Records of the Australian Museum

RECORDS OF THE
AUSTRALIAN MUSEUM

Volume 62 Numbers 2 & 3 24 November 2010

Uncovering local endemism in the Kimberley, Western Australia: description of new

species of the genus Amplirhagada Iredale, 1933 (Pulmonata: Camaenidae).

.Frank Kohler 217

A revision of the Australian funnel-web spiders (Hexathelidae: Atracinae).

.Michael R. Gray 285

Note on Hydroides malleolaspinus from the Kimberleys of Western Australia (Polychaeta:

Serpulidae). Anna Murray, Pat Hutchings and T. Gottfried Pillai 393

Contents Volume 62 . 395

Australian

museum

nature culture discover

Journal compilation © 2010 Australian Museum
The Australian Museum, Sydney

No part of this publication may be reproduced without
permission of The Editor.

Volume 62 Numbers 2 & 3

Published 24 November 2010

Price: AU$60.00

Printed byRodenPrint Pty Ltd, Sydney

ISSN 0067-1975

The Australian Museum is a statutory authority of,
and principally funded by, the NSW State Government.

NSW

GOVERNMENT

Communities

Cover images —Comparison is an essential aspect of discovery
in the natural sciences. In the example shown on the cover, the
skulls of two indigenous rodent species are depicted. Australian
Museum zoologist Troughton (1923, Records of the Australian
Museum 14[1]: 23-41) determined them to belong to the
species Leporillus conditor (Gould) (Figs. lc,d) and L. jonesi
Thomas (Figs. 2c,d). He noted that the genus, at that time, was
represented by species from west of the Lower Darling River in
New South Wales, from Alice Springs to the Nullarbor Plains,
and on islands of the Nuyts Archipelago in South Australia.
Their large stick nests had been noted by early explorers Sturt
and Mitchell and figured by Gould.

Since 1999 the primary scientific literature published in print by
the Australian Museum has also been freely accessible online at our
website. The entire legacy of Records of the Australian Museum , back
to volume 1 of 1889, has now been digitized and is freely available
online. The above work by Troughton can be read in its entirety at
http://dx.doi.Org/10.3853/j.0067-1975T4.1923.824

Records of the Australian Museum is covered in the Thomson
Scientific services: Current Contents ® / Agriculture, Biology,
and Environmental Sciences, and Science Citation Index
Expanded (also known as SciSearch ® )

* We encourage cross-linking in the scientific literature by
applying doi registration to Australian Museum publications via
CrossRef ®

The Australian Museum houses some of the world’s most important
collections of Australian animal, fossil and geological specimens
and cultural objects. Research on these millions of specimens and
artefacts yields insights into how our world changes through time and
how its diversity can be classified and interpreted. This knowledge,
when shared among the scientific and broader community—initially
through publication—helps us understand human impact on our
environment and what reasonable steps society can take now for
the well-being of future generations. Our responsibility is to inspire
the exploration of nature and cultures; our vision is a beautiful and
sustainable natural world with vibrant and diverse cultures.

Since 1889 the Records of the Australian Museum (ISSN 0067-
1975) has published the results of studies that derive in large
part from Australian Museum collections or studies that more
generally lead to a better understanding of nature and cultures in
the Australasian region. Issues of the Records are published three
times a year and circulated to 84 countries. All that is published in
print is, soon afterwards, also freely available online. Records of the
Australian Museum, volume 60, was published in 2008, volume 61
in 2009. Monographic works of particular significance are published
irregularly as Records of the Australian Museum, Supplements.
Catalogues and checklists have, in the past, been published in print as
numbered Technical Reports of the Australian Museum (ISSN 1031-
8062 print). From number 20 (December 2007) onwards, Technical
Reports of the Australian Museum, online (ISSN 1835-4211 online)
has been published online only. Australian Museum Memoirs (ISSN
0067-1967) ceased in 1983.

Librarians or publishers of similar scientific and academic journals
may propose exchange agreements with the Australian Museum
Research Library. Subscription to Records of the Australian Museum is
available. Back issues may be purchased from the Australian Museum
Shop

http://shop.australianmuseum.net.au

Authors are invited to submit manuscripts to The Editor. Manu¬
scripts meeting subject and stylistic requirements outlined in the
Instructions to Authors (see inside back cover) are assessed by external
referees.

The Editor, Records of the Australian Museum

Australian Museum

6 College Street

Sydney NSW 2010, Australia

editor@austmus.gov.au

http://publications.australianmuseum.net.au

Our logo is based
on a distinctive and
local Aboriginal rock
engraving of the
echidna. This image has
special meaning for the
Australian Museum as it
represents both nature
and culture—a fitting
symbol for all that we do.

nature culture discover

© The Author, 2010. Journal compilation © Australian Museum, Sydney, 2010
Records of the Australian Museum (2010) Vol. 62: 217-284. ISSN 0067-1975
doi:10.3853/j.0067-1975.62.2010.1554

Uncovering Local Endemism in the Kimberley,
Western Australia: Description of New Species of the
Genus Amplirhagada Iredale, 1933
(Pulmonata: Camaenidae)

Frank Kohler

Department of Environment and Conservation of Western Australia, Science Division,
Wildlife Place, Woodvale WA 6026, Australia; and

Australian Museum, 6 College Street, Sydney NSW 2010, Australia
frank.koehler @ austmus.gov.au

Abstract. In this work twenty-six species of the camaenid land snail Amplirhagada, which is endemic to
the Kimberley region in Western Australia, are newly described. In addition, the anatomy of two further
species, A. katerana and A. combeana, is described and a further, yet undescribed species is reported from
Boongaree Island based on dry shell material. Most of these species occur on islands of the Bonaparte
Archipelago off the mainland coast. The patterns of distribution and differentiation of these island species
are comparable, however, with those found on the mainland. Mainland species are usually restricted to
single rainforest patches. Frequently, single patches harbour only one or two congeneric species. Similarly,
smaller islands usually support one endemic Amplirhagada species whereas on larger islands up to four
species are found to occur in sympatry. Species are distinguishable particularly by the characteristic
anatomy of the inner penial wall. Sympatric species generally exhibit marked morphological differences
in shells and genital anatomy. A molecular phylogeny based on partial sequences of the mitochondrial
cytochrome c oxidase unit 1 gene (COI) reveals a basal polytomy among species of the genus, which
are generally genetically well differentiated. Relationships among species in the molecular tree mainly
reflect geographical patterns.

Kohler, Frank, 2010. Uncovering local endemism in the Kimberley, Western Australia: description of new species of
the genus Amplirhagada Iredale, 1933 (Pulmonata: Camaenidae). Records of the Australian Museum 62(2): 217-284.

Camaenid land snails of the genus Amplirhagada Iredale,
1933, are endemic to the Kimberley region of Western
Australia, where they have radiated extensively. Thirty
species are currently considered valid, most of which usually
occupy small distributional ranges along the Kimberley
coast from the Buccaneer Archipelago in the south to
Kalumburu in the north as well as in inland areas of the
Napier, Harding, and King Leopold Ranges, the Drysdale

River Reserve, Mt Elizabeth Station and the region south of
Wyndham (Solem, 1981a, 1988). Many of these species are
restricted to single localities, such as rainforest patches or
more open woodlands (Solem, 1991). Supposed exceptions,
with species occupying larger ranges, are likely to result
from the inadequate delimitation of morphologically cryptic
species (Solem, 1981a). Earlier accounts were predominantly
based on dry shells (e.g., Smith, 1894; Iredale, 1933,

