are deposited in the Queensland
Museum, Brisbane (QM). Measurements are of
post-orbital carapace length in millimeters (mm).

SYSTEMATICS

Family AYIDAE De Haan, 1849
Genus Pycnisia Bruce, 1992
Pvcnisia bunvip sp. nov.

(Figs 1-4)

MATERIAL. Holotype: QMW26722, 9 (7.0mm), Forbes
Inferno Cave, Riversleigh, Lawn Hill National Park,
northwestern Queensland, in shallow subterranean pools,
S. Williams and B. Ehrlich, 22/6/1994. Paratypes:
QMW26723, 9 (2.6mm), data as for holotype.
QM W26724, 9 (specimen damaged), data as for holotype.
QMW26725, 9 (6.2mm,), same location, coll. S.
Williams, 14/6/1993. QMW26725, 9 (3.7mm,), data as
for preceding specimen.

ETYMOLOGY. A mythical predatory creature said to live
in Australian freshwater pools. A noun in apposition.

DIAGNOSIS. Rostrum small, short, reaching to
only half length of eyes, but projecting beyond
level of inferior orbital angle. Eyes reduced,
pyriform, lacking pigment. Telson with one pair
of dorsal spines (2 pairs in paratypes) and one pair
of sub-dorsal spines, 7 spines (6-8 in paratypes)
on posterior margin. Pereiopods 1-5 with l, 1, 1, 1
and 2 setobranchs, respectively.

DESCRIPTION. Small species (postorbital
carapace length <7.0mm), but largest recorded
Australian troglophilic atyid species. Carapace
thin, flexible, smooth, without spines or setae
(Fig. 1); rostrum small, acute, with short, faint,
dorsal ridge, reaching beyond level ot inferior
orbital angles to half length of eyes (Figs 1 , 2A-B,
3M); pteryogstomian angle broadly rounded (Fig.
2B).

448

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 1. Pycnisia bunyip, sp. nov., 9 holotype (QMW26722), lateral
view. Scale bar = 5mm.

Abdomen smooth, pleura of abdominal
somites 1-5 rounded (Fig. 1). Telson slightly
shorter than sixth abdominal somite, rectangular,
with 1 pair of dorsal spines (2 pairs or 4
irregularly arranged dorsal telsonic spines on
paratypes) and 1 pair of small subdorsal spines
near posterolateral comer (Figs 2C, 3N,0);
dorsal spines placed slightly behind middle of
telson, second pair, if present, placed close to
posterolateral margin (Fig. 3N,0); posterior
margin broad, faintly biconvex, with 7 plumose
and 5 simple setae (6-8 plumose setae in
paratypes).

Eyes reduced, pyriform, devoid of pigment,
cornea rounded (faint papilla on tip of eyes in
small specimen with postorbital carapace length
of 2.6 mm) (Figs 1, 2A-B. 3M).

Antennular peduncle stout, slightly exceeding
distal margin of scaphocerite (Figs 2 A, 3A,M);
proximal segment broad, with stylocerite acute,
laterally divergent, reaching beyond middle of
proximal segment, outer anterolateral angle
weakly forwardly produced (acutely produced in
young paratype), medial margin relatively
straight; intermediate and distal segments sub-
cylindrical, medial margins setose; antennular
flagella slender, subequal.

Antennal scaphocerite (Figs 2 A, 3C,M) well
developed, broad; outer margin with large broad
acute distolateral tooth; inner margin convex;
bluntly rounded distally; antennal peduncle (Fig.
3B) without spines; flagellum long (Fig. 1),
distinctly longer than whole body length.

Mouthparts similar to P. raptor , Mandible
without palp (Fig. 3D-E); molar process small,
occlusal surface with marginal setal fringe;

incisor process short, broad, with 4
stout marginal teeth. Maxillula with
slender palp, with several setae
distally (Fig. 3F); upper lacinia
narrow, with numerous short
spinules on inner and distal margins;
lower lacinia subrectangular with
short setae on inner margin. Maxilla
(Fig. 3G) with slender palp con-
cealed behind endites, basal endite,
coxal endite, and scaphognathite
having 20 multidenticulate long setae
on posterior end. First maxilliped
(Fig. 3H) with simple tapered palp,
not reaching to distal margin of basal
endite; basal endite lamellar,
elongate, medial margin densely
fringed with setae; coxal endite stout;
exopod well developed, caridean lobe large,
broad, tapering distally; epipod vestigial. Second
maxilliped (Fig. 31) with well-developed exopod,
with numerous long plumose setae on distal half,
and shorter, thicker setae placed about quarter
distance from base; small rounded epipod with
multilamellar podobranch. Third maxilliped
reaching beyond antennular peduncle, stout with
small single arthrobranch (Figs 1, 3J); exopod
well developed, with long setae distally; epipod
strap shaped.

Pleurobranchs present on pereiopods 1-5,
exopods on maxillipeds only, and epipods on all
maxillipeds and pereiopods 1-4.

First pereiopod short, robust, chelate (Fig. 3K);
chela about 1.5 x carpus length, palm of chela
swollen, posteriorly expanded, longer than
dactylus; fingers robust, tapering, curved,
distally with tuft of sparse setae, claw-like tip;
carpus expanded distally, distodorsal margin
deeply excavate; merus subequal to chela, about
1 .5 x carpus length, slightly broadened distally;
coxa stout, with strap-like distally hooked epipod
and one setobranch. Second pereiopod (Fig. 3L)
with chela, chela about 0.9 x carpus length;
palm feebly expanded proximally, faintly bowed;
dactylus as long as palm length, unarmed,
otherwise similar to first pereiopod; fingers
subequal, stout, pigmented distally, fixed finger
similar to dactylus; carpus feebly expanded
distally, distodorsally feebly excavate; merus
about 1 .5 x carpus length; coxa similar to that of
first pereiopod, with strap-like epipod and one
setobranch. Third pereiopod very robust (Figs
2D, 4A); dactylus stout, tapering distally, unguis
demarcated with fringe of setae, ventral margin
with 9 stout, moveable spinules (Fig. 2E);

A NEW SPECIES OF PYCNISIA

449

FIG. 2. Pycnisia bunyip , sp. nov., $ holotypc (QMW26722): A, dorso-frontal view of carapace; B, latero- frontal
view of carapace; C, dorsal view of telson; D, left third pereiopod; E, dactylus of left third pereiopod; F, left
fourth pereiopod. Scale bars = 1 mm.

propodus about 0.9 x (range 0.8- 1.1 in paratypes)
carpus length, slightly bowed, ventral margin
with numerous short, stout, moveable spinules;
carpus slightly convex ventrally, with single row
of setae medially; merus very robust, about 1 .9 X
(range 1 .9-2.7 in paratypes) carpus length, oval in
section, 2.8 x (range 3. 8-5.0 in paratypes) longer
than greatest width, dorsal and ventral margins
convex; ischium short, distal half of ventral
border with rounded patch of dense setae; basis
short, robust; coxa robust, with strap-like distally
hooked epipod and one setobranch. Fourth
pereiopod (Figs 2F, 3B) robust as third
pereiopod; propodus subequal to carpus (range

1.0- 1.3 in paratypes); merus about 1.9 x (range

2.0- 2.9 in paratypes) carpus length, 2.9 x (range
3. 3-7.0 in paratypes) longer than greatest width;
otherwise similar to third pereiopod. Fifth
pereiopod slender (Fig. 4C); dactylus slender,
feebly tapering distally, unguis faintly
demarcated; propodus about 1.7 X (1.6 in
paratype QMW26725) carpus length,
ventrolateral border with row of stout setae;
carpus with small distodorsal lobe; merus about

1.5 x (1.7 in paratype QMW26725) carpus
length; ischium and basis normal; coxa with two
setobranchs but without epipod.

First pleopod (Fig. 4D) with slender, distally
attenuated endopod. Second (Fig. 4E) to fifth
pleopods similar, endopod not attenuated, with
appendix interna. Uropodal protopod with
posterolateral angle acute (Fig. 4F); diaeresis ot
exopod distinct, but incomplete; endopod
slightly shorter than exopod.

Color In Life. Photographs show a yellow
carapace with colour extending into anterior
abdomen, probably representing the hepato-
pancreas, and possibly ovary, beneath. Abdomen
mostly transparent.

REMARKS. This new species, P. bunyip , is
similar to P. raptor in having reduced eyes
without pigmentation, a small rostrum, and
robust pereiopods. However, in P. bunyip the
rostrum reaches slightly past the level of orbital
angle (shorter in R raptor ), and the telson has
only one or two pairs of dorsal spines and one pair

450

MEMOIRS OF THE QUEENSLAND MUSEUM

antenna; C, scaphocerite of left antenna; D, ventral view of mandible; E, dorsal view of mandible; F, left
maxillule; G, left maxilla; H, First maxilliped; I, second maxilliped; J. third maxilliped; K, First pereiopod; L,
second pereiopod. M-N, 9 paratype (QMW26723); M, dorso-frontal view of carapace; N, dorsal view of telson!
O, 9 paratype (QMW26724), dorsal view of telson. Scale bars = 1mm.

of subdorsal spines (four pairs of dorsal spines
and one pair of subdorsal spines in P. raptor).

The setobranch and multidenticulate scapho-
gnathite setae are known to have an important
gill-cleaning role in some caridean shrimps
(Bauer, 1981, 1989; Suzuki & McLay, 1998).
Pycnisia bunyip has one or two setobranchs on
the coxae of the pereiopods, and 20 multi-
denticulate scaphognathite setae similar to other
atyid species. Four Australian troglophilic
shrimps, Parisia gracilis , P. unguis , Pycneus
morsitans and Pycnisia raptor , also have
setobranch and several long multidenticulate
setae clearly belonging to the scaphognathite

(Bruce, 1992; Holthuis, 1986; Williams, 1964).
These species, along with P. bunyip , are thought to
have passive gill-cleaning mechanisms as occur in
other atyid species. However, Stygiocaris
lancifera and S. stylifera have no long setae on
the posterior portion of the scaphognathite
(Holthuis, 1960). These two species are thought
to have either an active gill-cleaning mechanism
using a pereiopod setal brush, or a different form
of passive gill-cleaning using branchiostegal
setae (Bauer. 1998; Batang & Suzuki, 2000).
Future study of gill-cleaning mechanisms of
cavernicolous shrimps may be useful for

A NEW SPECIES OF PYCNISIA

451

FIG. 4. Pycnisia bunyip , sp. nov., $ paratype (QMW26725): A, third pereiopod; B, fourth perciopod; C, fifth
pereiopod; D, first plcopod; E, second pleopod. Scale bars = 1mm.

understanding adaptive selection pressures and
phylogenetic relationships.

DISTRIBUTION. Only known from the type
locality.

ACKNOWLEDGEMENTS

We are very grateful to Stefan Williams for the
collection and donation of the specimens, and the
Japanese Government for a travel grant to H.
Suzuki. A.J. Bruce and Xinzheng Li are thanked
for their useful comments on the manuscript.

LITERATURE CITED

BATANG, Z.B. & SUZUKI, H. 2000. Gill structure and
gill-cleaning mechanisms of the redclaw crayfish
Cherax quadricarinatus (Decapoda, Astacidea,
Parastacidae). Journal of Crustacean Biology
20(4): 699-714.

BAUER, R.T. 1981. Grooming behaviour and morph-
ology in the decapod Crustacea. Journal of
Crustacean Biology 1: 153-173.

1989. Decapod crustacean grooming: functional
morphology, adaptive value, and phylogenetic
significance. Pp. 49-73. In Felgenhaucr, B.E. et
al. (eds) Functional morphology of feeding and

grooming in Crustacea. Crustacean Issues 6.
(A.A. Balkema: Rotterdam).

1998. Gill-cleaning mechanisms of the crayfish
Procambarus clarkii (Astacidea: Cambaridae):
experimental testing of setobranch function.
Invertebrate Biology 117: 129-143.

BRUCE, AJ. 1992. Pycnisia raptor , anew genus and
species of predatory troglobic shrimp (Crustacea:
Decapoda: Atyidae) from Northern Australia.
Invertebrate Taxonomy 6: 553-566.

HOLTHUIS, L.B. 1960. Two new species of atyid
shrimps from subterranean waters of N. W.
Australia (Decapoda Natantia). Crustaceana 1:

47-51. r t

1986. A new genus and species of subterranean
shrimp from Western Australia (Crustacea:
Decapoda: Atyidae). Zoologische Medcdelingen,
Leiden 60(7): 103-1 1 1.

SUZUKI, H. & McLAY, C.L. 1998. Gill-cleaning
mechanisms of Paratya curvirostris (Caridea:
Atyidae) and comparisons with seven species of
Japanese atyid shrimps. Journal of Crustacean
Biology 18(2): 253-270.

WILLIAMS, W.D. 1964. Subterranean freshwater
prawns (Crustacea: Decapoda: Atyidae) in
Australia. Australian Journal of Marine and
Freshwater Research 15: 93-106.

452

MEMOIRS OF THE QUEENSLAND MUSEUM

A NOTE ON THE HABITAT REQUIREMENTS OF
THE SWAMP CRAYFISH ON BRIBIE ISLAND,
SOUTHEASTERN QUEENSLAND Memoirs of the
Queensland Museum 49(1): 452, 2003 > Tenuibranchiurus
glypticus (Riek, 1969) is a little studied freshwater crayfish
with a distribution restricted to acidic wallum (sedge-heath)
swamps along the southeastern coast of Queensland. This
report describes the physical conditions and habitats in which
T. gypticus are found.

Observations were made at 38 sites on Bribie Island in
southeastern Queensland from February to March of 2001.
Bribie Island is a low-lying sand island with many areas
dominated by wallum heath vegetation and characterised by
pools of tannin-stained water. Sampling sites were swamps
and small artificial dams, and pools and gutters occurring
beside and across sandy vehicular tracks. Sampling was
conducted with dip nets using standard sweeps and all
animals captured were identified, enumerated and released at
point of capture. A range of physical parameters were
recorded (pH, substrate type, turbidity and presence of
vegetation) in order to ascertain whether T glypticus was
associated w ith any pool characteristics.

T glypticus was caught at 20 of the 38 sites sampled ( 1 9 of
34 temporary pools and 1 of 4 permanent pools). Most pools
were tannin-stained (n = 35) and pH ranged from 2.6-5.65. T.
glypticus were found in pools with pH ranging from 3.2-4.80,
and were not found in any of the three pools with clear water.
T. glypticus were more likely to be found in pools w'ith a sandy
substrate(62% of 29 pools) than in pools with a thick layer of
organic matter (22% of 9 pools; Fishers Exact test, p=0.058).

More pools with vegetation in the middle contained T.
glypticus (85% of 13 pools) than pools without vegetation in
the middle (36% of 25 pools; Fishers Exact test = 6.28,
p=0.012). T glypticus were more common in pools with the
sedge Rastio pollens (80% of 24 pools with R. pollens v 44%
of 14 pools without; Fishers Exact test, p=0.035). T. glypticus
presence was positively associated with the presence of insect
predators (Anisopteran odonate nymphs and/or Nepid

hemipterans) (100% of 11 pools with insects v 33% of 27
pools without; Fishers Exact test - 1 1.4, p<0.001), and the
crayfish Cherax robustus (76% of 1 7 pools w'ith C. robustus v
33% of 21 pools without; Fishers Exact test = 5.39, p=0.020),
but these associations were most likely due to common
habitat requirements of these species. The only fish species
found in samples was the Striped gudgeon ( Gobiomorphus
australis ) (6 pools, 2 with T. glypticus).

In 7 of the 20 pools in which T. glypticus were found, catch
rates (number/standard sweep) were compared between
sweeps through sedges (24 sweeps) and sweeps over bare
substrate (50 sweeps). Catch rate w-as significantly higher in
sedges (mean saiv . 1 . 12,s.d- 0.39, n =7; mean non sait , c = 0.00,
s.d=0, n=7; Wilcoxon z - -2.46, p=0.014).

Of the 20 sites where T. glypticus was collected, all were
w ithin wallum heath areas containing tannin-stained pools. T.
glypticus was only captured within vertical standing sedges.
This may indicate that sedges provide protection from
predators, or provides a food requirement. The data suggest
that T. glypticus has specific habitat requirements w'ilhin the
wallum areas in which it is found. These habitat
characteristics need further investigation in order to
understand the ecology of this unique crustacean.

Acknowledgements

Animals were collected under Queensland Parks and
Wildlife Service Permit E6/0000/0 1 /S A A and Queensland
Fisheries Permit PRM0049I J. We are grateful to the people
that helped in the collection of field data, and assisted in plant
and animal identification and finding sampling sites.

Literature Cited

RIEK. E.F. 1969. The Australian Freshwater Crayfish (Crustacea:

Decapoda: Parastacidae) with descriptions of new species.

Australian Journal of Zoology 17: 101-106.

Douglas Harding & Ian Williamson, School of Natural
Resource Sciences, Queensland University of Technology,
Gardens Point 4001, Australia.

NESTING STRATEGIES OF THE WATER MOUSE XEROMYS MYOIDES IN
SOUTHEAST QUEENSLAND

STEVE VAN DYCK AND IAN GYNTHER

Van Dyck, S. & Gynther, I. 2003 06 30: Nesting strategies of the Water Mouse Xeromys
myoides in southeast Queensland. Memoirs of the Queensland Museum 49(1): 453-479.
Brisbane. ISSN 0079-8835.