218

Records of the Australian Museum (2010) Vol. 62

124 ° 126 ° 128 °

- 14 °

- 16 °

- 18 °

Figure 1. Distribution of Amplirhagada species in the Kimberley region, northwestern Australia. (1) A. euroa n.sp., Adolphus Island. (2)
A. solemiana n.sp., Middle Osborn Island. (3) A. indistincta n.sp., Southwest Osborn Island, Kidney Island. (4) A. combeana, Cassini
Island. (5) A. mckenziei n.sp., Oliver Island. (6) A. ponderi n.sp., Kingsmill Island. (7) A. montesquieuana n.sp., Fenelon Island. (8) A.
descartesana n.sp., Descartes Island. (9) A. katerana, Katers Island. (10) A. puescheli n.sp., unnamed island near Prudhoe Island. (11) A.
decora n.sp., A. kessneri n.sp., Bigge Island. (12) A. berthierana n.sp., Berthier Island. (13) A. lamarckiana n.sp., Lamarck Island. (14) A.
anderdonensis n.sp., unnamed island, Anderdon Islands. (15) A. tricenaria n.sp., Prince Frederick Harbour, north of Hunter River mouth.
(16) A. regia n.sp., A. boongareensis n.sp., A. gibsoni n.sp., and Amplirhagada sp., Boongaree Island. (17) A. yorkensis n.sp., Coronation
Island. (18) A. bujfonensis n.sp., Buffon Island. (19) A. uwinsensis n.sp., Uwins Island. (20) A. sphaeroidea n.sp., St. Andrews Island.
(21) A. basilica n.sp., and A. camdenensis n.sp., Augustus Island. (22) A. kimberleyana n.sp., NW of Wilson Point. (23) A. gemina n.sp.,
1.5 km SE of Hall Point. (24) A. dubitabile n.sp., Steep Island. (25) A. napierana, north Napier ranges. (26) A. percita, Napier ranges.
(27) A. castra, A. mitchelliana, and A. varia, Mitchell Plateau.

1938, 1939). However, Solem (1981a, 1988) demonstrated
that Amplirhagada species frequently show only subtle
differences in shell features, such as colour, shape, and
sculpture. In addition, shells are prone to homoplasy, which
renders the delimitation of species by shell features alone
difficult or impossible. In fact, proper species delimitation
requires the study of genital anatomy, which was identified

as a convenient source of key morphological characters for
the recognition of species not only in this genus but also in
other camaenid genera (Solem, 1979, 1981a, 1981b, 1984,
1985; Willan et al., 2009; Kohler, 2010). In Amplirhagada
structure of the inner penial wall and development of the
main stimulatory pilaster provide particularly valuable
diagnostic characters for delimiting species (Solem, 1988).

Kohler: New Amplirhagada species from the Kimberley

219

A comparative study, mainly of the genital anatomy,
enabled Solem (198 la) to identify a large number of species
that were unrecognized by previous authors, who relied
exclusively on shell features. Solem showed that relatively
restricted areas in the Kimberley, such as the Mitchell
Plateau, may support a surprisingly large number of species
that can be differentiated on morphological criteria only by
their genital anatomy. Solem’s milestone publications on
the Camaenidae in Western Australia (Solem, 1979, 1981a,
1981b, 1984,1985,1988,1991; Solem & Christensen, 1984)
highlighted the remarkable diversity and patterns of local
endemism within this group of land snails. Even though he
published extensively on Western Australian Camaenidae,
Solem was not able to approach a complete documentation
of their extant diversity (Cameron et al., 2005).

In addition to thirty named species of Amplirhagada,
Solem (1991) listed 25 undescribed species from rainforest
patches and woodlands throughout the Kimberley region.
When Alan Solem died in 1990 he left numerous examples of
undescribed Amplirhagada species that are currently housed in
the Field Museum in Chicago. Given the poor documentation
of camaenid land snails from vast areas, such as most of
the islands off the Kimberley coast, and with documented
but undescribed material being kept in the collections of
various museums, many more species undoubtedly remain
to be discovered (Solem, 1988). Taking this high proportion
of undiscovered diversity into consideration; Amplirhagada
is surely the most species diverse camaenid genus in
northwestern Australia, with only a small fraction of its actual
species diversity having been described.

The aim of the present work is to enhance the
documentation of diversity of this endemic genus in the
Western Australian Kimberley region. Special attention
is paid to islands off the Kimberley coast, which have
previously been neglected. Twenty-seven Amplirhagada
species are newly described herein based on a comparative
study of samples from various sources. The core material
was collected in 2007-2008 during the Kimberley
Island Survey of the Department of Environment and
Conservation, Western Australia. This material is
supplemented by samples that were collected as early as
1987 and are housed in museum collections.

In addition to the study of morphological differentiation,
molecular phylogenies have been reconstructed based
on sequences of a partial fragment of the mitochondrial
cytochrome c oxidase gene (COI) in order to compare
rates of genetic and morphological differentiation and to
assess the diagnostic value of anatomical characters in a
phylogenetic context.

Material and methods

This study is primarily based on ethanol preserved specimens
and supplementary dry shell material collected during
the Kimberley Island Survey (KIS) of the Department of
Environment and Conservation, Western Australia (DEC)
in 2007-2009, the Kimberley Rainforest Survey of the then
Department of Conservation and Land Management, Western
Australia (RFS) in 1987-1988 and additional field work
conducted between 1988 and 2002 by various parties. Types
and other voucher material are deposited in the Western
Australian Museum, Perth (WAM), the Field Museum
of Natural History, Chicago (FMNH), and the Australian
Museum, Sydney (AMS).

Morphological descriptions focus on characters of the
shell, the mantle cavity, the genital organs, and the radula.
Morphometric shell parameters, such as height of shell (H),
diameter of shell (D), height of last whorl (LW), and width of
umbilicus (U) were measured with callipers precise to 0.1 mm.
The numbers of whorls (N), including the protoconch, were
counted as described Barker (1999: Fig. 6). The parameter
“angle of aperture” describes the angle formed between aperture
and the horizontal in degrees when the shell is in an upright
position. The morphometric shell parameters H, D, LW, U, N,
H/D were subjected to morphometric analyses when considered
necessary for the delimitation of species with similar shells.

Anatomy of mantle and genital organs was studied
using a binocular microscope with drawing mirror. If not
stated otherwise, the final inking of drawings was done by
Martin Piischel (Sydney). Radulae and jaws were extracted
manually, cleaned by soaking in 10% KOH solution for about
six hours followed by rinsing in water and ethanol. They
were mounted on carbon specimen tabs for electron scanning
microscopy. Radular tooth formula gives the numbers of
teeth as follows: C (central row of teeth)+number of lateral
rows of teeth+number of transitional rows of teeth+number
of marginal rows of teeth. The anatomy was studied in two
(in small series) to five specimens per sample in order to
confirm that morphological observations are consistently
found among conspecific specimens.

DNA was extracted from small pieces of foot muscle
by use of a QIAGEN DNA extraction kit for animal
tissue following the standard procedure of the manual.
A fragment of the mitochondrial cytochrome c oxidase
gene (COI) was amplified by PCR using the standard
primers LCO1490 and HC02198 of Folmer et al. (1994).
Reactions were performed under standard conditions with
an annealing temperature of 50°C. Phylogenetic trees were
reconstructed by application of Maximum Likelihood (ML)
using the software Treefinder version October 2008 (Jobb
et al., 2004), Bayesian Inference (BI) using the software
MrBayes 3.1.2 (Ronquist & Huelsenbeck, 2003), and
Maximum Parsimony using the Ratchet implemented in
WlNCLADA (Nixon, 1999) with 1,000 iterations, 10 trees
kept at each step. Nodal support of topologies was inferred
by applying MP bootstrapping (Felsenstein, 1985) or by
calculating Bayesian posterior clade probabilities (Larget
& Simon, 1999) and Expected-Likelihood Weights of Local
Rearrangements of tree topology (LR-ELW) (Strimmer
& Rambaut, 2002). Pair-wise genetic distances were
calculated with the software DAMBE (Xia & Xie, 2001).
Sequences have been deposited with GenBank under the
accessions GU302247 to GU302299.

220

Records of the Australian Museum (2010) Vol. 62

Systematic descriptions
Gastropoda
Heterobranchia
Stylommatophora
Camaenidae Pilsbry, 1895

Amplirhagada Iredale, 1933

Amplirhagada Iredale, 1933: 52; Solem, 1981a: 147-320;
Solem, 1988:28-32; Solem, 1991:187-202. Type species
Helix (Hadra) sykesi Smith, 1894 by original designation.

Tenuigada Iredale, 1939: 68. Type species Tenuigada percita
Iredale, 1939 by original designation.

Diagnosis

Rock or free sealer with medium sized (15-30 mm in
diameter), thin and translucent to moderately thick, broadly
conical to dome-shaped shell with moderately to strongly
elevated spire. Umbilicus concealed by columellar reflection
or narrowly open. Protoconch with weak to strongly
developed radially elongated pustulations; transition to
teleoconch inconspicuous. Teleoconch with very weak axial
growth lines that may become almost invisible on last whorl
or sculptured by well-developed, regular axial lirae. Last
whorl moderate to wide in cross-section; periphery well
rounded to slightly angulate (often transitions are found
within single populations). Shell colour usually variable
even within populations, background often yellowish brown
to horn, frequently with darker, brown bands located below
suture and on periphery of whorl; some species are uniform
in colour. Genitalia typically; development depends on
seasonal activity and maturity. Penis with well-developed
sheath, extending entire length, thin proximally, thick
distally. No well-developed epiphallus present. Penial
retractor muscle attached at apex of penial complex. Vas
deferens entering sheath from halfway up to almost apically;
entering penial chamber through verge. Inner penial wall
supports characteristic pustulation and basal pilasters;
a main stimulatory pilaster varies in development from
undifferentiated to very large, cone-shaped. Spermatheca
relatively simple, short reaching base of spermoviduct; duct
and head usually well differentiated; i nn er spermathecal
and vaginal wall with longitudinal pilasters that vary in
development and finer structure. Albumen gland elongate.
Hermaphroditic duct tightly undulating. Radula rectangular
in shape, usually between 3.5 and 5.5 mm long with 120-170
rows of teeth. Tooth formula variable, C+12-20+3-4+15-22.

Amplirhagada euroa n.sp.

Type locality (Fig. 1). Western Australia, eastern Kimberley,
Cambridge Gulf, eastern section of Adolphus Island,
15°06'32"S 128°09'08"E; KIS 3-113. East-facing gully with
ephemeral stream and patches of vine thicket, fig trees. Scree
on very steep slopes. In loose soil under large boulders (leg.
V. Kessner, 7 August 2008).

Type material. Holotype WAM S34601 (PL 1.1). Paratypes AMS
C463680 (6 preserved specimens, as holotype), WAM S37083 (12

preserved specimens, as holotype), WAM S37390 (3 shells, 15°06'18"S
128 o 09'04"E), WAM S37391 (shell, 15°06'35"S 128°09’57'’E), AMS
C463723 (6 shells, 15°06'32”S 128°09'08"E), WAM S37392 (11 shells,
15°06'32"S 128°09’08"E), WAM S37393 (3 shells, north of Adolphus
Island, 15°04'19"S 128°08’18"E).

Etymology. From euroa (Latin = eastern), for this species
occurring at the easternmost limits of the known range of
the genus.

Sealing strategy. Rock sealer.

Shell (Fig. 2A-B, PI. 1.1-2). Broadly conical with low to
medium high spire, thin to translucent. Periphery evenly
rounded to slightly angulate; upper and basal sectors of
whorls rounded. Umbilicus open, narrowly winding, 50-90
percent concealed by columellar reflection. Background
colour yellowish brown with chestnut brown, thin to
moderately broad, diffuse to well defined sub-sutural and
mid-whorl bands, clearly visible on last whorls only; ventral
colour brownish horn to whitish; outer lip colour tends
to be lighter than shell till whitish; inner lip translucent.
Protoconch c. 3 mm in diameter, comprising about 1.7
whorls, with faint, indistinct axial riblets. Teleoconch with
fine axial lirae, curved if viewed from above, pointed in
cross-section, irregularly spaced, spaces as wide as thickness
of lirae. Lirae evenly distributed across whorl diameter, with
reduced height underneath suture; present on all whorls.
Angle of aperture 45°. Outer lip simple rounded, sharp,
slightly to well expanded, slightly reflected; basal node
absent or very weak; palatal node absent. Parietal wall of
inner lip absent or inconspicuous.

Pallial morphology. Pallial cavity deep, extending one
whorl; mottled or spotted black mantle pigmentation. Kidney
extending about half of pallial cavity.

Genital morphology (Figs. 3-4). Penis straight, more or
less of same length as anterior part of oviduct. Vas deferens
coils before entering penis. Penial retractor muscle shorter
than penis complex. Penial verge short, less than Vs of
length of penial chamber, slender with rounded tip. Penial
wall pustules normal to elongated, arranged in rows over
entire length of inner penial wall. Main stimulatory pilaster
large, cone-shaped, comprising entire length of inner penial
wall; sculptured by ridges with smooth, flattened pustules.
Vas deferens entering penial sheath in upper third. Vagina
of medium length, posteriorly inflated; inner vaginal wall
supports undulating longitudinal pilasters. Spermatheca
short, reaching base of spermoviduct; duct wide, inner
wall with smooth longitudinal pilasters; head globular to
elongately inflated, connected with oviduct by connective
tissue, wall of head delicate, smooth. Free oviduct rather
straight comprising about half of length of anterior part of
oviduct. Spermoviduct longer than anterior part of oviduct.
Talon embedded in albumen gland close to anterior end of
albumen gland.

Radular morphology (Fig. 2C-E). Rectangular. Tooth
formula C+12-13+3-4+20-21. Average number of rows
of teeth 146±7 with 36.2±0.7 rows of teeth per mm (n = 2).
Central teeth with sharply pointed triangular mesocones,
shorter than base of tooth; ectocones vestigial. Lateral teeth
with bluntly pointed triangular mesocones, length equal to

Kohler: New Amplirhagada species from the Kimberley

221

Figure 2. SEM photographs of Amplirhagada euroa n.sp., paratype AMS C463680: (A) apical whorl viewed from above (scale 200 pm);
( B ) penultimate whorls viewed from above (scale 200 pm); (C) central and lateral radular teeth (scale 20 pm); (D) details of central and
lateral teeth (scale 10 pm); ( E ) details of outer lateral and inner marginal teeth (scale 10 pm); (F) jaw (scale 100 pm).

base of tooth; small ectocones; endocones absent. Marginal
teeth multicuspic, mesocone and endocone similar in length,
ectocone smaller than endocone, occasionally subdivided.

Comparative remarks. Amplirhagada euroa is geographic¬
ally well separated from most other congeneric species. Ampli¬
rhagada cambridgensis Solem, 1988 (from the western bank
of Cambridge Sound) and A. questronana Solem, 1981a (from
El Questro Station near Wyndham, c. 100 km S of Adolphus

Island) occur in closer proximity. The shell of A. cambridg¬
ensis is very similar but this species differs in the morphology
of the inner penial wall with pustules being arranged to form
corrugated longitudinal pilasters. Amplirhagada euroa differs
from A. questronana by umbilicus forming a chink instead
of being open and by its long main stimulatory pilaster that
supports flattened pustules (A. questronana has a short pilaster
with corrugations). Otherwise, both species have rather similar
shells with regard to shape and size.

222

Records of the Australian Museum (2010) Vol. 62

Figure 3. Genitalia of Amplirhagada euroa n.sp., paratype AMS
C463680 (7 August, scale 10 mm). Labelling of structures: ag,
albumen gland; hd, hermaphroditic duct; p, penial complex (penis
and penis sheath); ov, oviduct (free anterior part); rm, penial
retractor muscle; sd, spermathecal duct; sh, spermathecal head; sp,
spermoviduct (uterus, prostate); t, talon; va, vagina; vd, vas deferens.