Studies of Xeromys myoides nests on three islands of Moreton Bay and at ten coastal sites on
mainland southeast Queensland have revealed a variety of nesting strategies ranging from
the construction of large, free-standing, termitarium-like mounds up to 66cm high, to the
excavation of inconspicuous tunnels in the supralittoral bank at the marine/terrestrial
boundary. Techniques employed to locate nests and useful features for confirming the
identification ofX. myoides nesting structures arc provided. Information from a total of 1 10
nests was compiled. Of these, 21 were free-standing structures within areas of sedgeland,
chenopod shrubland, Sporobolus virginicus grassland or mangroves. Others were associated
with small, slightly elevated ‘islands' standing away from the supralittoral bank (20 nests) or
with the supralittoral bank itself (20). Thirty-one examples of nests constructed in living or
dead trees situated in the intertidal zone (or at its landward edge) were documented. Another
eighteen nests were recorded in spoil heaps of human origin. Information about the height of
nest structures and the number of holes providing access to nests is supplied. Where mound
structures were present, their height was built up over time with repeated plastering of
‘mortar' brought from within or below the nest and smeared from one or more entry holes to
the mound top in clearly defined tracks. Well-established mounds were rarely inundated
entirely. Nest location and. therefore, nest type were interpreted as resultant compromises
between the ability to withstand spring tides versus proximity to the most highly productive
resources of the mangrove zone. □ Xeromys, False Water-rat, rodents, survey , southeast
Queensland.

Steve Van Dyck, Queensland Museum, PO Box 3300 , South Brisbane 4101,
(stevevd@qm.qld.gov.au); Ian Gynther, Conservation Service, Queensland Parks and
Wildlife Service, Southern Region, POBox64, Bellbo\vrie4070, Australia; 12 April 2002.

Magnusson et al. (1976) described an extra-
ordinary, 60cm-high mud structure resembling a
termite mound, built at ground level against the
trunk of a living Bruguiera parviflora in a
mangrove forest on Melville Island, Northern
Territory. From this structure they extracted an
adult female Xeromys myoides and two young,
thereby documenting the first record of a nest for
this poorly known species. In 1991 , a number of
sedge-covered peat mounds attributed to X.
myoides were found on North Stradbroke Island,
southeast Queensland (Van Dyck, 1992; Van
Dyck & Durbidge, 1 992). None of these occurred
in mangroves but rather in immediately adjacent
areas of sedgeland or on the more landward
supralittoral bank (Van Dyck, 1997). The present
investigation of X. myoides nesting elsewhere on
North Stradbroke Island, as well as on South
Stradbroke and Bribie Islands and at ten
mainland sites in coastal southeast Queensland,
has revealed a variety of nesting strategies for the
species. These ranged from the construction of
large free-standing termitarium-like mounds to
the exploitation of hollow trunks within (or at the

landward edge of) the tidal zone and the
excavation of inconspicuous tunnels in the
supralittoral bank at the marine/terrestrial
boundary. The information presented here has
been compiled from a total of 110 nests
documented by us since 1991. It presents a broad
range of nesting strategies hitherto unrecorded
for this threatened species.

METHODS

Nesting structures were documented as part of
an ongoing survey of X. myoides in southeastern
Queensland and northeastern New South Wales.
Nests were recorded at sites examined between
the Great Sandy Strait, Queensland (25°47’S,
1 52°58 , E) and the Richmond River area of New
South Wales (28°54’S, 153°3UE), 345km to the
south (Table 1, Fig. 1). Nests were generally
located by searching in the intertidal zone between
the supralittoral bank and the outer (frequently
seaward) edge of the mangroves. However, some
nests involving simple holes excavated in the
supralittoral bank or in spoil heaps were revealed
only during the radio-tracking of individuals that

454

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1 . Xeromys myoides nest localities, abundance and nest reference numbers in Queensland, and localities
searched (unsuccessfully) in New South Wales (ordered by increasing latitude).

Locality

Lat. (S)

Long. (E)

Total Nests

Ref. Nos.

Kauri Creek Conservation Park

25°47’

152°58’

6

1-6

Noosa North Shore

26°23’

153°04’

6

7-12

Maroochy River

26°38’

153°04’

1

13

Pumicestone Passage

26°59’

153°04*

8

14-21

Gallagher Point, Bribie I.

27°00’

153°06’

5

22-26

Bullock Creek Conservation Park

27°00’

153°04’

3

27-29

T3

c

Donnybrook

27°or

153°03’

17

30-46

1

White Patch, Bribie I.

27°or

153°07*

1

47

8

3

Amity, N. Stradbroke I.

27°25’

153°26*

4

48-51

O'

Rainbow Channel, N. Stradbroke I.

2T2T

153°25’

10

52-61

Canalpin Creek, N. Stradbroke I.

27°36’

153°24’

1

62

Stockyard, N. Stradbroke I.

27°43’

153°24’

1

63

Steiglitz

27°45’

1 53°20’

4

64-67

Jacobs Well

27°46’

153°21’

1

68

Pimpama River

27°48’

1 53°20’

1

69

Coomera River

27°50’

153°22’

22

70-91

South Stradbroke I.

27°5r

153°25’

19

92-110

Cobaki Broadwater (3 sites)

28°H’

153°30’

Terranora Creek A (2 sites)

28°H’

153°32’

-•

.

Ukcrebagh Island

28°ir

153°33’

-

.

Terranora Creek B (3 sites)

28°12’

153 0 31’

-

.

Ukerebagh Mainland

28°12’

153°33’

-

.

Fingal

28°12’

153°34’

-

_

v>

Terranora Broadwater (3 sites)

28°13’

153°30’

-

.

s

£

Banora Point (4 sites)

28°13’

153°33’

-

.

•5

Chinderah Bay (3 sites)

28°14’

153°33’

-

.

o

KA

Cudgen Creek, KingsclifT

28°17’

153°34’

-

.

£

b

2

Hastings Point (2 sites)

28°22’

153°34’

-

.

Brunswick River

28°32’

153°32’

-

.

Marshalls Creek, Brunswick Heads

28°32’

153°33’

-

.

Simpsons Creek, Brunswick Heads

28°33’

153°33*

-

.

Belongil Creek, Byron Bay

28°38’

153°35’

-

.

North Creek, Ballina

28°5r

153°34’

-

.

South Ballina

28°53*

153°33*

-

.

Hermans Wharf, Richmond River

28°54’

153°3r

-

-

used these tunnels (Van Dyck, 1997). A more
detailed description of search techniques is
provided below.

Up to four different vegetation communities
occurred in the intertidal search area and,
wherever possible, the location of each X.
myoides nest was recorded with respect to these
communities. Based on the definitions of
Clifford & Specht (1979), the communities
encountered were:

1) sedgeland — an often well-defined zone of
rushes and sedges growing to about lm and
typically including Juncus kraussii and Baumea
juncea. The Mangrove Fern Acrostichum
speciosum occasionally grows here.

2) chenopod shrubland — a less frequently
encountered low, open shrubland of succulents
with a dwarf shrub habit growing on soils that dry
out and crack between inundations. Plant species
typically include Enchylaena tomentosa ,

NEST TYPES OF XEROMYS MYOIDES

455

Inset 1

Pumicestone/Rassa'ge a

Bullock Creek CP,
Donnybroolc

2.5 5 Iqn

v Gallagher Point

White Patch*

Inset^3.

Cobaki Broadwater*/" ( (Jweed Heads
J^V t v£^UkeVebagh Island

wrannra.r.rAok. &x£\ &J \\

Fingal

Terra nora^C reek B Ukerebagh Mainland

3 y Sf

TerrangjaBroadwater ^j*J^ anora Point

Noosa North Shore\ A ^Noosa Heads
River*?

Maroochy River

Maroochydore

Caloundra

<8ri6i* Island

IN

A

Moreton Island

Amity a^

Rainbow Channel

^^^Canalpin Creek a / StradBrofy Island
/stockyard

Southport

Tweed Heads

Cudgen Creek
Hastings Point

Belongil Creek
Byron Bay

Ballina

North Creek
South Ballina
Hermans Wharf 8

i

FIG 1. Distribution of Xeromys myoides nesting localities in Queensland (A), and localities searched
(unsuccessfully) in New South Wales (V).

s.o.o.zz

456

MEMOIRS OF THE QUEENSLAND MUSEUM

Sarcocornia quinqueflora , Suaeda arbus-
culoides and Suaeda australis.

3) Sporobolus grassland — a salt meadow of
Marine Couch Sporobolus virginicus closed
grassland, usually found closest to the extreme
high water spring tide mark and associated with
freshwater drainage.

4) mangroves — a community of varying
structural type and complexity, but usually
comprising one or more of Avicennia marina var.
australasica, Rhizophora stylosa , Bruguiera
gymnorhiza , Aegiceras corniculatum and, less
commonly, Ceriops tagal. Dowling (1986) and
Van Dyck ( 1 997) provide additional details of the
many mangrove communities occurring in
Moreton Bay.

In situations where more than one of these
intertidal communities was present at a site,
distinct zonation was often apparent. This made
assignment of a X. myoides nest to a particular
community easy. At other times the boundaries
between the various communities were blurred or
the communities interdigitated such that clear
zonation of the different vegetation types was not
obvious. In these cases, a nest was associated
with the dominant vegetation community in its
proximity.

Each nest was assigned to one of five nest
categories (below) and its location determined
with a GPS navigator. The vegetation cover on
the nest, nature of the mound material, number
and position of entrance/exit holes and height and
circumference of the mound were recorded. The
degree of moating by high tides was also
assessed. Finally, the nest’s position in the
intertidal zone was put into perspective in
relation to the vegetation communities occurring
along a linear transect that started at the terrestrial
boundary and passed through the nest to
terminate at the closest deep channel or large
body of water out into or beyond the associated
mangroves.

SEARCH TECHNIQUES. Techniques employed
to locate nests of X. myoides are described. As
previously stated, manual searching was con-
ducted across the entire intertidal zone. Particular
attention was paid to areas of higher ground
abutting or lying within the various intertidal
vegetation communities, i.e. places that offer
some elevation and, therefore, refuge against the
high tide. Where a defined supralittoral bank
existed, this was searched thoroughly for mud
moundings or other signs of X. myoides. Other
areas of high ground that were potentially

suitable for nesting were detected by the different
nature of the vegetation they supported. Small
‘islands’ at the same elevation as the supralittoral
bank often existed in the landward sections of the
intertidal zone. These supported terrestrial trees
or shrubs such as Melaleuca quinquenervia ,
Casuarina glauca and Baccharis halimifolia ,
and were surrounded by Sporobolus grassland,
sedgeland or chenopod shrubland. Locations
seaward of the supralittoral bank where such
trees occurred were investigated closely.

Local topography at each site was also
carefully considered. At some localities, for
example, narrow tongues or even large islands of
coastal woodland lay partly or entirely encircled
by mangroves or other intertidal vegetation
types, offering many nesting opportunities for A".
myoides. These terrestrial isolates were located
by scanning across the canopy of the mangrove
community to detect the obvious crowns of
Casuarina glauca or other terrestrial tree species.
Routine study of colour aerial photography
(1:12,000 scale or better) of each survey site
ensured that the discovery of such areas of high
ground was not left to chance.

In addition to searching for these obvious
topographical features offering nesting potential,
subtler evidence was sought of raised areas within
the intertidal zone created directly by X myoides
activity or by human disturbance. Amidst
Sporobolus grassland or chenopod shrubland,
mounded nest structures constructed by Water
Mice or mounds of artificial origin (e.g. human
spoil piles) were usually obvious. Within taller
vegetation, such as sedgeland or stands of
Acrostichum speciosum , this was not always the
case. Nevertheless, because the tops of such
mounds are seldom, if ever, inundated by high
tides, they often bore a lush growth of Sporobolus
virginicus. Consequently, stands of Juncus
kraussii , Baumea juncea or A. speciosum were
scanned for these tell-tale clumps of S. virginicus.
Where these clues to possible nest structures
were lacking, extensive areas of J. kraussii , B.
juncea or A. speciosum were systematically
traversed using parallel transects to locate
otherwise concealed nest mounds. Minor contour
changes in the overall height of the sedge or fern
stands were closely investigated to determine
whether these were due to raised substrate or a
nest mound. Within the intertidal zone, bund
walls, piles of spoil material from earthworks and
bulldozed trees with associated root clods were
examined carefully for evidence of colonisation
by X. myoides.

NEST TYPES OF XEROMYS MYOIDES

457

Reward for search effort was greatly increased
if surveying for nests in dense intertidal vegetation
or lush ground cover on the supralittoral bank
was undertaken after recent fires had swept
through an area. At such times, signs of X.
myoides activity including access holes, mud
tracks and daubing (below) were more readily
observable. In some cases, nest structures were
revealed that had been overlooked during
previous surveys.

Nest searching within the mangrove zone was
conducted less methodically due to the often
extensive area needing to be covered. Dead trees
and stumps and hollow, living mangroves
encountered while conducting such searches or
while setting Elliott trap transects were inspected
for evidence of X. myoides nesting activity. Signs
of occupation sought included mounded mud
structures located at ground level within hollow
trunks, mud packing against the bases of trunks
or any mud or peat material in tree trunks and
limbs above ground level.

RESULTS

DISTRIBUTION OF NESTING RECORDS. A
total of 1 10 nests belonging to X. myoides was
discovered at 1 7 of 28 localities searched along
the coastline of southeastern Queensland (Fig. 1 ,
Table 1 ). These searched localities were scattered
from Kauri Creek, Great Sandy Strait (25°47'S,
152°58’E), south to Cumimbin Creek on the
Gold Coast (28°08’S, 1 53°28'E). No evidence of
X. myoides nesting activity was found at four
mainland sites south of the Coomera River in
Queensland or at any of the 3 1 sites (from 1 8
localities) surveyed in New South Wales (Fig. 1,
Table 1).

NESTING STRATEGIES. Nesting structures
of X. myoides encountered at sites surveyed in
southeast Queensland were categorised into one
of the following five broad types: 1) free-
standing nests, 2) island nests, 3) supralittoral
bank nests, 4) tree trunk nests, and 5) spoil heap
nests. Photographs (Figs 2-15) and relevant
details of nests from each class should aid
recognition of these structures by field workers.
Figure 16 illustrates the diversity of X. myoides
nest types and locations within the different
intertidal vegetation communities. Although
these five categories offer a useful scheme for
documenting the range of A', myoides nesting
structures, the classification proved to be
somewhat arbitrary with, in some situations, the
divisions between certain nest types being

unclear. For example, free-standing mounds built
against tree bases or against clods of soil between
the roots of upturned trees could be classified as
tree trunk nests or spoil heap nests, respectively
(below). Such difficulties, however, were the
exception rather than the rule.

/. Free-standing Nests. Free-standing nests were
solitary, termitarium-like mounds. They were not
associated with either the supralittoral bank,
areas of substrate elevated above their
surroundings (‘islands' and spoil piles) or (except
in rare instances) hollow tree trunks or stumps.
They occurred in: (i) the mangrove zone (Fig. 2);
(ii) sedgeland (Fig. 3); (iii) chenopod shrubland;
or (iv) Sporobolus grassland (Fig. 4).

The locations and physical features of free-
standing mounds documented during the study
appear in Table 2. Free-standing nests were
always more conspicuous than other nest types,
often being large constructions up to 66cm high
(mean = 42cm, SD = 12cm, n = 20; minimum
height of occupied nests 25cm). All experienced
360° moating at high tide. This nest type was
recorded mainly from areas of sedgeland and
Sporobolus grassland (18 out of 21 cases), with
only one example from chenopod shrubland and
two noted inside the mangrove zone. One of these
mangrove mounds (Stockyard #63) had been
abandoned at some point up to 3.5 years after it
was first discovered (below). The other (Pumice-
stone Passage #17) was situated in an area of
minimal tidal influence. Occupied nests were
thickly covered with Marine Couch ( 1 4 out of 20
cases), the sedges Juncus kraussii or Baumea
juncea (5 out of 20 nests) or a combination of
sedge and couch (one case). A smaller additional
component of cover was contributed in some
instances by Suaeda arbusculoides , S. australis ,
Fit ex bicolor or Acrostichum speciosum. When
first recorded, nest #63 at Stockyard, North
Stradbroke Island, was partially covered with S.
australis. However, when revisited 3.5 years
later, this vegetation had all died and the nest was
abandoned.

Free-standing nests occurred either in areas
receiving infrequent flooding by tides or areas
that experienced more regular inundation but
offered a high degree of protection from erosional
action (wind-induced waves and/or tidal
currents). This protection was due to the
buffering effect of an adjacent broad mangrove
zone or because the areas were situated along
calm waterways. The sheltered Marine Couch
and Sarcocornia quinquejlora flats of the
western shores of Pumicestone Passage,

458

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. A free-standing nest in the mangrove zone (nest #17, Pumiccstone
Passage, November 1996). Photo Ian Gynther.

intersected by extensive natural ponds and
shallow drainage ditches, and located far from the
deep water of the Passage itself, provided the
most numerous examples of this nest type. In
such areas, given the limited exchange of surface
waters, mangrove community composition was
limited to one species ( Avicennia marina) that
grew no taller than 5m.