Amplirhagada solemiana n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, Admiralty Gulf, west
coast of Middle Osborn Island, 14°18T8"S 125°59'35"E;
KIS 2-030. Small vine thicket at base of hill, west facing
slopes. Common on trees and bushes (leg. V. Kessner, 13
February 2008).

Type material. Holotype WAM S34602 (PL 1.3). Paratypes AMS
C463681 (8 preserved specimens, as holotype), WAM S36860 (20
preserved specimens, as holotype), AMS C463683 (4 shells, 14°18'37.4"S
125°59'18.4"E), WAM S36572 (10 shells, 14°18'37.4”S 125°59'18.4"E),
AMS C463682 (2 preserved specimens, east coast, 14°18’53.5"S
126°02'06.7"E), WAM S36859 (6 preserved specimens, east coast,
14°18'53.5"S 126°02'06.7"E).

Additional, non-type material. WAM S36471, WAM S36566-71, WAM
S36573, WAM S36585-99, WAM S36858, WAM S36861, WAM S36971,
AMS C463684-5 (Middle Osborn Island).

Etymology. Named in honour of Alan Solem, in recognition
of his achievements in camaenid systematics.

Sealing strategy. Rock sealer.

Shell (Fig. 5A-E, PL 1.3-4). Semi-globose with moderately
high spire. Thin to solid, translucent. Periphery evenly
rounded to slightly angulate; upper and basal sectors of
whorls well rounded. Umbilicus open, forming a chink,
to 80-90 percent concealed by columellar reflection.
Background colour horn to yellowish brown, with brown,
well defined to diffuse, moderately broad, sub-sutural and

Figure 4. Interior of penial chamber of Amplirhagada euroa n.sp.,
paratype AMS C463680 (7 August, scale 5 mm). Labelling of
structures: iw, inner penial wall; lp, longitudinal (main) pilaster;
rm, penial retractor muscle; sh, penial sheath; vd, vas deferens;
vg, penial verge.

mid-whorl bands, visible on entire shell; ventral and outer
lip colour horn; inner lip translucent, pale. Protoconch c. 1.8
mm in diameter, comprising about 1.5 whorls, sculptured by
comparatively strong axial ribs. Teleoconch sculptured by
coarse, regular lirae, rounded in cross-section; spaces equal
to thickness of lirae; sculpture evenly distributed across
whorls of shell and across whorl diameter, height of lirae
reduced underneath suture. Angle of aperture 45°; outer
lip sharp to moderately thick, rounded, slightly expanded,
slightly reflected; basal and palatal node absent. Parietal wall
of inner lip inconspicuous.

Figure 5 (facing page). SEM photographs of Amplirhagada solemiana n.sp. (A-E). Shell, paratype AMS C463683: (A) apical whorl viewed
from above (scale 200 pm); ( B ) first four whorls viewed from above (scale 1 mm); (C) shell, lateral view (scale 1 mm); (D) details of axial
sculpture on last whorl, lateral view (scale 1 mm); (E) close-up of axial lirae on last whorl (scale 100 pm). (F-I) Radula, paratype AMS
C463681: (F) rows of central and lateral teeth (Scale 20 pm); (G) details if central and lateral teeth (scale 20 pm); (H) outer lateral and
inner marginal teeth (scale 10 pm); (/) middle and outer marginal teeth (scale 10 pm). (7) jaw, paratype AMS C463681 (scale 100 pm).

Kohler: New Amplirhagada species from the Kimberley

223

224

Records of the Australian Museum (2010) Vol. 62

Figure 6. Genitalia of Amplirhagada solemiana n.sp., paratype
WAM S36581 (13 February, scale 10 mm). Compare with Fig. 3
for labelling of structures.

Pallial morphology. Pallial cavity deep, extending one
whorl. Pigmentation on mantle consists of sparsely spaced
dark grey-black spots. Kidney extending about half of pallial
cavity or slightly more.

Genital morphology (Figs. 6-7). Penis straight to slightly
curved; same length as anterior part of oviduct or slightly
longer. Vas deferens forms simple loop before entering penis.
Penial retractor muscle about as long as penis. Penial verge
long to very long (c. Vs to Vi of penial chamber), slender to
spatulate with pointed tip. Pustulation comprising entire
length of inner penial wall; pustules small to moderate in
size, densely arranged in rows, some of which form four to
five corrugated longitudinal pilasters along entire length of
penial chamber; main stimulatory pilaster not differentiated.
Vas deferens entering penial sheath apically. Vagina of
medium length, tubular or posteriorly inflated. Inner vaginal
wall densely ciliated, ciliae arranged to form smooth
longitudinal pilasters, pilasters may be weakly developed.
Spermathecal duct of medium thickness; internally with
smooth longitudinal pilasters. Spermathecal head elongately
inflated, connected with oviduct by connective tissue; inside
entirely smooth; wall moderately thick. Length of free
oviduct equivalent to about half of anterior part of oviduct;
coiled underneath entrance to spermoviduct. Spermoviduct
longer than anterior part of oviduct. Talon embedded in
albumen gland close to anterior end.

Radular morphology (Fig. 5F-I). Rectangular. Tooth
formula C+9-12+3-4+19-21. Average number of rows of
teeth 131±22 with 29.9+0.6 rows per mm (n = 3).

Central teeth with bluntly pointed, ovate mesocones,
shorter than base of tooth; ectocones vestigial. Lateral teeth
with bluntly pointed ovate mesocones, length equal to base
of tooth; small ectocones; endocones absent. Marginal teeth
multicuspic, mesocone and endocone similar in length,
ectocone smaller than endocone, occasionally subdivided.

Comparative remarks. Shells are close to Amplirhagada
imitata (E. A. Smith, 1894) in overall shape, colouration, and
sculpture, but see Fig. 8. The type locality of A. imitata was
restricted to Baudin Island by Solem (1981a). Amplirhagada
imitata and A. solemiana exhibit the most sharply defined
and prominent axial sculpture among all congeners except
for A. indistincta. The anatomy of A. imitata is unknown for
only dry shells were available to Solem (1981a). In between

Figure 7. Interior of penial chamber of Amplirhagada solemiana
n.sp., paratype AMS C463681 (13 February, scale 3 mm). Compare
with Fig. 4 for labelling of stmctures.

Baudin Island (A. imitata ) and the two Osborn Islands (A.
solemiana and A. indistincta) there are numerous other islands
that are occupied by Amplirhagada species (e.g., Kingsmill,
Corneille, Fenelon, and Descartes Islands) (Fig. 1). Given the
usually very restricted occurrence of Amplirhagada species
and the marked genetic differentiation of A. solemiana and
A. indistincta , it is considered very unlikely that either of
them could be conspecific with A. imitata from Baudin
Island, which is located in a considerable distance (c. 50 km
NW of Osbom Islands). Based on shell features only, Solem
(1981a) also synonymized A. burrowsena Iredale, 1938 from
Vansittart Bay (c. 80 km E of Baudin Is) with A. imitata and
reported this species to further occur in rainforest patches on
the mainland (RFS 03/3,11/1). These reports, however, almost
certainly refer to extralimital populations. Species such as A.
imitata, A. solemiana, A. indistincta, A. burrowsena and likely
an undescribed species from the rainforest patches on the
mainland may possibly form a group of sibling species that are
not clearly differentiated by means of their shell morphology
as is also known from the species group of A. mitchelliana
Solem, 1981 from the Mitchell Plateau.

Kohler: New Amplirhagada species from the Kimberley

225

15 ■

14 -

13-

12 -

1 111

g> 10-

CD

O 0 0

o O A

*

*

o

o o
o

o A. indistincta
m A. solemiana

16 17 18 19 20 21 22

shell diameter (D) [mm]

Figure 8. Comparison of A. solemiana and A. indistincta by means
of shell parameters. Scatter-plot showing the ratio of shell height
(H) versus diameter (D).