Nests on the Noosa River (#s 9, 1 0) and Coomera
River (#s 76,77,87) were subject to more
extensive tidal inundation than those at Pumice-
stone Passage but occurred in similarly sheltered
areas amid broad expanses of Sporobolus
grassland or sedgeland and, in
these cases, adjacent to calm
river channels. The free-
standing nests at Kauri Creek
Conservation Park (#s 2,6),

Rainbow Channel (#57) and
South Stradbroke Island (#s
95,102,103) were all recorded
closer to potentially destructive
tidal influences, but occurred
on the landward side of 153-
400m-wide mangrove stands
that included Rhizophora
stylosa as a significant
component. The dense tangles
of prop roots typical of this
mangrove species would offer
an effective barrier against
strong tidal currents, storm
surge and wind-induced
waves. As would be expected

in such regularly inundated
sites, these nests were densely
consolidated by species with a
greater salt water tolerance,
namely sedges and Mangrove
Fern, and were closer to more
diverse mangrove communities.
The remaining nest, at Bullock
Creek Conservation Park
(#29), represented an inter-
mediate situation. Although
the surrounding sedgeland
here was not extensive, a
290m-broad mangrove zone
stood between it and the
relatively sheltered waters of
Pumicestone Passage.

Free-standing nests were
often constructed at great
distances from both the ter-
restrial woodland community
and deep water, further emphasizing the typically
sheltered nature of the locations at which these
nests occurred. For example, nests #17
(Pumicestone Passage) and #63 (Stockyard)
were 131m and 200m, respectively, from the
marine/terrestrial boundary, and many nests (#s
2,6,10,29,63, 95,102,103) were at least 250m
from the nearest body of deep salt water. Those at
Kauri Creek Conservation Park (#s 2,6) were
427m and 520m from the closest channel. With
one exception ( Rainbow Channel #57), all Type 1
nests were located adjacent to sections of the
shoreline that lacked a distinct supralittoral bank.

FIG. 3. A free-standing nest in sedgeland (nest # 1 0, Noosa North Shore, April
1 997). Photo Ian Gynther.

NEST TYPES OF XEROMYS MYOIDES

459

FIG 4. A free-standing nest in Sporobolus grassland (nest # 1 8, Pumicestone
Passage, November 1996). Photo Steve Van Dyck.

The greatest number of access holes (25) of any
nest type was recorded from a free-standing nest
mound (Pumicestone Passage #14),

2. Island Nests, island' nests were constructed
away from the supralittoral bank in areas of
substrate that were slightly higher than their
surroundings and generally above the level of
spring tides. They were often consolidated by the
roots of trees such as Melaleuca quinquenervia
and Casuarina glauca , or thickly covered with
sedges and/or Sporobolus virginicus. These
islands’ may represent vestiges of the supra-
littoral bank, eroded by the combined effects of
spring tides, wind-induced
waves and storm surge. Most
‘islands’ were, therefore,
closer to the supralittoral bank
than to the mangroves. Island
nests occurred in: (i) the
mangrove zone (Fig. 5); (ii)
sedgeland (uncommonly
including Acrostichum
speciosum) (Fig. 6); (iii)
chenopod shrubland; or (iv)

Sporobolus grassland. They
sometimes comprised simple
holes with no other signs of
working by X. myoides , but
more often were complex
constructions with additional
mounding.

Locations and physical
features of island nests are
shown in Table 3. Nests

constructed on islands were
second to free-standing nests
in their ease of detection. The
maximum recorded size of
such an island was approx-
imately ISm" (Donnybrook
#44). The mean height of
island nests above the
surrounding littoral substrate
was 51cm (range = 3 0-7 5 cm,
SD = 13cm. n = 20). All
islands were fully moated at
high tide and most (19 out of
20 examples) were consolidat-
ed by the roots of a few
salt-tolerant shrubs and trees
such as Casuarina glauca ,
Baccharis halimi folia and
Melaleuca quinquenervia or
the mangroves Avicennia
marina and Aegiceras
corniculatwn. The only island nest not associated
with shrubs or trees (Donnybrook #38) was
situated in the middle of an extensive area of low,
Sporobolus virginicus- covered plateaux,
intersected by a labyrinth of natural, shallow
channels and poorly draining pools.

All islands were thickly covered with ground
layer vegetation: Marine Couch (11 out ot 20
cases, including nest #38), sedges (five out of 20),
couch and sedges (two out of 20) or sedges and
Mangrove Fern (two out of 20). Marine Couch
cover generally characterised more sheltered
locations (e.g. Donnybrook; sections of Gallagher

FIG. 5. An island nest with obvious mounding in the mangrove zone (nest
#37, Donnybrook, November 1996). Photo Ian Gynther.

460

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 6. An island nest in sedgeland (nest #72, Coomera River, November
2001). Photo Ian Gynther.

Point on Bribie Island) whereas sedges or the
combination of sedges and fern, i.e. species
tolerant of a higher frequency of inundation by
salt water, occurred in areas potentially more
prone to erosion by spring tides, wind-induced
waves and storm surge (e.g. certain sections of
Kauri Creek Conservation Park, Amity, Rainbow
Channel and South Stradbroke Island). Nest #27
at Bullock Creek Conservation Park, situated in
an area exposed to only moderate erosional
forces, consisted of a Sporobolus- covered island
(with a single Casuarina glauca) within dense
sedgeland. Nest #74 at Coomera, another area of
intermediate shelter, was covered by a
combination of Marine Couch and sedges (with
an individual C. glauca). Couch and sedges also
covered one Donnybrook nest (#44), although
Marine Couch dominated, as was consistent with
the nest’s sheltered location.

Island nests were usually located closer to the
supralittoral bank or, where this was poorly
defined, the marine/terrestrial boundary (median
distance = 12m) than were free-standing nests
(median distance = 41m) or tree trunk nests
(median distance = 75m), most probably because
the island landlbrms bearing X myoides nests
originated through erosional processes operating
on the supralittoral bank. One exception, nest #37
at Donnybrook (Fig. 5), was located on an island
in an isolated, raised area of sparse Sporobolus a
distance of 195m into the mangroves from the
landward edge of the intertidal zone. Although
couch-covered, undermining of the structure by

spring tides was apparent at
around 20cm above the
substrate level.

The tops of all but one island
nest (Amity #50) were
plastered by X. myoides with
successive layers of mud or
peat daubing which, over time,
had produced mounds,
effectively increasing their
height against spring tides.

The greatest number of
access holes recorded from
island nests was seven (nest #s
37,48,110).

3. Supralittoral Bank Nests.
Supralittoral bank nests were
built into or on the earth bank
formed by erosional action at
the marine (mangrove, sedge-
land, chenopod, Sporobolus )/
terrestrial (swamp, wallum, coastal woodland)
ecotone by the highest of tides (Fig. 7). Such
nests were either: (i) simple holes excavated into
the vertical bank; or (ii) more elaborate
constructions with additional mounding (Fig. 8).

Twenty supralittoral bank nests were recorded.
The physical features of these are provided in
Table 4. Type 3 nests were more difficult to locate
than other types because banks were naturally
uneven in profile and thickly covered with
Marine Couch, sedges or shrubs, and because
mounding associated with such nests was either
nonexistent or occurred in various stages of
development. In the former case, inconspicuous
holes were built among peat and roots in the
bank. In the absence of peat or mud plastering
above these nests, the three recorded examples
(Rainbow Channel #s 54,55; Canalpin Creek
#62) were discovered only during the course of
radio-tracking studies.

In one case, at Donnybrook, a recent fire that
had burned to the supralittoral bank and into the
fringes of the Sporobolus grassland exposed
three nests that had not been detected during an
earlier survey (Fig. 17).

Supralittoral bank nests, being located at the
marine/terrestrial boundary, were not as prone to
inundation and so experienced less moating than
other nest types. The usual extent of moating of
nests in the supralittoral bank was 1 80°, although
the maximum recorded (270°) occurred in
situations where the bank formed small
promontories jutting out into the adjacent

NEST TYPES OF XEROMYS MYOIDES

461

TABLE 2. Free-standing nest mounds (Type 1) of Xeromys myoides from southeast Queensland. * The term
‘sedge’ refers to the combination of Juncus kraussii and Baumea juncea. Abbreviations: CP, Conservation
Park; H, height of nest mound; Circ., circumference of nest mound at base.

Ref.

No.

Locality

Lat (S)

Long (E)

Veg. Zone

H

(cm)

Circ.

(m)

Material

Holes

Veg. Cover

2

Kauri Ck CP

25°46 , 57”

152°58’28 M

scdgeland

60

4.5

peat/mud/

sand

12; 10 0cm, 10cm,
20cm

S. virginicus

6

Kauri Ck CP

25°47 , 14"

152°57'44"

sedgeland

35

1.5

black soil

2; 2 @ 0cm

J. kraussii

9

Noosa North
Shore

26°23*31”

153°03’58”

sedgeland

31

2.5

grey sand

6; 2 @ 0cm, 2 @ 2cm, 2
@ 3cm

J. kraussii,

S. virginicus

10

Noosa North
Shore

26°23’35”

153°03’47”

sedgeland

36

3.0

sand

3; 2 @ 0cm. 5cm

S. virginicus

14

Pumicestone

Passage

26°59’10”

153°03’46”

Sporobolus

60

6.0

dark loam

25; 6cm, 2 @ 8cm, 9cm,
10cm, 2 @ 13cm, 14cm,

1 5cm. 2 @ 1 6cm, 3 @
17cm, 2 @ 18cm, 19cm,

2 @ 20cm. 2 @ 21cm, 2
(2) 22cm, 31cm, 32cm

S. virginicus

15

Pumicestone

Passage

26°59 , 18”

153°03'51”

chenopod

35

4.7

peat/mud

9; Ocm, 3cm, 5cm. 6cm,
8cm, 3 (2) 10cm. 1 1cm

S. virginicus

16

Pumicestone

Passage

26°59 , 21”

153°03'59”

Sporobolus

40

3.8

peat/mud

13; 6 @ Ocm, 8cm, 10cm,
14cm, 15cm, 16cm,

1 8cm, 25cm

S. virginicus,

S. arbusculoides

17

Pumicestone

Passage

26°59’2r

isswos"

mangrove

60

5.1

peat/mud

23; 12 @ Ocm, 3 @
12cm. 3 @ 15cm. 3 @

1 8cm, 22cm, 24cm

S. virginicus,

S. australis

Pumicestone

Passage

26°59’2r

1 53°04’09”

Sporobolus

48

4.7

mud/sand/

loam

10; Ocm, 3cm, 5cm, 8cm,
10cm, 12cm. 14cm,
20cm, 34cm, 48cm

S. virginicus

19

Pumicestone

Passage

26°59’26”

153°04’0r

Sporobolus

48

4.1

mud

8; 3cm, 8cm, 1 1cm,
12cm, 2 @ 13cm, 24cm,
46cm

S. virginicus

20

Pumicestone

Passage

26°59 , 26”

153°04’08”

Sporobolus

27

2.4

peat/loam

8; 2 @ Ocm, 6cm, 2 @
8cm, 10cm, 2 15cm

S. virginicus

21

Pumicestone

Passage

26°59’3P

153°03’42”

Sporobolus

40

4.5

mud/loam

19; 13 @ Ocm, 5cm, 3 @
8cm, 10cm, 1 1cm

S. virginicus

29

Bullock Ck CP

27°00'47”

153°04’ir’

sedgeland

48

3.2

clay/mud/

sand

17; 13 @ Ocm, 10cm,
13cm, 14cm, 15cm

S. virginicus

57

Rainbow

Channel

27°27’35”

153°25’38”

sedgeland

66

4.7

peat/mud/

sand

6; 3 @ Ocm, 20cm, 40cm,
45cm

sedge* (1.6m),
Vitex bicolor

63

Stockyard

27°43 , 29"

1 5r24'26"

mangrove

23

3.7

mud

none (abandoned)

dead S. australis

76

Coomcra R

27°50’27”

153°22’41”

Sporobolus

25

3.4

mud

1 ; 20cm

S. virginicus

77

Coomera R

27°50 , 30"

153°22’45”

Sporobolus

25

6.0

mud

7; 5 @ 5cm, 2 20cm

S. virginicus

87

Coomcra R

27°50’52”

153°22’22”

Sporobolus

36

3.6

mud

4; 10cm, 2 @ 25cm,
31cm

S. virginicus

95

S Stradbrokc

27°5 1 ’34"

153°25’06”

sedgeland

37

4.2

black peat

5; 4 @ 0cm, 33cm

J. kraussii (1.3m),

A. speciosum

102

S Stradbroke

27°51’39* t

153°25’08”

sedgeland

40

4.4

peat/grey

sand

6; 3 @ Ocm, 24cm, 35cm,
40cm

J. kraussii ,

A. speciosum

103

S Stradbroke

27°5r39”

153°25M0”

sedgeland

50

4.8

peat/grey

sand

13; 5 @ Ocm, 7 @
20-25cm, 40cm

J. kraussii ,

A. speciosum

intertidal area (Donnybrook #s 31,32). An
intermediate degree of moating at high tide was
noted for two nests, #s 56 and 57, at Rainbow
Channel (210° and 200° moating, respectively).
The mean height of supralittoral bank nests was
55cm (range = 35-80cm, SD = 15cm, n = 17).

Nests were documented up to 32m from the
mangrove community and up to 11 access holes
were recorded. Ten of the nests were incorporated
among the roots of living or dead trees or shrubs.

4. Tree Trunk Nests. Tree trunk nests relied on a
hollow tree or stump to provide the supportive

462

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3. Island nests (Type 2) of Xeromys myoides from southeast Queensland. Abbreviations: CP,
Conservation Park; H, height of the island plus any additional mounding (often it was impossible to dissociate
the two); Circ., circumference of the island. * The term ‘sedge’ refers to the combination of Juncus kraussii and
Baumea juncea.

Ref.

No.

Locality

Lat (S)

Long (E)

Veg. Zone

H

(cm)

Circ.

(m)

Material

Holes

Veg. Cover

5

Kauri Ck CP

25°47’10”

152°58’24”

chenopod

45

5.1

mud/sand

4; 3 @ 0cm, 10cm

J. kraussii , C. glauca

22

Gallagher Pt

27°00’18”

153°06’03”

sedgeland/

Sporobolus

45

3.9

peat/loam

4; 0cm, 2 @ 8cm, 1 1cm

dead C. glauca
( 1 .2m), S. virginicus

23

Gallagher Pt

27°00’18"

153°06’03”

sedgeland/

Sporobolus

32

3.1

peat/loam

4; 3 @ 0cm, 12cm

S. virginicus, dead

C. glauca (0.7m),

M quinquenervia

27

Bullock Ck
CP

27°00’43”

153°04’1 1”

sedgeland

55

10.2

mud/grey

sand

3; 3cm, 2 @ 10cm

S. virginicus, C.
glauca (7.0m & 1 .2m)

34

Donnybrook

27°00’59”

153°02’57”

Sporobolus

45

6.2

clay-mud/

loam

6; 2 @ 10cm, 2 @
13cm, 22cm, 45cm

S. virginicus, A.
corniculatum (lm)

37

Donnybrook

27°01’08”

153°03’09”

mangrove

75

5.6

peat/mud

7; 3 @ 20cm, 23cm, 2
(2) 30cm, 45cm

S, virginicus,

C. glauca (4m)

38

Donnybrook

27°01W*

153°03’03”

Sporobolus

74

9

clay/

humus/sand

3; 29cm, 39cm, 72cm

S. virginicus

42

Donnybrook

27°0ri6"

153°03’00”

Sporobolus

58

12

clay/loam

4; 2 @ 0cm, 3cm, 37cm

S. virginicus,

B. halimi folia

43

Donnybrook

27°0ri7"

153°03’08”

sedgeland/

Sporobolus

58

12

clay/loam

6; 8cm, 9cm, 10cm,
20cm, 29cm, 31cm

S. virginicus,

C glauca (8m)

44

Donnybrook

27°0r21”

153°03’13*’

chenopod

59

13.7

clay/loam

5; 0cm, 8cm, 2 @
10cm. 21cm

S. virginicus, sedge*,

A. marina (1.6m)

45

Donnybrook

2V0V2T'

153°03’12”

chenopod

60

7

clay/loam

2; 27cm, 42cm

S. virginicus,

M. quinquenervia (4m)

48

Amity

27 0 24'41"

153°26’23”

sedgeland

45

2.7

grey-black

peat/sand

7; 7 @ 0cm

S. virginicus,

M. quinquenervia (4m)

49

Amity

27°25’25”

153°26’14”

sedgeland

60

4.1

grey

peat/sand

3; 2 @ 0cm, 20cm

B. juncea,

M. quinquenervia (4m)

50

Amity

27°25’26”

153°26’15”

sedgeland

40

7

nil

2; 2 @ 25cm

sedge* (lm),

C. glauca,

M. quinquenervia

1 51

Amity

27°25’31”

153°26’13”

sedgeland

60

3.1

grey

peat/sand

5; 2 @ 0cm, 2cm, 4cm,
48cm

sedge*,

M. quinquenervia (5m)

60

Rainbow

Channel

2V2V52"

153°25’39”

sedgeland

60

2.4

peat/mud

5; 3 @ 0cm, 2 @ 60cm

sedge* (1.6m),
Phragmites australis,

M. quinquener\’ia
(4m), C. glauca (6m)

72

Coomera R

27°50’23”

153°22’25”

sedgeland

40

9.8

black sandy
peat

5; 3 @0cm, 12cm,
35cm

S. virginicus,

C glauca (9m)

74

Coomeca R

27°50’24”

153°22’24”

sedgeland

30

6.4

black sandy
peat

2; 0cm, 30cm

S. virginicus,

J. kraussii.

C glauca (4m)

106

S Stradbroke

27°5r41”

153°25’09"

sedgeland

38

5.1

grey sand

6; 5 @ 0cm, 1 5 cm

J. kraussii ( 1 .3m),

A. speciosum ( 1 .2m),

M quinquenervia (4m)

1 110

S Stradbroke

27°5 1 ’44”

153°25’07”

sedgeland

45

8.8

peat/mud/
grey sand

7; 6 @ 0cm, 1 7cm

J. kraussii (1.3m),

A. speciosum (1.3m),

M. quinquenervia (5m)

frame for the mud structure built within. These
mostly involved dead stags of Eucalyptus
tereticornis (Fig. 9) or living or dead Avicennia
marina situated within the mangrove zone (Figs
10-13). Additional examples of tree trunk nests
involved living or dead Casuarina glauca ,

Melaleuca quinquenervia or Excoecaria
agallocha growing at or near the marine/
terrestrial boundary.

In spite of the number of tree trunk nests
recorded in or adjacent to the mangrove

NEST TYPES OF XEROMYS MYOIDES

463

FIG. 7. A supralittoral bank where tunnels made by Xeromys myoides are
either hidden among roots or are indistinguishable from crab holes (nest
#62, Canalpin Creek, North Stradbroke Island, September 1997). Photo
Steve Van Dyck.

community (31), this nesting strategy was not
documented widely throughout the survey area
(Table 5). Numerous examples (14) were
recorded from a limited area (approximately 60m
x 530m) on South Stradbroke Island inside the
hollowed bases of large, decayed Eucalyptus
tereticomis stumps, now completely surrounded
by a mangrove open woodland. Although an
almost unlimited number of hollow-trunked
mangroves is available, only ten records (Noosa
North Shore #12; Donnybrook #36; Coomera
River #s70,71,73, 75,78-81)
were made of nests inside the
trunks of living mangroves
( Avicennia marina). Two other
records were of nests inside
the trunks of dead mangroves.

In one case (Noosa North
Shore #11), the tree involved
was the rotting stump of a
Milky Mangrove Excoecaria
agallocha. In the other
(Donnybrook #30), a nest was
discovered in the small,
leaf-lined (leaves of Aegiceras
corniculatum) trunk of a dead
mangrove, possibly Avicennia
marina. The remaining five
tree trunk nests were located
inside dead or hollow-trunked
but living Melaleuca
quinquenervia (#s 1,3) or

Casuarina glauca (#s
4,28,52) growing at the
marine/terrestrial boundary or
within the uppermost zone of
tidal influence.

Tree trunk nests assumed a
variety of forms. In most cases
cavities within living or dead
trees were either packed with
mud or contained a mounded
mud structure visible from the
outside (Figs 9,10). An
exception was discovered
within a living Avicennia
marina at Coomera River
(nest #75). Here, the basal
hollow was not entirely mud-
filled but instead contained a
60cm-high, ramped mud
structure built against the
tree’s sloping, interior wall.

Other tree nests were
located within relatively small
trunks that lacked large holes and so precluded
the structure of the nest being observed from the
outside. Consequently, it was impossible to
determine whether or not the internal cavity was
mud-filled. In some cases, it was not even
obvious that such trunks were hollow'. Even so, X.
myoides was clearly occupying these trees
because of additional mud working including
mounds with at least one access hole built against
the tree’s base (Fig. 11), plastering of the tree’s
exterior surface, footprints creating tracks along

bank nest with additional mounding (nest #24,
March 1 999). Photo Ian Gynther.

FIG. 8. A supralittoral
Gallagher Point, Bribie Island,

464

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG 9. A tree nest at the base of a Eucalyptus tereticornis stag (nest #105,
South Stradbroke Island, June 1995). Photo Ian Gynther.

the uppermost surface of sloping trunks,
especially near ground level, and plugging of
knot holes or the ends of broken trunks or
branches (Fig. 12). In one example (Coomera
River #80), the plugging of a gap in the upper
surface of a dead, horizontal trunk of a living A.
marina apparently led to the construction of a
small mound of mud (10cm high) atop the
broken-off trunk at a height of 86cm above ground
(Fig. 13). When examined on a subsequent visit,
this mound had been destroyed and a nesting
chamber of leaves inside the trunk's cavity was
visible. Change over time in the extent of mud
working associated with tree
trunk nests was not un-
common and, in certain cases
involving smaller diameter
mangrove trees in particular
(e.g. Coomera River nest #s
70,73,78), nests were not
active on later visits or could
not be relocated at all as no
signs of former occupation by
X. myoides were detectable.

A final variation in tree trunk
nests was seen in situations
involving the broken and
decaying stumps of Casuarina
glauca or Melaleuca quin -
quenervia near the edge of the
sedge zone (Kauri Creek
Conservation Park #s 1,4;

Bullock Creek Conservation
Park #28). Here, mud mounds

were constructed in and around
the remains ofthe stump such that
the timber appeared to act as
internal reinforcing for the com-
pleted structure.

Occasionally the local landform
at sites with tree trunk nests
prevented individual nests fitting
neatly into the standard habitat
zonation scheme. In these cases,
Table 5 provides simple
descriptive terms for the physical
location/vegetation at the nest
site. For example, "woodland/
sedgeland’ was applied to
situations where a distinct supra-
littoral bank was lacking at the
boundary between the intertidal
area and adjacent Melaleuca
quinquenervia or Casuarina
glauca woodland. The term
‘woodland tongue' was applied to a promontory
of dry land that lay between areas of mangrove
and saltmarsh. The overall landform and the size
of the tongue made it too big to be considered an
island and its situation within the intertidal zone
ruled out the possibility of it being termed a true
supralittoral bank.

Visible mud heights in Type 4 nest structures
reached 86cm above the surrounding littoral
substrate (Coomera River #80), but in some cases
may have been higher in the concealed cavities
inside the trunks. Nest #98 at South Stradbroke
Island, a small mound inside a wide, hollow

FIG 1 0. A tree nest in the trunk of a living Avicennia marina (nest #12, Noosa
North Shore, April 1997). Photo Ian Gynther.

NEST TYPES OF XEROMYS MYOIDES

465

TABLE 4. Supralittoral bank nests (Type 3) of Xeromys myoidcs from southeast Queensland. Abbreviations: M,
height of mound structure, if present; B, height of supralittoral bank; H, total nest height, i.e. M+B; Circ., basal
circumference of any mounding; indet., indeterminate. Hole heights are measured from the bank base. * The
term ‘sedge' refers to the combination of Juncus kraussii and Baumea juncea.

1

Ref.

No.

Locality

Lat (S)

Long (E)

M/B

(cm)

H

(cm)

Circ.

(m)

Material

Holes

Veg. Cover

7

Noosa North

Shore

26°23’10"

i53°04’i0”

16/64

80

1.9

sand

2; 63cm, 68cm

C. glauca (8m)

8

Noosa North

Shore

26°23’12”

153°04’1 1”

20/43

63

1.6

peat/sand

2; 2 @ 43cm

S. virginieus, sedge*

24

Gallagher Pt

27°00’20”

1 53°05'59"

43/27

70

4.1

loam/sand

2; 27cm, 45cm

S. virginictis,

J. kraussii,

C. glauca (8m)

25

Gallagher Pt

27°00’29”

153 o 05’52”

?/?

38

3.8

loam/white

sand

5; 2 @ 2cm, 6cm,
17cm, 38cm

S. virginieus, J.
kraussii . B. halimifolia

26

Gallagher Pt

27°00 , 31”

153°05’52”

21/39

60

4.1

loam/white

sand

2; 2 @ 0cm

S. virginieus,

J. kraussii,

C. glauca (4m)

31

Donnybrook

27 o 00'56”

153°02’56”

44/?

7

4.3

peat

1 1 ; 5 @ 8+?cm, 4 @

1 5+?cm, 2 @

1 8+?cm

burnt ('IS. virginieus),

C. glauca (4m)

32

Donnybrook

27°00’57 v

153°02’56”

25/32

57

5.0

clay/humus

9; 0cm, 2cm, 4cm,
15cm, 33cm, 34cm,

2 @ 38cm, 47cm

S. virginieus

33

Donnybrook

27°00’58”

153°02’55”

38/?

7

6.8

?peat (burnt
out)

5; 2 @ 8+?cm,
15+?cm, 16+?cm,
30+?cm

burnt (IS. virginieus),

C. glauca (4.5m)

35

Donnybrook

27°oror

153°02’59”

25/50

75

3.4

sand/clay

3; 2 (a). 25cm, 30cm

nil

39

Donnybrook

27°0r09”

153°03*08”

20/30

50

2.2

sand/clay

5; 2 @ 0cm, 1 5cm,
35cm, 40cm

S. virginieus

40

Donnybrook

27°onr

153°03’13”

?/?

35

3.1

sand/loam

5; 2 @ 0cm, 2cm,
10cm, 15cm

nil

41

Donnybrook

27°0ri 1”

153°03’14”

24/40

64

2.8

sand/clay

4; 0cm, 40cm,
45cm. 55cm
(plugged)

S. virginieus

53

Rainbow Channel

27°27’28”

153°25’43”

10/30

40

2.2

pcat/mud/

sand

2; 0cm, 30cm

J. kraussii (lm),
Imperata cvlindrica ( 1 m)

54

Rainbow Channel

27°27’29”

153°25’43”

0/35

35

N/A (no
mound)

nil

1; 8cm

sedge*

55

Rainbow Channel

27°27’30”

153°25’43”

0/35

35

N/A (no
mound)

nil

undetected

tree roots

56

Rainbow Channel

27°27’34”

153°25’43”

30/40

70

3.9

peat/mud/

sand

6; 4 @ 0cm, 2 @
70cm

J. kraussii (1.6m),

M. quinquenenia (5m) |

58

Rainbow Channel

27°27'40”

153 o 25’40"

30/30

60

3.9

peat/mud

6; 0cm, 4 @ 30cm,
60cm

J kraussii

59

Rainbow Channel

27°27’44”

153°25*49”

10/?

7

1.1

peat/sand

3; ?cm, 2 @ 10+?cm

J. kraussii,

Caustis blakei,
Gahnia sieberiana

61

Rainbow Channel

27°28 , 01”

153°25’36”

15/20

35

indet.

pcat/mud

2; 0cm, 35cm

J. kraussii,

B. halimifolia

62

Canalpin Ck

27°36’19”

153°24’38”

0/60

60

N/A (no
mound)

nil

1; 32cm

Gahnia sp.,

M. quiiu{uener\ia (11m),

B. halimifolia (2m)

trunk, was the lowest recorded tree nest at only
25cm. Mean nest height inside tree trunks was
59cm (SD = 1 8cm, n = 26). Additional plastering

of mud against the interior or exterior surfaces of
the tree often extended much higher than the nest
heights indicated in Table 5. Furthermore, in

466

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 5. Tree trunk nests (Type 4) of Xeromys myoides from southeast Queensland. Abbreviations: CP,
Conservation Park; H, height of visible mud structure only (actual nests may be higher inside trunks); Circ.,
maximum circumference of the tree nest at ground level (where relevant, including extent of any mud mounding
or buttress roots used as nest access points); indet., indeterminate.

Ref.

No.

Locality

Lat(S)

Long (E)

Vcg. Zone

H

(cm)

Circ.

(m)

Material

Holes

Tree Species

1

Kauri Ck CP

25°46’57”

152°58’24”

sedgeland

35

3.2

peat/black

soil/sand

5; 2 @ 0cm, 8cm, 2
@ 20cm

M. quinquenervia , dead,
lit. ?, CBH N/A

3

Kauri Ck CP

25°47’0r

152°58’22”

woodland/

sedgeland

28

3.3

peat/sand

5; 5 @ 0cm

M. quinqueneivia , live,
Ht. ?, CBH ?

4

Kauri Ck CP

25°47W

1 52°58* 0”

sedgeland

25

1.8

black

loam/sand

1; 0cm

C. glauca, dead, Ht. ?,
CBH?

11

Noosa North
Shore

26°23*41”

153°03’43’*

woodland

tongue

51

2.7

grey sand

3; 0cm, 8cm, 15cm

E. agallocha , dead, Ht
0.6m, CBH N/A

12

Noosa North
Shore

26°23'4P

153°03’45”

mangrove

63

1.9

sand

2; 0cm, 9cm

A. marina , live, Ht
6.5m, CBH 1 .9m

28

Bullock Ck

CP

27°00'47*’

153 o 04*ir

sedgeland

45

3.4

mud

6; 2 @ 0cm, 5cm,
15cm, 16cm, 23 cm

C. glauca , dead, Ht
0.45m, CBH N/A

30

Donnybrook

27°00’55”

153°03’0r

mangrove

80

0.9

peat/mud

2; 2 @ 0cm

? A. marina, dead, Ht
1.8m, CBH 0.95m

36

Donnybrook

27 o 01’06”

153°03*09"

mangrove

48

3.6

mud

3; 2 @ 0cm, 32cm

A. marina, part-live, Ht
8m, CBH 2.8m

52

Rainbow

Channel

2T2T2T

153°25*45”

woodland/

sedgeland

indet.

1.3

peat

1 ; 0cm

C. glauca, live, Ht 12m,
CBH 1.2m

70

Coomera R

27°50’17”

153°22’49"

mangrove

85

1.1

mud

2; 0cm, 85cm

A. marina, live, Ht 6m,
CBH 0.8m

71

Coomera R

27°50*22”

153°22*51”

mangrove

indet.

0.7

mud

2; 0cm, 30cm (in
wood)

A. marina, live, Ht 5m,
CBH 0.8m

73

Coomera R

27°50’23”

153°22’53”

mangrove

indet.

0.9

mud

1; 0cm

A. marina, live, Ht 5m,
CBH 0.8m

75

Coomera R

27°50’24 M

153°22’31"

mangrove

60

2.4

mud

1; 30cm

A. marina, live, Ht 6m,
CBH 0.85m

78

Coomera R

27°50’31”

153°22 , 22"

mangrove

indet.

1.8

mud

2; 1cm, 2cm

A. marina, live. Ht 6m, j
CBH 0.7m

79

Coomera R

27°50’32 , ‘

153°22’37”

mangrove

80

1.7

mud

2; 2 @ 0cm

A. marina, live. Ht 6m,
CBH 1.1m

80

Coomera R

27°50 , 35’*

153°22’19”

mangrove

86

1.6

mud

3; 3 @ 0cm

A. marina, live, Ht 5m,
CBH 0.57m

81

Coomera R

27°50’35”

153°22’19”

mangrove

indet.

2.0

mud

4; 0cm, 2 @ 15cm,
30cm (tree hole)

A. marina, live, Ht 6m,
CBH 0.68m

92

S Stradbrokc

27°5P2 T

153°25*01"

mangrove

75

1.8

mud/grey-
black sand

undetected

E. tereticomis, dead, Ht
3m, CBH 1.25m

93

S Stradbfoke

27 # 5r30”

153°25*00"

mangrove

66

3.5

mud/grey-
black sand

2; 30cm, 40cm

E. tereticomis, dead, Ht
2.1m, CBH 1.3m

94

S Stradbrokc

27°5r30”

153°25 , 00"

mangrove

63

2.6

mud/grey-
black sand

2; 2 @ 63cm

E. tereticomis, dead, Ht
4m, CBH 1.7m

96

S Stradbrokc

27°5 1 ’35~

153°25*Or

mangrove

72

3.1

mud/grey-
black sand

3; 3 @ 0cm in but-
tresses

E. tereticomis, dead, Ht
1.9m, CBH 1.25m

97

S Stradbrokc

27°51*36"

153°25’0r

mangrove

56

4.7

mud/ grey-
black sand

5; 4 @ 0cm, 20cm

E. tereticomis, dead, Ht
lm, CBH lm

98

S Stradbroke

2T5VYT

153°25’02”

mangrove

25

1.1

mud/grey-
black sand

2; 2 @ 0cm under
logs

£. tereticomis, dead, Ht
1.6m, CBH 1.9m

99

S Stradbroke

2TSV2>T'

153°25 , 03”

mangrove

65

3.1

peat/grey
sandy mud

2; 2 @ 25cm

E. tereticomis, dead, Ht
1.3m. CBH 1.2m

100

S Stradbroke

27°5r38 , ‘

153°25'04”

mangrove

54

3.0

mud/grey

sand

2; 0cm, 40cm

E. tereticomis, dead, Ht
2.8m, CBH 1.5m

NEST TYPES OF XEROMYS MYOIDES

467

TABLE 5 (Com.)

Ref.

No.

Locality

Lat (S)

Long (E)

Veg. Zone

H

(cm)

Circ.

(m)

Material

Holes

|

Tree Species

101

S Stradbroke

27°51’38”

153°25’04”

mangrove

62

4.2

mud/grey

sand

8; 8 @ 0-32cm

E. tereticornis , dead, Ht
2.5m, CBH 1.6m

104

S Stradbroke

27°5r40”

153°25’04”

mangrove

35

2.7

mud/grey-
black sand

5; 0cm, 4 @ 10cm

E. tereticornis , dead, Ht
9m, CBH lm

105

S Stradbroke

27°5T4r

153°25’04”

mangrove

60

3.1

mud/grey

sand

4; 3 @ 0cm, 1 5cm

E. tereticornis , dead, Ht
2.5m, CBH 1.5m

107

S Stradbroke

27°5 1 ’42”

153 0 25W

mangrove

60

3.1

mud/grey

sand

2; 2 @ 0cm

E. tereticornis, dead, Ht
3m, CBH 2.1m

108

S Stradbroke

27°5 1 ’42”

153°25'05”

mangrove

60

2.5

mud/grey-
blaek sand

5; 5 @ 0-25cm

E. tereticornis, dead, Ht
3m, CBH 2.3m

109

S Stradbroke

27°5r44"

153°25'04"

mangrove

85

4.3

grey sand

5; 5 @ 0cm ( 1 in
buttress)

E. tereticornis, dead, Ht
8m, CBH 1.55m

small diameter trees mud plugging of knot holes
and other gaps in the tree's outer walls were
sometimes seen at considerable heights. In one
case at Coomera River (#8 1 ), a plugged knothole
was noted 1 .75m above ground, while other holes
at heights of 1 .4m and 1.1m were also blocked
with mud.