Amplirhagada indistincta n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, Admiralty Gulf, West
coast of South West Osborn Island, 14°22’26"S 125°56'13"E;
KIS 3-7. Isolated vine thicket patch on dune behind sand
beach, on trees, under logs (leg. V. Kessner, 29 July 2007).

Type material. Holotype WAM S34603 (PL 1.5). Paratypes AMS
C463686 (5 preserved specimens, as holotype), WAM S36617 (10
preserved specimens, as holotype), AMS C463688 (5 shells, 14°22'26"S
125°56'13"E),WAM S36575 (15 shells, 14°22'26"S 125°56’13"E), AMS
C463687 (20 preserved specimens, 14°22’26.5"S 125°56'17.8"E), WAM
S36865 (40 preserved specimens, 14°22'26.5"S 125°56'17.8"E), AMS
C463689(ll shells, east coast, 14°22'47.3"S 125°56'00.6"E),WAMS36962
(25 shells, east coast, 14°22'47.3''S 125°56’00.6''E).

Additional, non-type material. WAM S28521, WAM S36491-501, WAM
S36574, WAM S36577, WAM S36612-16, WAM S36618-25, WAM
S36863^4, WAM S36866, AMS C463690-1 (South West Osborn Island);
WAM S41454, AMS C463724 (Kidney Island; 14.329°S 125.985°E).

Etymology. From indistincta (Latin = indistinct) referring
to the close morphological resemblance with A. solemiana.

Sealing strategy. Rock sealer.

Shell (Fig. 11A-D, PL 1.5-6). Semi-globose to broadly
conical with moderate to high spire; solid to thick, not
translucent. Periphery well rounded to slightly angulate.
Umbilicus completely concealed by columellar reflection.
Background colour yellowish to brownish white; with thin
to moderately broad brown to yellowish brown sub-sutural
and mid-whorls bands that are visible on most whorls, sub-
sutural band diffuse, mid-whorl band well marked; ventral
colour whitish to greyish white; outer lip colour differs
from shell, whitish; inner lip translucent, white. Protoconch

Figure 9. Genitalia of Amplirhagada indistincta n.sp., paratype
AMS C463687 (11 February, scale 10 mm). Compare with Fig. 3
for labelling of structures.

Figure 10. Interior of penial chamber of Amplirhagada indistincta
n.sp., paratype AMS C463687 (11 February, scale 5 mm). Compare
with Fig. 4 for labelling of structures.

226

Records of the Australian Museum (2010) Vol. 62

Figure 11. SEM photographs of Amplirhagadaindistinctan.sp. ( A-D ) Shell: (A) apical whorl viewed from above, paratypeAMS C463688
(scale 200 pm); ( B ) first four whorls viewed from above, paratypeAMS C463689 (scale 1 mm); (C) details of axial sculpture on last whorl,
lateral view, paratypeAMS C463689 (scale 1 mm); ( D ) close-up of sculpture on last whorl, paratypeAMS C463689 (scale 100 pm); (E)
jaw, paratypeAMS C463687 (scale 100 pm). ( F-G ) Radula, paratypeAMS C463687: (F) rows of central and lateral teeth (Scale 20 pm);
(G) outer lateral and inner marginal teeth (scale 10 pm).

c. 2 mm in diameter, comprising about 1.5 whorls, with
strong axial sculpture. Teleoconch sculptured by coarse,
regular, curved, in cross-section rounded lirae; sculpture
evenly distributed across shell and whorl diameter; spaces
between lirae equal to thickness of lirae, height of lirae
reduced underneath suture. Angle of aperture 30°; outer
lip rounded, sharp to moderately thick, expanded, slightly
reflected; basal node of lip weak, palatal node absent.
Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity deep, extending one
whorl. Pigmentation on mantle consists of sparsely
distributed dark brown to black spots. Kidney extending
more than half of pallial cavity.

Genital morphology (Figs. 9-10). Penis straight to slightly
curved; same length as anterior part of oviduct or slightly
longer. Vas deferens forms simple loop before entering penis.
Penial retractor muscle about as long as penis. Penial verge

Kohler: New Amplirhagada species from the Kimberley

227

long to very long (c. Vs to Vi of penial chamber), slender to
spatulate with pointed tip. Pustulation comprising entire
length of inner penial wall; penial wall pustules small to
moderate in size, densely arranged in rows, some of which
form four to five longitudinal pilasters along entire length of
penial chamber; main stimulatory pilaster not differentiated.
Vas deferens entering penial sheath apically. Vagina of
medium length, tubular or posteriorly inflated. Inner vaginal
wall densely ciliated, ciliae arranged to form smooth
longitudinal pilasters, pilasters may be weakly developed.
Spermathecal duct of medium thickness; internally with
smooth longitudinal pilasters. Spermathecal head elongately
inflated, connected with oviduct by connective tissue; inside
entirely smooth; wall moderately thick. Length of free
oviduct equivalent to about half of anterior part of oviduct;
coiled underneath entrance to spermoviduct. Spermoviduct
longer than anterior part of oviduct. Talon embedded in
albumen gland close to anterior end.

Radular morphology (Fig. 11F-G). Rectangular. Tooth
formula C+12+3-4+20-21. Average number of rows of teeth
126±14 with 26.3±1.5 rows per mm (n = 2). Central teeth
with bluntly pointed, ovate mesocones, shorter than base of
tooth; ectocones vestigial. Lateral teeth with bluntly pointed,
ovate mesocones, shorter than base of tooth; ectocones small,
endocones absent. Marginal teeth multicuspic; ectocones
shorter and narrower than mesocones, split into two denticles;
endocones of approximately same size as ectocones.

Comparative remarks. Most similar to A. solemiana in
shell and genital anatomy. Shells of A. indistincta are larger
than those of A. solemiana (Table 1); a one-way ANOVA
revealed that both species differ significantly in the shell
parameters H, D, and H/D (Fig. 8). Specimens from Kidney
Island, which is located in between Middle and Southwest
Osborn Island, tend to have slightly larger and more robust
shells than specimens from the type locality. In the molecular
phylogeny they cluster closely together with A. indistincta and
are therefore considered conspecific because their anatomy
corresponds with the specimens from Southwest Osborn
Island. Amplirhagada solemiana and A. indistincta exhibit
a virtually identical genital anatomy but cluster as clearly
distinct lineages in the mitochondrial phylogeny (see below).
For relationship with A. imitata see under A. solemiana.

Amplirhagada combeana Iredale, 1938

Helix (Hadra) imitata var. cassiniensis Smith, 1894: 92, pi. 7,
fig. 16 (Cassini Island). Amplirhagada combeana Iredale,
1938: 113 (nomen novum), Solem, 1981a: 310-312, pi.
12b, figs. 71h-i).

Nomenclatural remarks. The original specific epithet
“ cassiniensis ” employed by Smith (1894) is preoccupied
and has been replaced with “ combeana ” by Iredale (1938),
who elevated the taxon to the rank of an independent species.
Solem (1981a: 310-312) described details of the shell but
was not able to describe the anatomy because no preserved
material was available.

Material examined. WAM S41450, AMS C463725 (Western Australia,
northwestern Kimberley, Bonaparte Archipelago, Cassini Island,
13°57'04"S 125°38’39"), WAM S41455, AMS C463726 (13°57’22"S
125°37’53"E) (Fig. 1).

Sealing strategy. Rock sealer.