Up to eight entrance holes were recorded in
tree trunk nests but, given the number of exposed
'buttress' roots through which access to some
nests might have been gained, this total was
probably an underestimate. Other tree nests had
no visible access points in the trunk or roots but
did possess mud mounds with entrance tunnels
constructed against the base of the trunk.
Recorded examples of such mounds ranged in
height from 10-42cm and contained 1-3 access
holes, sometimes with fluted entrances. These
mounded structures were of insufficient height to
represent nests themselves but appeared to
provide access to one or more holes in the nest
tree at or near ground level. This was not con-
firmed in any of the documented cases because it
would have necessitated destroying the
associated mound.

Because of the location of many of this
extraordinary range of tree trunk nests deep
within the mangrove community (up to 265m
from the landward mangrove zone edge), most
experienced longer periods of inundation and
deeper moating than other X. myoides nest types.
The only tree trunk nests recorded that did not
receive 360° moating during the tidal cycle
(Noosa North Shore #11; Rainbow Channel #52)
involved trees standing on the supralittoral bank.
In both cases, the maximum extent of moating
experienced at high tide was 180°.

5. Spoil Heap Nests. Spoil heap nests were those
constructed in human-made piles of excavated or
bulldozed earth (Fig. 14), soil clods among roots
of bulldozed trees or in the bund walls associated
with drainage or flood mitigation works (Fig. 1 5).
Such artificially created features provided
elevation above the surrounding intertidal
communities and the level of spring tides.

FIG. 1 1 . Mounding at the base of a living (hollow)
Avicennia marina (nest #81, Coomera River,
November 2001). Photo Ian Gynther.

468

MEMOIRS OF THE QUEENSLAND MUSEUM

At Maroochy River (Fig. FIG 13. Mounding on dead, hollow, horizontal trunk of living Avicennia
14) and Donnybrook, nests marina (nest #80, Coomera River, November 2001). Photo Ian Gynther.

FIG 12. Details of mud plugging of hole in trunk of
same living Avicennia marina depicted in Fig. 11
(nest #81, Coomera River, August 2001). Photo Ian
Gynther.

Eighteen nests were recorded
in human-made bund walls or
spoil piles (Table 6). Spoil
heap nest heights ranged from
40-89cm (mean = 56cm, SD =
15cm, n = 18). Although ex-
amples of Type 5 nests were
discovered within each inter-
tidal vegetation community,
the majority was in Sporobolus
grassland, a community that
receives a high incidence of
human-related impacts because
of closer proximity to adjacent
land uses. All spoil heap nests
identified during this study
would have experienced 360°
moating during spring high
tides.

#13 and #46, respectively, were constructed in
combined soil and tree stump waste that had been
bulldozed to near the landward edge of the
mangrove woodland, presumably during
construction of vehicle tracks. The spoil heap
associated with the nest at White Patch on Bribie
Island (#47) resulted from a firebreak being
bulldozed through the wallum vegetation to the
edge of the intertidal zone. Similarly, all nine
Type 5 nests discovered on the north bank of the
Coomera River were in spoil piles created during
past clearing of the site for a development that
was then temporarily abandoned. Some of the
piles included rock, gravel and even concrete
debris (nest #s 86,88,90,91).

At Steiglitz, four nests were found in spoil
heaps originating from the soil associated with
the exposed roots of upturned trees or from
excavation activity during the construction of a
high-banked drainage channel. All piles were
thickly covered with Marine Couch. Large heaps
with circumferences of 7. 3-7. 6m (nest #s 65, 67)
were richly pocked with access holes (19 and 20
holes, respectively) and heavily scored (beneath
the couch) with mud tracks created by the
animals (see Fig. 20). These nests were close
(approximately 130m) to the site of an intensive
marina development. The structurally simple
mangrove community associated with the
Steiglitz nests was probably not older than thirty
years. In the mid-1960s, elevation of the nearby
existing main road (90m to the southwest),
together with the introduction of tidal gates on the

NEST TYPES OF XEROMYS MYOIDES

469

FIG 14. Spoil heap nest in bulldozed material (nest #13, Maroochy River,
March 1 996). Photo Ian Gynther.

Behm’s Creek bridge (700m to the southeast)
resulted in the site undergoing an ecological
succession from a Casuarina glauca/Sporobolus
virginicus community to one dominated by
Avtcennia marina/S. virginicus (G. Leiper, pers.
comm.). Numerous dead C. glauca stags remain
today. The lm-high drainage channel wall,
approximately 10m north of the spoil pile nests
did not show any evidence of nesting activity.
This was not the case at the Pimpama River
locality, where the single recorded nest (#69),
discovered by Peter Lehmann during a radio-
telemetry study, was constructed in the spoil bank
created during excavation of a drainage channel
(Fig. 15). This channel emptied directly into the
Pimpama River, 33m from the nest site.

NEST RECOGNITION.
With some experience, most
active or recently active X.
myoides nests belonging to
each nest class described
above could be identified with
confidence by considering a
combination of the following
features: the overall height of
the nest, the size and shape of
any associated mounding, the
existence of additional work-
ings including mud or peat
plastering and tracks, and the
presence and nature of access
holes. The small percentage of
X. myoides nests that could not
be detected or reliably
identified using visual search
techniques included those
constructed in the supralittoral
bank without any additional associated
mounding. It was necessary to locate these using
radio-telemetry techniques because the profusion

All Type 5 nest sites were within highly
disturbed areas or in close proximity to such
areas. In addition to the White Patch and
Coomera River sites mentioned above, the
Maroochy River nest (#13) was approximately
1 6m from a road skilling a sugar cane plantation,
nest #46 (Donnybrook) was approximately 5m
from a vehicle track and a now felled, exotic pine
plantation, and all Steiglitz nests occurred in an
area less than 200m wide between an artificial
channel draining an abandoned sugar cane
plantation and a road bordering a marina
development. Nest #68 (Jacobs Well) was built in
a large (9m x 11m) spoil heap, 17m from the
boundary of a commercial nursery on the main
Jacobs Well Road.

FIG. 15. Spoil heap nest in material excavated from a
drainage channel (nest #69, Pimpama River, July
1995). Photo Ian Gynther.

470

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 6. Spoil heap nests (Type 5)o fXeromys myoides from southeast Queensland. Abbreviations: H, height of
the spoil heap plus any additional mounding; Circ., basal circumference of the spoil heap.

Ref.

No.

Locality

Lat (S)

Long (E)

Veg. Zone

H

( cm )

Circ.

( m )

Material

Holes

Veg. Cover

13

Maroochy R

26°38’20”

153°04*18”

Sporobolus

67

16

stump/clay/

loam

8; 4 @ 0cm, 2 @ 1 8cm.
20cm, 22cm

*

46

Donnybrook

27°01’22"

153°03’12”

chenopod

89

24

stump/clay
(later burnt)

7; 2 @ 25cm, 30cm,
56cm, 70cm, 2 89cm

S. virginicus

47

White Patch

27°0r40”

1 53°06’59”

sedgeland

45

6.8

peat/loam

9; 3 @ 0cm, 8cm, 2 @

1 0cm, 30cm, 34cm, 40cm

J. kraussii ,
Phragmites
australis

64

Steiglitz

27°45 , 20”

153°20’29’'

Sporobolus

44

3.0

peat/sand

1 2; 2 @ 4cm, 5cm, 2 @
10cm, 3 @ 13cm, 14cm,
15cm, 17cm, 21cm

S. virginicus

65

Steiglitz

27°45’21”

lss^o^r

Sporobolus

60

7.6

sand/loam

1 9; 5 @ 0cm, 7cm, 8cm,
9cm, 2 @ 10cm, 2 @

1 1cm, 2 @ 12cm, 15cm,
20cm, 30cm, 37cm, 42cm

S. virginicus

66

Steiglitz

27°45’2P

153°20'28”

Sporobolus

53

2.6

sand/loam

7; 5cm, 7cm, 2 @ 8cm,
Ucm, 14cm, 15cm

S. virginicus

67

Steiglitz

27045 ’24”

153°20’25”

Sporobolus

80

7.3

stump/peat/

sand

20; 0cm, 2 @ 8cm, 2 @
Ucm. 3 @ 15cm, 16cm,

2 @ 17cm, 2 @ 18cm, 3
@ 20cm, 2 1 cm, 22cm,
53cm, 64cm

S. virginicus ,

C. glauca (4m)

68

Jacobs Well

27°46’22”

153°21’16"

Sporobolus

50

2.6

peat/sand

4; 10cm. 15cm, 21cm,
45cm

S. virginicus

69

Pimpama R

27 0 48’18”

153°20’21”

mangrove

45

12.6

peatmud

8; 5 (a). 0cm, 3 @ 45cm

nil

82

Coomera R

27°50’35”

I53°22’20”

mangrove

47

3.6

peat/mud/

sand

3; 15cm, 30cm, 33cm

S. virginicus

83

Coomera R

27°50’39”

I53°22’17”

mangrove

45

3.0

peat/mud/

sand

2; 24cm, 32cm

S. virginicus,

S. quinquejlora

84

Coomera R

27°50*39"

153°22’19”

mangrove

44

3.1

peat/mud/

sand

1; 0cm

S. virginicus,

S. quinqueflora

85

Coomera R

27°50’5r

153°22’20”

Sporobolus

40

3.8

peat/mud/

sand

2; 2 @ 0cm

S. virginicus

86

Coomera R

27°50 , 51”

153°22’22”

Sporobolus

45

3.3

clay/mud/

gravel

7; 0cm, 5cm, 19cm,
20cm, 24cm, 2 @ 29cm

S. virginicus

88

Coomera R

27°50’54 M

153°22’26”

Sporobolus

62

8.1

mud/gravel/

rock

8; 3 @ 10cm, 40cm,
43cm, 2 @ 49cm, 55cm

S. virginicus

89

Coomera R

27°50’55”

153022*21”

Sporobolus

40

4.5

heavy loam

4; 2 @ 0cm, 1 5cm, 30cm

S. virginicus

90

Coomera R

27°50’55”

153°22 , 26”

Sporobolus

64

7.2

mud/gravcl/

rock

6; 1 2cm, 1 7cm, 25cm,
30cm, 36cm, 47cm

S. virginicus

1 91

Coomera R

27 o 50 , 56 , '

153°22’28”

Sporobolus

80

11.1

mud/gravel/

debris

3; 5cm, 21cm, 31cm

S. virginicus,

S. quinquejlora

of crab holes that occurred in supralittoral banks
made positive visual identification of nest
entrances impossible. Also, nests that had been
abandoned for a long period of time were difficult
to identify with confidence because all external
signs of occupation (holes, tracks, plastering,
etc.) had disappeared. As an illustration of this,
only one abandoned nest (a free-standing mound
designated as Stockyard #63) could be reliably
attributed to X myoides during this study. This
was because its history of occupation was known.

The various features that aid in the identification
of X. myoides nesting structures are described in
more detail here.

Overall Nest Height and Moundings. For all X.
myoides nest types, a plot of the overall heights of
nests above the surrounding substrate of the
intertidal zone revealed an approximately normal
distribution. The mean height of extant nests
across all nest classes was 53cm (range =
25-89cm, SD = 16cm, n = 101). Two-thirds of
these occupied nests had heights within the range
of 3 1 -60cm, with only 7% and 27% of nests being
smaller or larger, respectively This typical size

NEST TYPES OF XEROMYS MYOIDES

471

SEDGELAND / CHENOPOD SI I Kl BLAND / SPOROBOLUS GRASSLAND

MARINE MUDFLATS

FIG. 16. Diagram of intertidal community zonation
from the supralittoral bank (top) to the marine
mudflats showing the variety of Xeromys myoides
nesting strategies documented from the 110 nests
encountered during this study. The numbers
represent totals for each nest type recorded within
each zone.

range provides a useful guide when assessing
whether potential nest structures encountered
during survey work belong to X. myoides.

All nest types involved, or could incorporate,
characteristic moundings of mud or other
substrate material. The mound structures
associated with free-standing nests ranged up to
66cm in height and were often conspicuous in
their surroundings because the mound accounted
for the nest's total height. However, those found
in association with island, supralittoral bank or
spoil heap nests were generally smaller in all
dimensions because the raised substrate on which
the nest was located already provided substantial
elevation and, therefore, protection of the nest
against high tides. In all cases, however, the
overall profile of the mounded structure was
similar — approximating an inverted paraboloid.

In situations of tall or dense vegetation,
mounded structures created by X. myoides were
much easier to detect and identify where fires had
recently burned the survey area. This was true for
mounds associated with island nests (Fig. 1 7) but
was particularly so for the large mud mounds of
free-standing nests and those on the supralittoral
bank that would otherwise have been concealed
by surrounding sedgeland (Fig. 18).

In the area of southeast Queensland in which
this study focused, the naturally occurring
structures most likely to be mistaken for X.
myoides nests were various mounds made by
intertidal crab species. The most frequently
encountered were the low, irregular mud mounds
found in the outer (more
seaward) portions of the
mangrove community, usually
amongst stands o iRhizophora
stylos a. These were created by
Neosarmcirtium trispinosum
and Perisesarma messa. Two
main indicators that these
were not Water Mouse nest
structures were the abundance
of such mounds (at times
covering large areas amid the
mangroves) and their limited
height (most <25cm). Given
their position in the intertidal
zone, it was quite apparent
that even the tallest of these
structures would be entirely
inundated at high tide. Never-
theless, such crab mounds may
offer valuable protection to X.
myoides because many

472

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 18. A mound associated with a supralittoral bank nest exposed by Fire
(nest #61, Rainbow Channel, North Stradbroke Island, September 1997).
Photo Steve Van Dyck.

captured individuals ran into these holes upon
release from our traps.

Plastering and Tracks. The tops of mounded
structures associated with active A', myoides nests
often bore signs of recent 'earthworks’ in the
form of plastering or daubing. This frequently
involved additions of a mud or peat slurry that,
over time, gradually served to increase the
mound’s overall height. In other cases, the
material added w'as not as fluid, instead forming a
peaty layer in which small (<lcm diameter),
roughly spherical balls of
substrate were compacted
together (see Fig. 1 8 ). In what-
ever form it took, the fresh
daubing was often worked
into and among the bases of
living stems of sedges or
Sporobolus to a height of many
centimetres (see Fig. 18).

The style of plastering used
by Water Mice to add height to
their mound structures was
also used to repair any damage
to the top or sides of the mound.

For example, where Feral Pigs
Sus scrofa had breached the
side of a free-standing nest at
Pumicestone Passage (#21),
the resulting hole was plugged
with new ly added peat material
of a markedly different colour
and texture to the surrounding

mound, indicating it had come
from a different source and
was added later. Another
example from Gallagher
Point, Bribie Island is shown
in Fig. 19.

Plastering and daubing were
also features of tree trunk
nests. In fact they were often
the only visible evidence of
nesting activity in trees
because the very nature of such
nests often made it impossible
to see the full extent of the
workings within the hollow
trunk or stump. At times, mud
or peat packing could be seen
inside the tree’s hollow cham-
ber when viewed from above
(see Fig. 10) or by examining
knot holes or openings left
where trunks or branches had
broken off. The use of plastering in such
situations was similar to the plugs constructed
following damage to other AT. myoides nest types.
Mud workings associated with tree trunk nests
also included mounds with access holes
constructed against the trunk at ground level (Fig.

1 1, see above). Nest #80 at Coomera River, with
its small mound of mud adorning the upper
surface of a horizontal section of trunk well
above ground (Fig. 13), represented a more
exaggerated example of plastering.

FIG 19. Repair work to damaged nest mound (nest #24, Gallagher Point,
Bribie Island, March 1999). Photo Steve Van Dyck.

NEST TYPES OF XEROMYS MYOIDES

473

FIG. 20. Free-standing nest showing slurry track
leading from access hole to mound top (nest #17,
Pumicestone Passage, November 1996). Photo Steve
Van Dyck.

The material used by X. myoides to plaster the
tops of nest mounds or repair breaches to mound
structures was brought from one or more access
holes and then smeared to the site where
additions were being made. This action, which in
captivity was observed (by SVD) to involve the
animal pushing the substrate along with its
forefeet, resulted in clearly defined, slurry tracks
5- 1 0cm wide leading to the top of the nest mound
(Fig. 20). Breaches in the nest structure were also
observed being plugged using the mouth only,
with a captive individual putting a plug in place
with the bottom of the snout. Often each mound
possessed multiple tracks (Fig. 21). By parting
the Marine Couch or sedges growing on the
mound, these obvious tracks could be traced back
to the hole from which the mud, peat or other
material had come. The plastering action that
must be associated with construction of such
slurry pathways was apparent from the way the
material used overlaid the bases of grass and
sedge stems along the route. Once the track
substrate had dried and hardened following
application, these tell-tale signs of energetic

FIG. 21. Multiple slurry tracks on top surface of nest
mound (nest #29, Bullock Creek Conservation Park,
March 1999). Photo Ian Gynther.

building activity persisted long after the work
was actually performed.

A further feature of X. myoides nests useful for
confirming identification is the frequent
inclusion in the daubing and tracks of whole or
partial carapaces of small crabs, particularly
Parasesarma erythrodactyla and He l ice leachii ,
upon which this rodent feeds (Van Dyck, 1997).
Although this was most likely an inadvertent
action on the part of the Water Mouse, the
carapaces would have reinforced the ‘mortar’
formed by mud, peat or other substrates used in
nest construction.

Finally, very fresh tracks, as well as mud or
peat daubing, often possessed a distinctive,
somewhat acrid aroma characteristic of X.
myoides. Whether this smell, detectable to us
only at close range, was due to the animal’s
droppings or to deliberate scent marking using
secretions from its anal glands was not
determined, nor was it ascertained whether fresh
workings always bore this odour. Nevertheless,
this olfactory evidence proved useful in
identifying nests of this species.

474

MEMOIRS OF THE QUEENSLAND MUSEUM

Access Holes . All active X. myoides nests
possessed external holes to provide the animals
with entry and exit points. The size, shape,
number and positioning of these holes offered
clues indicating a potential nest structure did
indeed belong to the Water Mouse. The
occasional exceptions to this were supralittoral
bank nests where no additional mound structure
was associated with the nest and some tree trunk
nests where ground level access was most likely
achieved via existing tunnels in hollow buttress
roots. Where present and visible, nest access
holes were generally larger in overall diameter
than those created by the various southeast
Queensland intertidal crabs encountered during
this study. They were usually circular or
horizontally elliptical. Hole dimensions were not
recorded for each nest observed in this study and
so summary statistics on size variation are not
available. Nevertheless, typical dimensions were
35-38mm in diameter for circular access holes
and 35mm wide x 28mm high for elliptical holes.
Access holes used by X. myoides always gave the
appearance of being open, although observations
made in captivity revealed that at least some
tunnels leading into the nesting chamber within
the mound were blocked some way down with a
plug of mud 5-8cm in thickness (SVD, pers.
obs.). This blockage was not visible from outside
the nest. On occasion we discovered holes that
had dome-shaped caps of mud or peat sealing the
external entrance, but in all such instances the
animal responsible was determined to be a crab.

The number of access holes per nest ranged
from one to 25. Across all nest types, however,
the majority (64%) of nests possessed between
one and five access holes, with a further 25% of
nests having six to ten holes. Of the remaining
nests, 5% had 11-15 holes, 4% had 16-20 holes,
and nests with 2 1 -25 holes accounted tor only 2%
of the total recorded. Based on this trequency
distribution v one would only occasionally expect
to encounter nests of X. myoides with more than
ten access holes when conducting nest searches.

Although access holes could be present at any
height from ground level to the nest top, the
distribution of hole heights for all nest types
involving mounded or elevated structures was
skewed towards the lower third section of the
nest. Consequently, where a structure suspected
of being a nest ofX. myoides possessed multiple
holes, additional evidence supporting these
suspicions was provided by a higher proportion
of holes near the base of the structure.

Access holes showed no obvious difference in
size or shape according to their position on the
nest. Those at ground level or near the base of a
nest structure, however, usually led to tunnels
filled with water. As was frequently observed (by
SVD) in the captive situation, animals negotiated
these Hooded passages before entering the nest
chambers within. Visible pathways with X.
myoides footprints leading away from the nest
were sometimes detected at ground level access
holes, particularly when the surrounding
substrate was boggy. These pathways, created
through frequent usage by resident animals, often
coursed beneath tree roots, dead fall or timber
flotsam.

Confusion with other Rodent Activity. It was
possible to mistake nests of two other rodent
species occupying saltmarsh and mangrove
habitats for those of X. myoides. Nests of the
introduced Black Rat Rattus rattus were
regularly encountered in the hollow branches and
trunks of mangrove trees at heights well above
the level of high tide. These nests usually
involved obvious collections of dried mangrove
leaves, which could be seen through knot holes,
broken ends of branches and occasionally down
the hollow centres of trunks. Mud. peat or other
substrate was never associated with nests of R.
rattus , but nests often contained crab claws and
carapaces. Collections of these crab remains
were also encountered in knot holes, branch forks
and other recesses in trees thought to represent
regular feeding stations used by Black Rats. This
suspicion was supported by the identification of
hair samples collected from such places as Rattus
sp. (B. Triggs, pers. comm.). These sites were
distinguished from the feeding middens created
by X. myoides by their presence above ground
and by the inclusion of the eaten remains of claws
belonging to the crab He l ice leachii. With a
carapace up to 25mm across, this species is
probably near the upper size limit of crustaceans
preyed upon by Water Mice. Observations in
captivity revealed that although X. myoides
attacked and devoured similar sized species (e.g.
Perisesarma mess a ), it did not usually consume
the claws, which consequently remained intact
(SVD, pers. obs.).

Evidence of activity of a second rodent species,
the Swamp Rat R. lutreolus , was occasionally
discovered in the landward portion of the
sedgeland and on the supralittoral bank. This
species typically made obvious runways through
sedgeland that could be traced for considerable
distances among dense ground layer vegetation.

NEST TYPES OF XEROMYS MYOIDES

475

FIG. 22. A pile of spoil created by tunnelling or feeding activity of Rattus
lutreolus in sedgeland (Coomera River, November 2001). Photo Ian
Gynther.

The runways were made more distinct by the
Swamp Rat's habit of chewing rushes or sedges
along the path, leaving only the stem bases
behind. In addition, low mounds or piles formed
from R. lutreolus tunnel spoil were sometimes
encountered. These consisted of a coarse mixture
of balls of often peaty substrate and short lengths
of chewed sedge stems. The resulting mixture
was always much more loosely packed and
friable than the substrate found on nests of X.
myoides (Fig. 22).

DISCUSSION

During our investigation of the Water Mouse's
southeast Queensland distribution, we
successfully discovered at least one nest structure
at all localities at which the species was trapped.
This illustrates the considerable value of em-
ploying nest search techniques when conducting
field surveys. Although the approach is not
entirely foolproof, careful searching for evidence
of nesting activity of X. myoides may represent a
more convenient and efficient survey method by
comparison to the usual technique of Elliott
trapping.

To our knowledge, no other small rodent
constructs conspicuous and relatively immense
mud nesting mounds after the manner of X.
myoides. Although comparison of nesting
techniques can only be made within a limited
field of semi-aquatic rodent genera (see Van
Dyck, 1997), it is very likely the preference of

Xeromys for low altitude, still
water, saline conditions and
regular tidal fluctuations in
water level that has led to the
evolution of this unique
nesting strategy. In some
broad principles of nest
construction, however, there
are counterparts in lodge-
building beavers Castor
canadensis and muskrats
Ondatra zibet hica from the
Northern Hemisphere. Both
species gather vegetation into
sizeable nesting piles (up to
1.8m and 60-90cm above
water level, respectively),
which are accessed through
underwater tunnels. Of even
greater relevance, where earth
banks are high enough for
dens to be well above water
level, or where streams are
swift with an accompanying increase in erosional
force, both beavers and muskrats dig tunnels into
the bank rather than building mounded lodges
( Walker, 1964; Burt & Grossenheider, 1976).

Considering the small size of A", myoides and
the inconvenience that high tides must bring to
initial mound construction in the littoral zone, we
hypothesise that in situations with a normal tidal
range and influence most nests begin in any
suitable ground offering sufficient height above
the upper tidal level. This would explain the
propensity of X. myoides for not only colonising
the supralittora! bank when one exists, but also
raised islands or hummocks, spoil piles, bund
walls and clods of earth amongst the roots of
upturned trees wherever such features occur in
the intertidal zone. However, in exposed
situations where minimal buffering is offered by
the mangrove community and where no sedges or
Marine Couch occur seaward from the supra-
littoral bank (e.g. Canalpin Creek with its
25m-wide, structurally open mangrove zone),
tunnels in the high supralittoral bank may be the
only type of nest present. The protection from
savage erosion provided by a broad mangrove
zone (up to 385m) and abutting zone of sedgeland
(up to 32m wide) in locations such as
Donnybrook and Rainbow Channel gives the
animals time to respond to minor erosion and wet
nest chambers by slowly building up nest height
with the repeated plastering of mud or peat. Thus
examples of additional daubing that formed small

476

MEMOIRS OF THE QUEENSLAND MUSEUM

mounds atop the nest were recorded from all nest
types encountered during the study. On occasions
of extreme high tides, these additions were
sometimes the only part of the nest above water
level (e.g. free-standing nest #s 76, 87, Coomera
River; tree trunk nest #105, South Stradbroke
Island) and, by observing through access holes,
one or more individuals could be seen occupying
these most elevated chambers within the nest
structure. During an unusually high daytime tide
associated with wind and storm surge at Coomera
River, an animal was observed by one of us
(SVD) escaping from the hole at the top of the
nest and being forced to swim to find alternative
shelter. Nearby, an adult female and two young
were seen sitting under S. virginicus cover on the
top of a second nest.

The ‘islands’ with which Type 2 nests are
associated very likely form through erosion of the
supralittoral bank. Presumably, the life of such
supralittoral offcuts is dependent upon their
stabilisation by vegetation cover and their
capacity to endure further erosion. When such
‘islands’ are eventually carved from the supra-
littoral bank, only those sufficiently consolidated
by the roots of trees, shrubs and ground cover
may remain as high points, maintaining their
integrity in the face of spring tides, wind-induced
waves and storm surge. Type 2 nests may then
originate through colonisation of such newly
available high ground within the tidal zone
following the island’s formation. Conceivably,
though, pre-existing supralittoral bank nests may
be sufficiently consolidated within the root mass
of trees to be able to persist with the island as it is
carved oft'. This would offer a second possible
origin of these island nests.

In highly sheltered locations (e.g.. Pumice-
stone Passage), where spring tides fail to
establish a supralittoral bank, free-standing nests,
constructed slowly in the absence of frequent
inundation or tide damage, predominate. This
strategy of constructing large, mounded nests in
an area lacking terrain features that would
provide sufficient height to offer protection
against tides enables X. myoides to colonise
otherwise uninhabitable locations.

Plastering of nests appears to be performed in
response to wet nesting chambers or breaching of
the nest’s outer wall. This conclusion is
supported by the infrequency with which such
mud-daubing occurs once a nest is established.
Plastering of approximately 20cm diameter
increased the height of a bank nest (#53) at

Rainbow Channel from nothing to 6cm in
1 7months. More dramatic plastering events were
also noted. On Bribie Island, the height of a
mound structure on a spoil pile nest (White Patch
#47) increased some 15cm over a maximum
period of four months, and perhaps over a time
span as short as 3-4 weeks (D. Cameron, pers.
comm.). This remarkable rate of construction
may have been in response to a period of
prolonged inundation due to the combined
effects of high rainfall (531.5mm) and tide
heights of up to 2.51m in February 1999.
Although the rate at which plastering and
daubing of a nest structure occurs is undoubtedly
dependent upon the number of nest occupants,
such events as this at Bribie Island are probably
atypical and large, free-standing mounds (e.g.,
nest #s 14, 1 7,57) may represent decades of effort
by generations of mice.

Based on the quantity of substrate that would
be required to construct a large, mounded
structure, we speculate that most, if not all, of the
material used to create the tracks and daubing on
the nest’s top must originate from substrate layers
beneath ground level, i.e. below the nest itself.
This would also account for the sometimes
different nature of the mound and plastering
material as compared to the surface substrate
immediately surrounding the nest. The
observations of Magnusson et al. (1976) support
this hypothesis — in the Melville Island X
myoides nesting mound, tunnels 3-5cm in
diameter were noted to extend as much as 90cm
below ground level. The significant volume of
material that must have been excavated during
their construction was presumably added to the
above-ground mud mound.

The use of spoil piles, clods of soil associated
with the roots of fallen trees and hollow tree
trunks for nesting provided an insight into the
opportunistic way X. myoides uses structures that
provide nest elevation in situations where it is
otherwise in short supply. In addition, tree trunk
nesting demonstrated that if a durable framework
of support were available Water Mice would
occupy the littoral zone well out into the
mangrove community. It was apparent, though,
that not all hollow trunks offer suitable nesting
locations for this rodent. Fourteen of the 3 1 tree
trunk records came from one location on South
Stradbroke Island, and these were all from
Eucalyptus stags. The ‘stranding’ within the
mangrove zone of these large upright stumps, up
to 125m from the marine/terrestrial boundary,
can be attributed to the rise in water level within

NEST TYPES OF XEROMYS MYOIDES

All

the Southport Broadwater and subsequent
erosion and flooding of the low-lying terrestrial
community caused by sand sedimentation inside
the Broadwater after the breaching of the hitherto
connected North and South Stradbroke Islands at
Jumpinpin in 1896 (Connah, 1946; Brooks,
1953). That the present day substrate level is
lower than it was previously is readily apparent
from the exposed root systems of the dead
eucalypts. Presumably, by the time the trunks
were hollow and accessible toX. myoides , the rise
in seawater level had not been so great as to
prevent nest mounds being initiated within the
trees’ protective walls. Furthermore, the ongoing
rise in water level must have been gradual enough
for the mud mounding process to keep abreast of it.

Although the hard-wearing nature of the
eucalypt trunks at this South Stradbroke Island
site still protects nests today from daily tidal
exacerbations, the building of mud structures
from ‘scratch’ in a regularly and deeply flooded
location such as this is probably only possible
under the special circumstances in which a
particular tree offers safe refuge above high tide
to a X. myoides individual prior to and during the
process of nest construction. Of the 12 cases of
nests in mangroves reported here, one (Noosa
North Shore #11) occurred in the stump of a
Milky Mangrove, which, as is typical of this
species, was growing at the marine/terrestrial
boundary, landward of the main mangrove
community. As a consequence, it would have
been inundated only very occasionally during the
highest of spring tides. The other mangrove nests
were, without fail, either in small to large
diameter, sloping trunks or in vertical trees of
small diameter. In both situations, the nature of
the internal hollows presumably enabled the
non-arboreal X. myoides to scramble up inside,
thereby providing dry shelter to the nest-building
individuals during the intervening periods of
tidal inundation when mound construction within
the trunk (or the stopping up of knot holes and
other gaps in the tree) could not be undertaken.
This may explain why so many apparently
suitable mangrove trunks, particularly those of
Avicennia marina , are not utilised for nesting by
X. myoides. Their large diameter, hollow bases
are usually vertical and simply don’t provide
opportunities for X. myoides individuals to climb
up inside.

Although it was obvious when larger diameter
nest trees were cither mud-filled or contained a
mounded mud structure that could house a
nesting chamber, it was never unequivocally

ascertained whether Water Mice also filled the
internal cavities of small diameter trees with mud
or other substrate material for this purpose. It is
possible that the only mud workings associated
with such nests are the plugged external holes
that provide nest security whereas the concealed
spaces within the narrow diameter trunks and
limbs of the tree itself serve as a nesting chamber.
This was suggested by the one tree nest example
(Coomera #80) where it was possible to view a
leaf-filled chamber within a horizontal trunk
following damage to the small mud mound that
had previously capped the roof. By contrast to
large, free-standing mound structures, which
would involve considerable effort to build and
maintain, the use of such trees that require
relatively little mud plastering or packing to
convert them into suitable nest sites may make it
possible for individual X. myoides to occupy
nests on a temporary basis or to maintain multiple
nest trees within a single home range. Such
simple refuges may be utilised by males or newly
recruited individuals, i.e. those animals simply
seeking shelter rather than somewhere to raise
young. Observations in captivity indicated that,
in stark contrast to an adult female, a male X.
myoides used a very basic nest with a chamber
lacking any leaves or grass for lining (SVD, pers.
obs.). Similarly, an adult male individual caught
by hand under a piece of corrugated iron beside
the Tomkinson River, Arnhem Land (Magnusson
et al., 1976) may have been using the site as a
temporary refuge since no nest was found. The
fact that certain tree nests were ephemeral in
nature was demonstrated by the finding that they
were no longer active on our subsequent visits,
with some trees even lacking their once tell-tale
signs of mud daubing. Presumably not long after
a tree nest is abandoned any mud additions fall
into disrepair, particularly when these are
incapable of being consolidated by vegetation
and are submerged and subjected to tidal currents
on a frequent basis.

The same gradual rise in sea level that very
likely led to the proliferation of tree trunk nests in
the eucalypt stags of South Stradbroke Island
may also account for the origin of the
free-standing mounds (nest #s 95,102,103) that
stand amid the sedges and mangrove fern
landward of the mangroves at this site, despite the
now regular inundation of this section of the
intertidal zone. Here, mound construction may
also have been able to keep pace with the long
term, incremental change in water height that
occurred following the break at Jumpinpin. The

478

MEMOIRS OF THE QUEENSLAND MUSEUM

extra shelter afforded by the seaward mangrove
community, together with the decreased period
and extent of tidal inundation in the sedgeland
areas as compared to the mangroves, may have
allowed nests with no external structural
framework, i.e. free-standing mounds, to be built.

Curiously, none of the nests encountered
during this study closely matched the description
of the original X. myoides nest structure from
Melville Island provided by Magnusson et al.
(1976). It involved a 60cm-high mud mound,
containing a nesting chamber, constructed at
ground level against the trunk of a living
mangrove (Bruguiera parviflora) tree. The nest
from southeast Queensland that most closely
resembled this Northern Territory example was
tree trunk nest #8 1 from Coomera River in which
a 42cm-high mud mound was built against the
base of a living Avicennia marina. Other tree
trunk nests with smaller mud mounds were also
discovered. However, given the limited size of
these mounds, as well as the position of the trees
in the littoral zone, the mounds associated with
these nests would have been entirely inundated
during high tide and so could not serve as nests in
their own right. Rather, the purpose of these
ancillary mounds may either have been to
provide secure access through mud tunnels to the
nest proper within the adjacent tree or to serve as
a buffer against the tide and thereby prevent mud
or organic matter within the tree from washing
out through any ground level hole in the trunk.
Both explanations may be true. Alternatively, it is
interesting to speculate about whether these small
mounds built alongside tree nests serve any
useful function at all. Because natural selection
will have favoured those X. myoides individuals
that build nest structures above the height of
spring tides (the dominant external factor that
ultimately must govern nest heights in any
region), perhaps animals construct mounds even
in situations in which they are not required. Based
on our hypothesis about the requirements for nest
construction to begin within the mangrove zone,
we would assume that the B. parviflora adjacent
to the nest structure described by Magnusson et
al. (1976) was hollow and so able to afford the
Water Mice individuals refuge during the
mound-building phase.