Shell (Fig. 14A-C, PI. 1.7). Broadly conical with low to
medium spire; solid (not translucent). Periphery well rounded
to slightly angulate. Umbilicus completely concealed by
columellar reflection or forming a chink. Background
colour horn to yellowish brown; with diffuse to well defined,
thin to moderately broad, chestnut brown sub-sutural and

228

Records of the Australian Museum (2010) Vol. 62

Figure 14. SEM photographs of Amplirhagada combeana, AMS C463725. (A-C) Shell: (A) apical whorl viewed from above (scale 200
pm); ( B ) first four whorls viewed from above (scale 1 mm); (C) details of axial sculpture on last whorl (scale 100 pm). ( D-E) Radula:
(D) close-up of central and lateral radular teeth (Scale 10 pm); (E) outer lateral and inner marginal teeth (scale 10 pm).

mid-whorl bands that are most conspicuous on last whorl;
ventral colour horn; outer lip colour same as shell; inner
lip translucent. Protoconch c. 1.7 mm in diameter with 1.5
whorls, comparatively strong axial sculpture. Teleoconch
sculptured by coarse, curved, in cross-section rounded,
irregularly spaced lirae; spaces equal to thickness of lirae;
lirae evenly distributed across shell and whorl diameter,
height reduced underneath suture. Angle of aperture 30°;
outer lip rounded, moderately thick, expanded, slightly
reflected; basal node of lip weak, palatal node absent.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl. Pigmentation on mantle consists of
sparse greyish patches or spots. Kidney extending not more
than half of pallial cavity.

Genital morphology (Figs. 12-13). Penis straight; more
or less of same length as anterior part of oviduct or longer.
Vas deferens forms simple loop before entering penis.
Penial retractor muscle shorter than penis. Penial verge
short (c'/s penial chamber), broad with pointed to rounded
tip. Penial wall covered by a number of smooth longitudinal
pilasters; pustulation absent. Main stimulatory pilaster well-
differentiated, extending apical to median portion of penial
chamber, cone-shaped, corrugated by ridges that support little
hooks. Vas deferens entering penial sheath in upper third.
Vagina of medium thickness, posteriorly inflated. Inner vaginal
wall with smooth, longitudinal pilasters. Spermathecal duct

moderately thick, inside with smooth longitudinal pilasters.
Spermathecal head globular, connected with oviduct by
connective tissue, inside entirely smooth with delicate
wall. Free oviduct comprising about half of anterior part
of oviduct; coiled underneath entrance to spermoviduct.
Spermoviduct longer than anterior part of oviduct. Talon
embedded in albumen gland at junction with spermoviduct.

Radular morphology (Fig. 14D-E). Rectangular. Tooth
formula C+12+2-3+18-20 with 162.5±3.5 rows of teeth,
33.5±1.9 rows per mm (n = 2). Central teeth with bluntly
pointed, triangular to ovate mesocones, shorter than base of
tooth; ectocones well developed. Lateral teeth with bluntly
pointed, triangular to ovate mesocones; length equal to base
of tooth; ectocones well developed, endocones vestigial.
Marginal multicuspic; ectocones shorter and narrower
than mesocones; endocones approximately same size as
ectocones; divided into two denticles.

Comparative remarks. This species differs from all previous
species by the absence of penial wall pustules and the presence
of a well-developed main stimulatory pilaster. Axial sculpture
is not as regular as in A. solemiana and A. indistincta but more
pronounced as in A. euroa. With respect to penial morphology
A. solemiana and A. indistincta differ most markedly by
absence of a well-differentiated main stimulatory pilaster.
For comparison with A. mckenziei and A. montesquieuana see
below, with other species see Solem (1981a: 310).

Kohler: New Amplirhagada species from the Kimberley

229

Amplirhagada mckenziei n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, Admiralty Gulf, Mont¬
esquieu Islands, Oliver Island, 14°05'42"S 125°44'30"E;
KC-062 (leg. V. Kessner & A. Longbottom, 18 June 1988).

Type material. Holotype WAM S34604 (PL 1.8). Paratypes WAM S41488
(3 preserved specimens), FMNH 219253 (3 preserved specimens).

Additional, non-type material. AMS C463727, FMNH 219250,
WAM S41487 (Western Australia, northwestern Kimberley, Bonaparte
Archipelago, Admiralty Gulf, Montesquieu Islands, unnamed island SE of
Oliver Island, 14°05'55"S 125°44'50"E; KC-061).

Etymology. Named in honour of Norman L. McKenzie,
Western Australian Department of Environment and Con¬
servation, in recognition of his promotion of malacological
research in Western Australia.

Sealing strategy. Rock sealer.

Shell (Fig. 17A-C, PL 1.8-9). Broadly conical to almost
flat with low to medium spire. Thin to solid, translucent.
Periphery rounded. Umbilicus concealed by columellar
reflection to 50-100 percent, forming a chink or narrowly
winding opening. Background colour brownish horn,
ventrally lighter; with diffuse to well marked, thin,
yellowish brown sub-sutural and mid-whorl bands that are
most conspicuous on last whorl; outer lip colour same as
shell; inner lip translucent, whitish. Protoconch c. 2.2 mm
in diameter with 1.5 whorls, sculptured by strong axial
elements. Teleoconch with coarse axial lirae; curved when
viewed from above; rounded in cross-section with irregular
spacing; spaces equal to thickness of lirae; distributed evenly
across shell and across whorl diameter, reduced underneath
suture. Angle of aperture 30°; outer lip rounded, sharp to
moderately thick, slightly expanded, not or slightly reflected;
basal node absent or weak; palatal node absent. Parietal wall
of inner lip inconspicuous.

Pallial morphology. Pallial cavity short, extending half
whorl. Mantle pigmentation consists of blackish spots.
Kidney extending more than half of pallial cavity.

Figure 15. Genitalia of Amplirhagada mckenziei n.sp., paratype
WAM S41488 (18 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Genital morphology (Figs. 15-16). Penis straight, slightly
longer than anterior part of oviduct. Vas deferens forms
simple loop before entering penis. Penial retractor muscle
shorter than penis. Penial verge very short (barely visible) to
short (<Vs penial chamber), slender to spatulate with pointed
tip. Penial wall pustules small, arranged in rows over entire
length of inner penial wall. Main stimulatory pilaster well-
differentiated, relatively thin, cone-shaped, sculptured by
ridges with little hooks, on apical portion of inner penial
wall. Yas deferens rather straight to undulated, rather thick,
entering penial sheath in upper third of penis. Vagina rather
thick, tubular to posteriorly inflated. Inner vaginal wall with
smooth longitudinal pilasters. Spermathecal duct wide, with
smooth longitudinal pilasters inside. Spermathecal head
globular to elongately inflated, connected with oviduct by
connective tissue; inside entirely smooth with thin wall. Free
oviduct more or less straight comprising less than half to
about half of length anterior part of oviduct of anterior part
of oviduct. Spermoviduct of same length as anterior part of
oviduct. Talon embedded in albumen gland at junction with
spermoviduct.

Figure 16. Interior of penial chamber of Amplirhagada mckenziei
n.sp., paratype FMNH 219253 (18 July, scale 10 mm). Compare
with Fig. 4 for labelling of structures.

230

Records of the Australian Museum (2010) Vol. 62

Figure 17. SEM photographs of Amplirhagada mckenziei n.sp., paratypes WAM S41488. (A-C) Shell: (A) apical whorl viewed from
above (scale 200 pm); ( B ) first four whorls viewed from above (scale 200 pm); (C) details of axial sculpture on last whorl (scale 100 pm).
( D ) Jaw (scale 100 pm). (E-F) Radula: (E) close-up of central and inner lateral radular teeth (Scale 10 pm); (E) outer lateral and inner
marginal teeth (scale 10 pm).

Radular morphology (Fig. 17E-F). Rectangular. Tooth
formula C+12-18+2-3+18-20. With in average 135±12.5
rows of teeth, 31.2±1.2 rows per mm (n = 2). Central teeth
with sharply pointed, triangular mesocones, shorter than
base of tooth; ectocones vestigial. Lateral teeth with sharply
pointed, triangular mesocones, length equal to base of
tooth; ectocones tiny, endocones vestigial. Marginal teeth
with triangular to ovate mesocones; ectocones shorter and
narrower than mesocones; endocones reduced in size.