Trapping and radio-tracking studies have
revealed that X myoides regularly follows the
receding tide out into the mangrove zone where it
feeds until rising water forces it back to the
shelter of its nest site (Van Dyck, 1997). This
apparent preference for foraging among

mangroves suggests this is where food resources
for the species occur in the highest densities. This
conclusion is supported by studies showing that
substrate-dwelling fauna of the mangrove zone is
richest in species at the lower tide levels where
regular inundation by tides occurs (McCormick,
1978). On first appearances, it would appear
sensible for X. myoides to nest within the
mangrove zone but additional factors come into
play. Of all the vegetation communities in the
littoral zone, the resource rich mangrove
community is the first to be inundated on the
flooding tide, and the last to be exposed when the
water recedes. Furthermore, the depth of
inundation is greater there than anywhere else in
the intertidal zone. As a consequence, the time
available for a mangrove-nesting X. myoides to
forage between tides is more limited. We suggest
that, overall, nest location, and therefore nest
type is a resultant compromise between prox-
imity to the most productive resources of the
mangrove zone and a suite of complicating
factors, namely the difficulty (or practicality) of
nest building in a regularly flooded site, the ability
of the nest to withstand tides, particularly spring
tides, and the period available for foraging.

In addition to proximity to rich food resources,
what other selective advantages may offset the
effort in constructing and maintaining large
mounds or tree trunk nests within or as close as
possible to the productive mangrove zone as
opposed to digging simple tunnels into the
supralittoral bank near the woodlands and
wallum? One possibility is that offspring survival
might be higher in complex nest structures
attached to high-yielding foraging areas as a
consequence of the cumulative effect of home
range defence by related adults. This is supported
by the incidence of agonistic encounters
witnessed between individuals away from nest
sites (SVD, pers. obs.). Bank nests, unlike other
nest types out in the intertidal zone, are not fully
moated at high tide and so may offer less
protection from snakes and more opportunities
for disruptive intrusion from conspecifics.
Additional disturbance may be caused by the
foraging activity of Rattus lutreolus and predation
by R. rattus and Hydromys chrvsogaster. The
evening journey between the supralittoral bank
and foraging areas within the mangroves may
also produce higher losses to nocturnal raptors,
Red Foxes Vulpes vulpes and Cats Felis catus.
Finally, fire, the dramatic event that revitalises
wallum and coastal woodland by triggering
germination, may select those mice that nest far

NEST TYPES OF XEROMYS MYOIDES

479

from its influence. In this regard we have direct
evidence in nests #33 and #46 (Donnybrook) that
wild fires can quickly convert supralittoral bank
nests and adjacent spoil heap nests to peat ash.

ACKNOWLEDGEMENTS

In addition to those acknowledged in the North
Stradbroke Island study (Van Dyck, 1997) we
thank Peter and Carol Lehmann (Pimpama) for
all their help and pikelets, Glenn Leiper and staff
(Jacobs Well Environmental Education Centre)
for accommodation and assistance, Steve Price
(Queensland Parks and Wildlife Service) for
alerting us to the impacts of a fire at Donnybrook
and for showing us so many Pumicestone
Passage nests, CSR Limited for permission to
enter their Donnybrook property, and Austcorp
International Ltd for permission to carry out
research on their Coomera property. We are also
most grateful to Ross Patterson, Sarah Parker,
Scott Rogers, Don Cameron and Brendan
McLarty (all Queensland Parks and Wildlife
Service) and Heather Janetzki, Andrew Baker,
Tamya Cox and Angela Frost (all Queensland
Museum) for their information about nest
locations and valuable assistance in the field. The
mighty efforts of Harriet Preece (Queensland
Parks and Wildlife Service) in preparing the map
used in Fig. 1 were greatly appreciated.

LITERATURE CITED

BROOKS, J.H. 1953. Stradbroke Island erosion and
Broadwater silting Southport. Queensland
Government Mining Journal, August, 1953:
566-569.

BURT, W.H. & GROSSENHEIDER, R.P. 1976. Afield
guide to the mammals of America north of
Mexico. (Houghton Mifflin Company: Boston).

CLIFFORD, ITT. & SPECHT, R.L. 1979. The vegetat-
ion of North Stradbroke Island, Queensland.
(University of Queensland Press: Brisbane).

CONNAH, T.H. 1946. Stradbroke Island erosion and
Broadwater silting, Southport. Queensland
Government Mining Journal, December, 1946:
370-373.

DOWLING, R.M. 1986. The mangrove vegetation of
Moreton Bay. Queensland Botany Bulletin 6.
(Queensland Department of Primary Industries:
Brisbane).

MAGNUSSON, W.E., WEBB, GJ.W. & TAYLOR,
J.A. 1976. Two new locality records, a new habitat
and a nest description for Xeromys myoides
Thomas (Rodentia: Muridae). Australian Wildlife
Research 3: 153-57.

McCORMICK, W.A. 1978. The ecology of benthic
macrofauna in New South Wales mangrove
swamps. (Unpubl. MSc thesis, University of New
South Wales: Sydney).

VAN DYCK, S.M. 1992. Parting the reeds on Myora’s
Xeromys kibbutz. Wildlife Australia 29(1): 8-10.

1997. Xeromys myoides Thomas, 1889 (Rodentia:
Muridae) in mangrove communities of North
Stradbroke Island, southeast Queensland.
Memoirs of the Queensland Museum 42(1):
337-66.

VAN DYCK, S.M. & DURBIDGE, E. 1992. A nesting
community of False water rats Xeromys myoides
on the Myora sedgelands of North Stradbroke
Island. Memoirs of the Queensland Museum
32(1): 374.

WALKER, E.P. 1964. Mammals of the world. Volume
2. (The Johns Hopkins Press: Baltimore).

480

MEMOIRS OF THE QUEENSLAND MUSEUM

ARTIFICIAL NESTING MOUNDS FOR THE WATER
MOUSE, XEROMYS MYOIDES. Memoirs of the
Queensland Museum 49(1): 480, 2003:- In intertidal
communities of southeastern Queensland the Water Mouse
Xeromys myoides constructs nests of five basic types: 1)
free-standing mud mounds within areas of sedgeland,
chenopod shrubland. Marine Couch Sporobolus virginicus
grassland or mangrove forest, 2) mounds associated with
small, slightly elevated ‘islands' standing away from the
supralittoral bank; 3) mounds or holes in the supralittoral
bank; 4) mud and leaf constructions inside hollow tree trunks;
and 5) nests made in spoil heaps of human origin (Van Dyck
& Gynther, 2003). Ultimately, the locations (and, therefore,
type) of nests constructed by these carnivorous rodents
appears to be a compromise between a nest’s ability to
withstand spring tides and its proximity to the most highly
productive resources of the mangrove zone.

Inevitably, some habitats that otherwise would appear
suitable for Water Mice either offer insufficient elevation
above the surrounding intertidal community or lack suitable
hollow tree trunks to allow nest construction and breeding to
proceed without inundation. Given the use by X. myoides of
human-generated spoil heaps for nesting (excavated material,
tree stump waste, bulldozed soil, gravel rock and concrete:
Van Dyck & Gynther, 2003), the provision of artificial nesting
mounds might represent a useful management tool to
encourage this vulnerable species to colonise new sites.

From 5 May - 24 June 1999, 12 artificial nesting mounds
were installed on intertidal wetland (centre at 27°50.857*S,
153°22.470’E) close to the Coomera River, southeastern
Queensland. The site, although relatively close to an
established X. myoides population (approximately 400m
northwest), presented few natural nesting opportunities. It
was characterised by extensive S. virginicus grassland flats, a
relatively immature mangrove community, a small Swamp
Oak Casuarina glauca component and an elevation
throughout below the 1 m contour.

Each artificial nest installation consisted of four copper
napthenate-treated hardwood stakes (70mm x 50mm x 2m)
set into the comers of a rough, 1 m square and driven into the
substrate until about 1 .5m of stake remained above ground.
Inside the stakes, a cylinder of heavy duty plastic garden
trellis (1.2m high, 75mm square lattice pattern) was secured
with tie wires. For mouse access, a
length of 1 00mm diameter agricultural
pipe with a number of 40mm diameter
holes cut along its length was laid on the
ground across the middle of the
cylinder, each end protruding from the
trellis material. (This pipe was later
deemed unnecessary see below.)

Commercial potting mix was then
tipped in to form a central core of earth
and commercial mulch (chipped tree
material) was packed around and on top
of it to fill the cylinder up to about lm
high (Fig. 1A).

Three artificial mounds were
installed among mangroves near the
river’s edge. Another was positioned on
a degraded mud flat, two were installed
on low S. virginicus/C. glauca islands
and the remainder (six) were erected in
the open 5. virginicus grassland.

Initially, the artificial mounds were
monitored regularly, and as early as 2
June 1999 (three weeks after
installation) dismembered crab remains
were found on top of two mounds. On 5

August 1999 rodent droppings w'ere retrieved from one
mound and later identified as Rattus sp. (B. Triggs pers.
comm.). By this time it was apparent that all but one mound
had been completely inundated by spring tides, and many
needed topping up. In particular, the two near the river's edge
were rapidly eroding. By 14 October 1999. more droppings
were retrieved from another mound, but the occurrence
nearby of chewed leaf galls from the mangrove Avicennia
marina indicated Rattus rattus was probably involved.
Thereafter, the artificial nests were left unchecked until
December 2002, when all 12 mounds were revisited. In all but
one. the entire mulch pile inside the trellis cylinders had been
w'ashed away. However, one cylinder, installed on a small S.
virginicus ‘island' (27°50.808’S, 1 53°22.479'E) contained a
functional X. myoides nest (Fig. IB). The total nest height
(nest mound plus 20cm mud bank) was 47cm, mound circum-
ference was 235cm and three access holes were incorporated
into the structure; two at ground level and one at 20cm (at
mound base). The mounding material w'as mostly sand and
mud but clearly contained traces of the original soil and mulch
provided. Much of the surface of the mound was consolidated
by S. virginicus , and, as is typical of this rodent, small claws
and carapaces of crabs (here, the Mottled Shore Crab
Paragrapsus taevis ) w ere incorporated into the external mud
daubing. The agricultural pipe access was completely
blocked with mud.

The provision of such nesting frames, furnished instead
with more robust trellis and a much less friable soil, may not
only expand the area of habitat available to the species but
may become increasingly important at sites where foxes and
feral pigs are a potential threat to the nesting success or
existence of local A', myoides populations.

The financial assistance of Austcorp International Limited
is gratefully acknowledged.

Literature Cited

VAN DYCK, S. & GYNTHER, l, 2003. Nesting strategies of the

Water Mouse Xeromys myoides in southeast Queensland.

Memoirs of the Queensland Museum, this volume.

S. V an Dyck & H. Janetzki, Queensland Museum, PO Box
3300, South Brisbane 4101; I. Gynther, Queensland Parks
and Wildlife Sendee, PO Box 64, Bellhowrie 4070, Australia
6 May 2003.

DIAB OLICORIS, A NEW GENUS IN THE TRIBE PLOIARIOLINI (HEMIPTERA:
REDUVIIDAE: EMESINAE) FROM NEW CALEDONIA

M.A. WALL AND G CASSIS

Wall, M. A. & Cassis, G 2003 06 30: Diabolicoris , a new genus in the tribe Ploiariolini
(Hcmiptera: Reduviidae: Emesinae) from New Caledonia. Memoirs of the Queensland
Museum 49(1): 481-483. Brisbane. ISSN 0079-8835.

Diabolicoris saicelloides new genus and new species is described and placed in the emesine
tribe Ploiariolini. The species is known from a single specimen collected in New Caledonia.
Diabolicoris saicelloides shows close affinities to Saicella Usinger, a flightless genus
endemic to the Hawaiian Islands. □ thread-legged bugs, assassin bugs, Diabolicoris
saicelloides, new species, Reduviidae, New Caledonia .

Michael A. Wall, Department of Ecology r and Evolutionary Biology, University of
Connecticut, Storrs, USA; Gerasimos Cassis, Centre for Biodiversity and Conservation
Research, Australian Museum, Sydney 2000, Australia; 3 November 2002.

The Ploiariolini (Hemiptera: Reduviidae:
Emesinae) is supported by several synapomorphic
characters in the wings, as well as those in the
male and female genitalia (Wygodzinsky, 1966).
While apparently monophyletic, the position of
the Ploiariolini within the Emesinae is ambiguous.
Wygodzinsky (1966) considers the Ploiariolini
unplaced in the Emesinae, but tentatively allied it
with the Emesini, Metapterini and Deliastini.
Based on its pronotal glands, we suggest the
Ploiariolini may be allied with the emesine tribe
Collartidini and the subfamily Saicinae. Further-
more, Saicella Usinger, a flightless ploiarioline
endemic to the Hawaiian Islands, shares several
characters with the Saicinae (e.g. elongate spines
on head, labium, procoxae and profemora;
procoxae relatively short; and anterior acetabula
sometimes directed ventrally) and the Ploiariolini
(Wygodzinsky, 1966; Polhemus, 2000). Until now,
Saicella has been the only ploiarioline known to
possess numerous transitional characters. Here
we describe a new genus of Ploiariolini from
New Caledonia, Diabolicoris , that has many of
the same transitional characters.

Diabolicoris Wall & Cassis gen. nov.

(Fig. 1)

TYPE SPECIES. Diabolicoris saicelloides Wall & Cassis.

ETYMOLOGY. Latin for devil and bug, due to general
diabolically spiny appearance of the bug.

DIAGNOSIS. Similar in appearance to Saicella ,
but with spines on trochanter of prolegs and
spines absent from procoxae. Also, forewings
with triangular subbasal cell, and discal cell
distally not lobed.

DESCRIPTION. Macropterous. Body shining,
moderately covered in golden flexuous setae. No
woolly setae present.

Head short; distinctly divided into anteocular
and postocular portions; anteocular region
roughly parallel-sided dorsally; postocular
region slightly shorter than anteocular, globular
both laterally and dorsally with shallow sulcus
dorsally, as wide as combined width of eyes and
anteocular region; eyes small, set laterally, well
below dorsum of head; antenniferous tubules
large;ocelli absent; buccula with single posterior
spine-like seta; gena with three spine-like setae;
labium 3-segmented, segment I subcylindrical
with 2 + 2 spine-like setae, equal in length to
segment II and III combined, segment II inflated
with 3 + 3 setae.

Pronotum covering mesonotum, anterior and
posterior lobes present, deeply constricted before
midline; anterior lobe subglobular, deeply
divided into two elevated lobes by deep median
longitudinal sulcus, ostiole of pronotal gland
present on sulcus; anterior acetabula directed
slightly forward; posterior lobe subquadrate,
lateral margins weakly divergent posteriorly.
Scutellum barely exposed, semicircular, slightly
raised medially. Metanotum as long as hind lobe
of pronotum, with a short, stout, apical spine.

Forewings broadly rounded distally with small
notch at M and wing margin. Discal cell large,
truncate proximally, curved apically. M and Cu
fused basally of discal cell forming a triangular
subbasal cell. Pterostigma narrow, short, barely
surpassing the costal margin of the discal cell.
Hind wings present.

Prolegs stout; coxae simple, less than 4 X
longer that wide; trochanters armed with two

482

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1 . Lateral view of holotype of Diabolicoris saicelloides with selected detail. A, flexuous setae of head and
pronotum; B, distally oriented stiff setae of meso- and metalegs; C, areolated pattern elements of forewing; D,
appressed setae of abdomen; E, capitate setae covering female genitalia.

obvious spines set on tuberculate bases; femora
with row of five enlarged elongate spines set on
large tuberculate bases along exterior lateral face,
length of longest spine half as long as femur, also
with row of six slightly smaller spines along
interior lateral surface, angle between interior
and exterior rows of spines greater than 90°, distal
third of femur with minute dark papillae; tibia
with two parallel rows of numerous spinelike
setae on ventral surface; tarsus 2-segmented,
second segment 2x longer than first; paired
claws minute. Middle and hind legs slender with
distally angled stiff setae, tarsi 3-segmented, first
segment tapered distally, shorter than other
segments, junction of second and third segment
difficult to discern.

Abdomen broadly attached to thorax, elliptical;
connexival margins curved inward; first visible
stemite subquadrate, posterior margin shallowly
concave, spiracle distinctly stalked.

REMARKS. Diabolicoris represents a unique
combination of character states, some shared

w ith Saicella , others with Emesopsis Uhler, and
others are possessed by all three genera. Diaboli-
coris resembles the Hawaiian endemic genus,
Saicella , in the possession of elongate spines on
the profemora, labium, and venter of the head.
Unlike Saicella , Diabolicoris is macroptcrous.
The proximally truncated discal cell of the
forewing allies Diabolicoris with Emesopsis and
related genera. Wygodzinsky (1966) concluded
that Saicella and Emesopsis are closely related
based on characters associated with the phallus,
vestiture, and abdomen. Although the question
bears further phylogenetic analysis, Diabolicoris
appears closely related to the Saicella-Emesopsis
clade proposed by Wygodzinsky (1966).