Comparative remarks. The studied museum material was
labelled as “Amplirhagada sp. 72” by Solem. This species
is closely related to A. montesquieuana (see below) and A.
combeana, both inhabiting islands in relative proximity.
Amplirhagada combeana is similar in exhibiting a relatively
pronounced but irregular axial sculpture but differs in having
smaller main stimulatory pilaster, penial verge and no penial
wall pustules.

Kohler: New Amplirhagada species from the Kimberley

231

Amplirhagada ponderi n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, Admiralty Gulf,
Kingsmill Island, 14°09’24"S 125°46'16"E; KC-067 (leg.
V. Kessner & A. Longbottom, 19 July 1988).

Type material. HolotypeWAMS34605 (PL 1.10). ParatypeWAMS41489
(1 preserved specimen), FMNH 219268 (1 preserved specimen).

Etymology. Named in honour of Winston F. Ponder, senior
fellow of the Australian Museum, in recognition of his
achievements in malacological research.

Shell (PL 1.10). Broadly conical, with low spire; thin
(translucent) to solid. Periphery slightly angulate; upper
and basal sectors of whorls rounded. Umbilicus open,
narrowly winding, c. 40-60 percent concealed by columellar
reflection. Background colour yellowish brown to horn; with
diffuse, thin, light brown sub-sutural and mid-whorl bands
being most conspicuous on last whorl; ventral colour whitish
horn; outer lip colour same as shell; inner lip colour horn to
translucent. Protoconch and teleoconch smooth except for
growth lines. Angle of aperture 45°; outer lip rounded, sharp
to moderately thick, expanded, slightly reflected; basal node
of lip weak to moderately developed; palatal node absent.
Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl. Blackish mottled pigmentation on
mantle. Kidney extending half of pallial cavity.

Genital morphology (Figs. 18-19). Penis bent; much
longer than anterior part of oviduct. Vas deferens forms
complex coiling before entering penis. Penial retractor
muscle stubby, much shorter than penis. Penial verge
medium sized to long (Vs-Vs of penial chamber), slender
to spatulate with pointed tip. Penial wall pustules very
small, arranged in rows over entire length of inner penial
wall. Main stimulatory pilaster well-differentiated, narrow
and quite long comprising apical to median portion of
penial chamber; corrugated by ridges of undifferentiated
pustules that support little hooks. Vas deferens entering

Figure 18. Genitalia of Amplirhagada ponderi n.sp., paratype
FMNH 219268 (19 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

penial sheath in upper third. Vagina relatively long, tubular,
posteriorly slightly inflated. Inner vaginal wall with smooth
longitudinal pilasters. Spermathecal duct moderately thick
with smooth longitudinal pilasters inside. Spermathecal head
elongately inflated, connected with oviduct by connective
tissue, entirely smooth inside; wall delicate. Length of free
oviduct less than half of anterior part of oviduct, zig-zag-
folded underneath entrance to spermoviduct. Posterior part
of genitalia unknown.

Radular morphology (Fig. 20A-C). Rectangular. Tooth
formula C+16+3+18. Average number of rows of teeth 165 (n
= 1) with 35.6 rows per mm (n = 1). Central teeth with sharply
pointed, elongate to triangular mesocones, shorter than base
of tooth; ectocones vestigial. Lateral teeth with sharply
pointed, elongate to triangular mesocones, not exceeding
base of tooth; ectocones well developed, endocones vestigial.
Marginal teeth with elongate to triangular mesocones;
ectocones shorter and narrower than mesocones; endocones
reduced in size.

Comparative remarks. The studied museum material
was labelled as “Amplirhagada sp. 73” by Solem. The
shell of this species is similar to those of other species
from the Montesquieu Islands, such as A. mckenziei, A.
montesquieuana or A. puescheli (for comparison with the
latter two see below), and is not readily differentiated from
the former in shell characters alone. The inner penial wall

Figure 19. Interior of penial chamber of Amplirhagada ponderi
n.sp., paratype FMNH 219268 (19 July, scale 5 mm). Compare
with Fig. 4 for labelling of structures.

232

Records of the Australian Museum (2010) Vol. 62

Figure 20. SEM photographs of the radula of Amplirhagada ponderi n.sp., paratype FMNH 219268: (A) central and inner lateral teeth (Scale
20 pm); ( B ) close-up of inner lateral teeth (scale 10 pm); (C) outer lateral and inner marginal teeth (scale 10 pm); (D) jaw (scale 100 pm).

of A. ponderi , however, is very characteristic. It differs
from A. mckenziei most markedly by its much longer and
narrow main pilaster, which extends almost to the anterior
end of the penial chamber. The penis of A. ponderi is
proportionally longer. For comparison with other species
from the Montesquieu Islands see below. Phylogenetically,
A. ponderi is more closely related to A. varia, A. solemiana,
and A. indistincta. From all these it differs by possessing a
well-developed, corrugated main pilaster.

Amplirhagada montesquieuana n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, Admiralty Gulf, Fenelon
Island, 14°08T6"S 125°4F55"E; FERT 04 (leg. Harvey,
28.08.2002).

Type material. Holotype WAM S34606 (PI. 1.11). Paratypes AMS
C463748 (9 preserved specimens), WAM S41453 (15 preserved
specimens).

Etymology. In reference to Montesquieu Islands, to which
Fenelon Island belongs.

Sealing strategy. Rock sealer.

Shell (Fig. 21A-B, PI. 1.11). Broadly conical with low spire;
thin (translucent). Whorls evenly rounded in cross-section.
Umbilicus open, forming a chink to narrowly winding

opening, 80-95 percent concealed by columellar reflection.
Background colour whitish or yellowish to golden brown;
with diffuse, thin, brown sub-sutural and mid-whorl bands
most conspicuous on last whorl; ventral colour whitish;
outer lip colour same as shell or lighter; inner lip whitish
and translucent. Protoconch c. 2 mm in diameter with 1.5
whorls, sculptured by fine but distinct axial lirae. Teleoconch
with coarse axial lirae, rounded in cross-section with regular
spacing, spaces equal to thickness of lirae. Sculpture evenly
distributed across shell and whorl diameter, height reduced
underneath suture. Angle of aperture 30°, outer lip rounded,
sharp to moderately thick, slightly expanded and reflected;
basal node of lip weak, palatal node absent. Parietal wall of
inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl. Mantle with mottled, black pigmentation.
Kidney extending about half of pallial cavity.

Genital morphology (Figs. 22-23). Penis straight, of more
or less same length as anterior part of oviduct to slightly
longer. Vas deferens coils once before entering penis. Penial
retractor muscle clearly shorter than penis. Penial verge
very short, slender to spatulate, with pointed tip. Penial
wall pustules of normal size, slightly elongated, arranged in
sparsely distributed, distinct rows over entire length of inner
penial wall. Towards base of penial chamber, pustules fuse to
form corrugated, narrow pilasters. Main stimulatory pilaster

Kohler: New Amplirhagada species from the Kimberley

233

Figure 21. SEM photographs of Amplirhagada montesquieuana n.sp., paratypes AMS C463748. (A-B) Shell: (A) apical whorl viewed
from above (scale 200 pm); ( B ) sculpture on last two whorls viewed from above (scale 200 pm). ( C-F) Radula: (C) central and lateral
teeth (Scale 20 pm); (D) close-up of central and inner lateral teeth (scale 10 pm); (E) outer lateral and inner marginal teeth (scale 10 pm);
(F) close-up of middle marginal teeth (scale 10 pm). (G) Jaw (scale 100 pm).

differentiated, forming cone-shaped, prominent ridge covered
by enlarged pustules at apical to median portion of penial
chamber. Pilaster ridges carry little toe-shaped extensions
that support hooks. Vas deferens entering penial sheath in
upper third. Vagina moderately long, tubular. Inner vaginal
wall with smooth longitudinal pilasters. Spermathecal duct
wide; inner wall with transversely structured, longitudinal
pilasters. Spermathecal head globular to elongate, connected
with oviduct by connective tissue; wall delicate. Free oviduct
comprising±half of anterior part of oviduct, rather straight.
Spermoviduct longer than anterior part of oviduct. Talon

embedded in albumen gland at junction with spermoviduct.