DISTRIBUTION. Currently only known from
the type locality in southern New Caledonia.

Diabolicoris saicelloides Wall & Cassis sp. nov.
(Fig. 1)

MATERIAL. Holotype. 9 , NEW CALEDONIA 22 14 S
166 52 E, Chute Madeleine, 230m, 12 November 2000,

NEW ASSASSIN BUG

483

GB Monteith, pyrethrum, trunks and logs, 9917. (Museum
National d’Histoire Naturelle - Paris; don no. 1097 du
23.05.2003)

ETYMOLOGY. Named for the genus Saicella which it
resembles.

DESCRIPTION. (All measurements in mm).
Length: 2.4. Ground color aureous to stramineous;
antenniferous tubercles and tarsi white; antennae
testaceous; meso- and metalegs becoming
testaceous distally. Wing with conspicuous
testaceous pattern elements.

Head length/width: 0.48/0.28; covered with
appressed and upright flexuous golden setae;
eyes small, elliptical, longest along dorsoventral
axis, approximately 25 ommatidia each; antennal
segments length: 1.14/0.84/0.48/0.32, first
segment widest distally.

Pronotum length/width: 0.42/0.36, procoxae
length/width: 0.28/0.08, profemora length: 0.62,
protibiae length: 0.46, protarsi length: 0.12,
mesofemora length: 1.28, mesotibiae length:
1.66, mesotarsi length: 0.14, metalemora length:
1.74, metatibiae length: 2.44, metatarsi length:
0.14, and forewing length: 2.14.

Abdomen elliptic in dorsal view, length: 1.5;
external female genitalia obscured by thick
hyaline capitate hairs; male unknown.

DISTRIBUTION. Known only from Chute
Madeleine, New Caledonia. Although this area is
primarily dry/sclerophyllous maquis-type
vegetation, Diabolicoris saicelloides was
collected from tree trunks in a nearby pocket of
rainforest. Subsequent pyrethrum collecting at
the same location by the same collector in
November 2002 failed to yield more specimens.

ACKNOWLEDGEMENTS

Sincere thanks to Geoff Monteith of the
Queensland Museum, who collected and loaned
this material to us. This project was completed
while the senior author was in Australia on a U.S.
Department of State Fulbright Fellowship and
supported by the Australian-American Fulbright
Commission. The Australian Museum Trust is
thanked for supporting this work.

LITERATURE CITED

POLHEMUS, D.A. 2000. A revision of the endemic
Hawaiian reduviid genus Saicella Usinger, with
descriptions of four new species (Heteroptera:
Reduviidae: Emesinae). Proceedings of the
Entomological Society of Washington. 100: 1-20.
WYGODZ1NSKY, P. 1966. A monograph of the
Emesinae (Reduviidae, Hemiptera). Bulletin of
the American Museum Natural History. 133:
1-614.

MEMOIRS OF THE QUEENSLAND MUSEUM

485

VOCALISATION BY THE COMMON EASTERN
AUSTRALIAN GECKO, GEHYRA DUBI A. Memoirs of
the Queensland Museum 49( I): 485-6. 2003:- Information on
vocalisation in Australian geckos is scant. A handful of native
species are known to call and one introduced to Australia, the
house gecko, Hemidactylus frenatus Dumcril & Bibron,
1836, has a loud distinctive call. Most calls reported for
Australian geckos have been related to fighting, {e.g.
Diplodactylus Bustard, 1965; Gehyra - Bustard, 1969;
Frankenberg & Werner, 1984), or confrontation with a
predator (e.g. Christinas Daniels et al., 1986; Nephrurus
asper - Bustard, 1967; Greer pers. obs M 1989; Phyllurus
platurus - Greer pers. obs., 1989). On Boyne Island, in
Central Queensland, Gehyra dubia Macleay. 1877 is
commonly found in acacia, eucalyptus woodlands and the
surrounding urban areas. This gecko is usually arboreal and is
frequently observed on the walls of houses, both in and
outside, and other domestic structures.

During late September, 1 997, 1 recorded the calls produced
by Gehyra dubia specimens for the first time. This was
achieved by holding 4 captive G. dubia (sample 1,3$ , 1 <$) in
two small, transparent containers. Two geckos were placed in
each container and microphones were secured to the lids. The
outputs from these two microphones were connected to a
stereo cassette recorder. At night, between the 23-27/9/97, a
scries of !4hr recordings was made. This was followed by
recordings during late October 1 997 (sample 2-26,29). mid
February' 1998 (sample 3- 26 ), late January 2001 (sample 4-
26), and late September 2001 (sample 5- 6 , $). Sample 6,
early June 2003 (4$, 26), was not recorded, but closely
observed to establish if both sexes produced all three calls. Four
$ $ were placed in one container, and the 2 6 6 in another.

Examination of these recordings and my observations
between 1989-2003 indicate that Gehyra dubia has at least
three distinct calls: a distress call, a chirp and a fast chatter.
These recordings have been digitised (AWE Sound Blaster
Card; Sampling rate, 44,100, 22,050, 1 1,025; sampling size
16 bit), and reproduced into audiographs (Fig. 1A-C). This
has enabled accurate measurements of the call duration (in
seconds) and a comparison of the amplitude of pulses within a
call. Individual pulses and number of pulses per call were
verified by reducing the play-back sampling rate to 1 1 ,025Hz.
The dominant frequency band of individual pulses (the
frequency at which most sound energy is transmitted), and
pulse width was achieved by using a freeware dual channel
audio spectrum analyser (Gram 50 program). This provided a
scrolling time-frequency display (Fig. 2A-C). Only
recordings of high resolution were used for frequency
analysis and pulse width measurements.

Although these recordings are of the calls of captive
geckos, calls are consistent with those heard in the wild.
Reasons for these vocalisations are unknown. It was possible
to distinguish between the sexes of adults by examining the
neck and cloacal regions of each specimen: males lack
obvious endolymphatic calcium stores; have distinct prcanal
pores aligned in an inverted ‘V’; distinct postcloacal sacs and
enlarged postanal tubercles. Females have distinct
endolymphatic calcium stores, moderate postanal tubercles,
but lack prcanal pores and expanded postcloacal sacs.

Gehyra dubia calls

Type 1 - distress: when fighting, specimens often release a
one pulse clamor which is louder and more drawn out than
other calls. This was not recorded, but has been displayed by
both sexes (sample 6).

Type 2 - the chirp (eee...eee...eec) has been heard
frequently. Both 6 6 & 99 produce this call (sample 6).
However, it has been possible to record this call on only three
occasions (CH 1 , CH 1 a and CHFC 1 ). This call is displayed as

a single pulse, ora series of chirps with long intervals between
pulses. Recording CH 1 (Fig. 1 A ), 2 9 $ , consisted of 6 pulses/

1 1 ,2s; recording CH 1 a (2 9 9 ), is of a single pulse; recording
CHFC1 (Fig. IB), 6 & 9, multiple chirp portion, 4 pulses/
5,6s. Estimated, dominant frequency band of individual pulses
varies between 2.6-3. 8Khz. The pulse width is between
140-228 milliseconds. Pulses peter out at 6.3-8. 7KHz.

Type 3 - the fast chatter (ee.ee.ee) is, by far, the most
frequently heard, and has been recorded in FC I -11. Adult 6 6
often exhibit this call (Table 1).

Total pulses per call ranged from 9-23. Pulses per second
(pulse rate) varied from 6-8.3. The call duration varied from
1.2-3. 6s. Pulse width range 72- 167ms. Dominant frequency
range 2.6- 3.8kHz. The first pulse of recordings FC 2,5,8,10
and 1 1 varied between 72- 125ms and have a lower frequency
range than proceeding pulses. Pulses peter out at 3.7-7kHz.

An analysis of these recordings has established that 6 6
produce the fast chatter (FC5-FC10). 9 9 have produced the
chirp (CHI and CHI a). When the combination of 6 6 and
9 9 was used for a recording, I was unable to ascertain which
gecko called.

Conclusion

Gehyra dubia does frequently call. The calls heard and/ or
analysed can be categorised into three distinct groups: 1 ) the
distress, 2) the chirp, and 3) the fast chatter. Both sexes exhibit
the distress call. The chirp call has also been produced by both
6 6 & 9 9. Pulses of the chirp call are often long (up to
228ms), lack intensity, and are exhibited as a single pulse or a
series of pulses with long interv als between The fast chatter
has been strongly exhibited by 6 6 only, and is frequently
heard throughout my house. The amplitude of these pulses is
high, and typically intensifies in the latter portion of the call
(Fig. 1C). Pulses of the fast chatter are short (< 167ms), and
are carried out over a short period (<3.6s). Estimated
dominant frequencies of the chirp and fast chatter calls are
quite similar (2.6-3. 8kHz). Differentiation between both types
of calls includes, pulse rate, pulse width and call intensity.

Only one recording consisted of both the chirp and fast
chatter calls - CHFC 1 , 6 & 9, sample 1 (four chirps followed
instantly by 1 7 fast chatter pulses. Fig. 1 B). By comparison to
other recordings (CH 1 , CH 1 a and FC5- 1 0), and the very short
transition period from chirps to chatter, it is quite reasonable
to suggest that the call of recording CHFC1 consists of a 9
followed by a 6 . However, this would need to be verified by a
higher sample rate.

The reasons for these calls are unknown. However, it does
not seem unreasonable to speculate that they relate to sex rec-
ognition and territory definition and defence as has been
postulated for many other animal calls.

Acknowledgements

For assistance in preparing this paper, thanks are due to
Jeanette Covacevich for advice on the manuscript; Dr Mark
Hutchinson (South Australian Museum) and Peter Robertson
(Wildlife Profiles) both imparted their knowledge on sexual
dimorphism within the gecko family; Jon Butterworth
(Electrical Engineer, Queensland Cement Ltd) for guidance
on the Gram 50 program and analysis on sound waves; Assoc.
Prof. Ross Alford and Dr Lin Schwarzkopf of James Cook
University for forwarding the Gram 50 program and sharing
their knowledge on the analysis and identification of
biological sounds, Special thanks to my wife Inge for
tolerating the disturbances and inconvenience of such a study.
John Toop of the Department of Environment and Heritage
provided study permits.

Literature Cited

BUSTARD, H.R. 1 965. The systematic status of the Australian geckos
Gehvra variegata (Dumeril and Bibron, 1836) and Gehyra

MEMOIRS OF THE QUEENSLAND MUSEUM

486

A

I { !

1 , ?

i

_J.il illM.

B

1 1 1 1 1 1

^+trti

c

Ij J

FIG. 1 . Audiographs. A, CH 1 , sample 1,299,6 pulses, call
duration - 11.2s. B, CHFC1, sample 1, 6 & 9, call
duration ~8.2s. C, FC10, sample 4, 2d 6, 21 pulses, call
duration -3. 6s.

australis (Gray, 1845). Herpctologia 20(4): 259-72.

1967. Defensive display behaviour of the Australian gecko
Nephrurus asper. Herpetologia 23(2): 126-29.

1969. The population ecology of the gckkonid lizard Gehyra
variegata (Dumcril and Bibron) in exploited forests in northern
New South Wales. Journal of Animal Ecology 38: 35-51

DANIELS, C.B., FLAHERTY, S.P. & SIMBOTWE, M.P. 1986. Tail
size and effectiveness of autotomy in a lizard. Journal of
Herpetology 20(1): 93-96.

FRANKENBERG, E. & WERNER, Y.L. 1984. The defensive vocal
“distress*’ repertoire of gekkonid lizards: intra-and-
inter-specific variation. Amphibia-Rcptilia 5(2): 109-24.

GREER, A. 1989. The biology and evolution of Australian lizards.
(Surrey Beatty & Sons: Chipping Norton).

Peter R. Tremul, 5 Kilman Court, Boyne Island 4680,

Australia ; 10 June 2003.

FIG. 2. Spectrographs. A, FC10, sample 4, 26 6, pulses
14-19, dominant frequency of pulses -2. 6kHz. B, CHI,
sample 1,299, pulses 1 -3, dominant frequency of pulses
-3.8, 2.6 & 2.6kHz. C, CHFC1, sample 1, 6 & 9, pulses
4-20, chirp and chatter transition, dominant frequency of
pulses -3.8kHz. Darker shades indicate higher decibel
readings.

TABLE 1. Recording data. LR, low resolution.

Rec.

No.

Sex &
sample No.

Pulses/

call

duration

( sec )

Pulses/

sec.

Pulse

width

(msec.)

Estimated

dominant

frequency

(kHz)

FC1

6 & 9, si

12/1.7

7

87-133

3.0

FC2

6 & 9, si

9/1.5

6

87-156

2.6

FC3

6 & 9, si

11/1.5

7.3

LR

LR

FC4

d & 9, si

17/2.5

6.8

LR

LR

FC5

2 d, s3

10/1.2

8.3

76-114

3.8

FC6

2 d, s3

18/2.2

8.2

LR

3.1

FC7

2 d, s3

23/2.9

7.9

LR

LR

FC8

2 d, s4

19/3.4

5.6

72-167

3.8

FC9

2 d, s4

20/3.4

5.9

76-167

2.6

FC10

2 d, s4

21/3.6

5.8

125-156

2.6

FC1 1

d & 9,s5

18/3.3

5.5

87-163

3.6

CONTENTS (continued)

FREEMAN, A.B.

An observation of calling northern tinker frogs on Mount Bellenden Ker 295

HEATWOLE, H., DE BAVAY, J. & WEBBER, P.

Faunal survey of New England. V. The lizards and snakes 299

HOCKNULL, S.A.

Etnabatrachus maximus gen. et sp. nov., a Plio-Pleistocene frog from Mount Etna, central

eastern Queensland * ^27

JOHNSON, P.T.J. & RITCHIE, E.G. „

Macroinvertebrate fauna of an iron-rich stream in the Wet Tropics of Australia: a comparative

analysis of communities using a rapid bioassessment protocol 331

KOFRON, C.P. . ^ ™

Case histories of attacks by the Southern Cassowary in Queensland jj?

LEE, A.C. & ARANGO, C.P. . , , _ , . .

Two new species and other records of sea spiders (Pycnogonida: Arthropoda) from tropical

North Queensland * 343

LIMPUS, C.J., MILLER, J.D., PARMENTER.C. J. & LIMPUS, D.J.

The Green Turtle, Chelonia my das, population of Raine Island and the northern Great Barrier

Reef: 1843-2001 349

Aoupinia , a remarkable new genus of Adeliini from New Caledonia (Coleoptera. Tcnebriomdae)

SUZUKI, H. & DAVIE, P.J.F. ^ _ ., .... .

A new cavemicolous shrimp in Pycnisia Bruce, 1992 (Crustacea: Decapoda: Candea. Atyidae)

from northwestern Queensland

VAN DYCK, S. & GYNTHER, I. M

Nesting strategies of the Water Mouse Xeromys myoides in southeast Queensland

WALL, M.A. & CASSIS, G.

Diabolicoris, a new genus in the tribe Ploiariolini (Hemiptera: Rcduvndae: Emesinae) from
New Caledonia

NOTES

RIX ’ M '° Blastobasine coleophorid moths as prey for the Australian araneid spider Celaenia calotoides Rain-
bow

HARDING, D. & WILLIAMSON, I. _ t „ ... . . , ocf<ir „

A note on the habitat requirements of the swamp crayfish on Bnbie Island, southeastern ^

Queensland

VAN DYCK, S., JANETZK1, H. & GYNTHER, I. . , 4 o 0

Artificial nesting mounds for the Water Mouse Xeromys myoides

TREMUL, P.R. , * » • 485

Vocalisation by the common eastern Australian gecko, G enyra

CONTENTS

BAEHR, B.

Three new endemic genera of the Asteron-comp\cx (Arancae: Zodariidae) from Australia:

Basasteron, Euasteron and Spinasteron 1

BAEHR, B.

Tropasteron gen. nov. of the Asteron - complex (Araneae: Zodariidae) from tropical Queensland . . 29
BAEHR, M.

Psydrine ground beetles (Coleoptera: Carabidae: Psydrinae), excluding Amblytelini, of eastern
Queensland rainforests 65

BRUCE, AJ.

Further information on two pontoniine shrimps from ascidian hosts, Dasella brucei Berggren,

1990 and Pseudopontonia minuta (Baker, 1907) (Crustacea: Decapoda: Palaemonidae) ... Ill

BRUCE, A.J.

Periclimenes species (Crustacea: Decapoda: Pontoniinae) from far north Queensland 115

CASSIS, G., SCHWARTZ, M.D. & MOULDS, T.

Systematics and new taxa of the Vannius complex (Hemiptera: Miridae: Cylapinae) from the

Australian region 123

CASTRO, P. & DAVIE, P,

New records of palicid crabs (Crustacea: Brachyura, Palicidae) from Australia 153

CATERINO, M.S.

New species of Chlamydopsis (Histeridae: Chlamydopsinae), with a review and phylogenetic

analysis of all known species 159

DAVIES, V.T.

Barahna , a new spider genus from eastern Australia (Araneae: Amaurobioidea) 237

DAVIES, V.T.

Tangana , a new spider genus from Australia (Amaurobioidea: Amphinectidae: Tasmarubriinae)

Miocene palynofloras from subsurface sediments in the Bundaberg district 261

EDGECOMBE, G.D.

A new genus of henicopid centipede (Chilopoda: Lithobiomorpha) from New Caledonia 269

FAIRFAX, R J. & FENSHAM, R.J.

Great Artesian Basin springs in southern Queensland 191 1-2000 285