Radular morphology (Fig. 21 C-F). Rectangular. Tooth
formula C+11+4+14. Average number of rows of teeth
150+32.5 with 38.1+0.3 rows per mm (n = 2). Central teeth
with sharply pointed, triangular mesocones, shorter than base
of tooth; ectocones small. Lateral teeth with bluntly pointed,
triangular mesocones, length equal to base of tooth; ectocones
well developed, endocones vestigial. Marginal teeth with
elongate mesocones; ectocones shorter and narrower than
mesocones, split into two denticles; endocones reduced in size.

234

Records of the Australian Museum (2010) Vol. 62

Figure 22. Genitalia of Amplirhagada montesquieuana n.sp.,
paratype AMS C463748 (28 August, scale 10 mm). Compare with
Fig. 3 for labelling of structures.

Fig. 4 for labelling of structures.

Comparative remarks. The shell of this species is smaller
than that of other species from the Montesquieu Islands
but otherwise difficult to differentiate by means of colour,
sculpture and shape (Table 1). Its axial sculpture is more
regular than that of A. combeana and A. mckenziei. From
A. mckenziei it also differs by a smaller and comparatively

flatter shell; a one-way ANOVA revealed that both species
differ significantly in the shell parameters H, D, FW and
H/D. The inner penial wall is characteristic by its rather
sparsely distributed rows of pustules that give rise to
corrugated, narrow, longitudinal pilasters towards the base
of the penial chamber, and the shape and development of the
main stimulatory pilaster being shorter and broader than in
A. ponderi and larger than in A. mckenziei. The possession
of well-developed “toe-shaped” extensions of the pilaster
ridges is unique amongst species from Montesquieu Islands.

Amplirhagada descartesana n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Admiralty Gulf, Institute Islands, Descartes
Island, 14°10'26"S 125°40’38"E; KC-071 (leg. V. Kessner
& A. Longbottom, 19 July 1988).

Type material. Holotype WAM S34607 (PI. 1.12). Paratypes WAM
S41491 (4 preserved specimens, as holotype), FMNH 219276 (5 preserved
specimens, as holotype), FMNH 219272 (16 preserved specimens, unnamed
island SW of Descartes Island, 14°10'45"S 125°40'00"E, KC/070), WAM
S41490 (10 preserved specimens, same as FMNH 219272), AMS C463749
(7 preserved specimens, same as FMNH 219272).

Additional, non-type material. WAM S41452 (Descartes Island,
125°40'47"E 14°09’50"S).

Etymology. In reference to Descartes Island, where this
species occurs.

Shell (Fig. 24A-D, PI. 1.12). Semi-globose to broadly conical
with medium high spire; thin (translucent) to solid. Periphery
evenly rounded to slightly angulate; upper and basal sectors
of whorls rounded. Umbilicus forming a chink to narrowly
winding opening, 30-100 percent concealed by columellar
reflection. Background colour brownish horn; uniform or
banded; if present sub-sutural and mid-whorl bands diffuse,
thin, brown, on last whorl(s) only; ventral colour, outer and
inner lip colour whitish to cream. Protoconch c. 2.8 mm in
diameter with 1.7 whorls, almost smooth, sculptured by faint
axial lirae. Teleoconch sculptured by coarse lirae, rounded
in cross-section, regularly spaced, spaces equal to thickness
of lirae, evenly distributed across shell and whorl diameter
even, reduced underneath suture. Angle of aperture 45°, outer
lip rounded, moderately thick, slightly expanded, slightly
reflected; basal node absent or weak, palatal node absent.
Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending % whorl. Mottled pigmentation on mantle dark
grey. Kidney extending about half of pallial cavity.

Genital morphology (Figs. 25-26). Penis straight to
slightly curved, longer than anterior part of oviduct. Vas
deferens forms simple loop before entering penis. Penial
retractor muscle shorter than penis. Penial verge short
(< X A penial chamber), slender to spatulate, with pointed
tip. Penial wall pustules present, of average size, slightly
elongated, arranged in rows across entire length of inner
penial wall. Main stimulatory pilaster well-differentiated,
large, cone-shaped, sculptured by smooth horizontal ridges,
comprising entire length of penial chamber. Vas deferens
entering penial sheath in upper third. Vagina comparatively

Kohler: New Amplirhagada species from the Kimberley

235

Figure 24. SEM photographs of Amplirhagada descartesana n.sp., paratype AMS C463749. ( A-D ) Shell: (A) apical whorl viewed from
above (scale 200 pm); ( B ) sculpture on first four whorls viewed from above (scale 1 mm); (C) sculpture, lateral view (scale 100 pm); (D)
close-up of sculpture, obliquely from above (scale 100 pm). ( E-G ) Radula: (E) central and lateral teeth (Scale 20 pm); (F) close-up of
central and inner lateral teeth (scale 10 pm); (G) outer lateral and inner marginal teeth (scale 10 pm). ( H) Jaw (scale 100 pm).

236

Records of the Australian Museum (2010) Vol. 62

Comparative remarks. The present material has been
labelled as “Amplirhagada sp. 74” by Solem. The species
is most readily distinguished from other species from the
Montesquieu Islands by its more turreted shell. Similar to A.
montesquieuana , its axial sculpture is more regular than that
of A. combeana and A. mckenziei. From A. montesquieuana
it differs by its larger size and more conical shape. The main
stimulatory pilaster is similar to that in A. montesquieuana
in shape but lacks “hooked toes”; broader than pilaster in
A. ponderi and A. mckenziei ; penial wall pustulation differs
markedly from that in A. montesquieuana.

Figure 25. Genitalia of Amplirhagada descartesana n.sp., paratype
WAM S41491 (20 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

short to moderately long, posteriorly inflated. Inner vaginal
wall with smooth longitudinal pilasters. Spermathecal duct
comparatively wide, internally with smooth longitudinal
pilasters. Spermathecal head elongately inflated, connected
with oviduct by connective tissue, internally smooth, with
delicate wall. Free oviduct comprising about half of anterior
part of oviduct, rather straight. Spermoviduct clearly longer
than anterior part of oviduct. Talon embedded in albumen
gland close to anterior end.

Radular morphology (Fig. 24E-G). Rectangular. Tooth
formula C+16-17+3-4+18-20. In average with 139±5.5
rows of teeth, 31.4+0.2 rows per mm (n = 2). Central teeth
with bluntly pointed, triangular mesocones, shorter than
base of tooth; ectocones small. Lateral teeth with bluntly
pointed, triangular mesocones, length equal to base of tooth;
ectocones small, endocones vestigial. Marginal teeth with
elongate mesocones; ectocones shorter and narrower than
mesocones, divided into two denticles; endocones smaller

Figure 26. Interior of penial chamber of Amplirhagada
descartesana n.sp., paratype FMNH 219276 (20 July, scale 5
mm). Compare with Fig. 4 for labelling of structures.

Amplirhagada hater ana Solem, 1981

Amplirhagada katerana Solem, 1981a: 198-201, figs. 37d,
41a, 43e-f.

Material examined. AMS C463692, WAM S36601 (preserved specimens,
Bonaparte Archipelago, Montague Sound, Rater’s Island, 14°26'51.6"S
125°31'07.1"E), WAM S36878 (preserved specimens, 14°26'52"S
125°31'13”E), AMS C463695, WAM S36879 (preserved specimens,
14°26'49"S 125°31T5"E), AMS C463696, WAM S36880 (preserved
specimens, 14°26'56.8"S 125°31'11.0"E) (Fig. 1).

Sealing strategy. Free sealer.

Shell (Fig. 27A-C, PI. 1.13). Semi-globose with medium
spire; solid. Periphery evenly rounded to angulate. Umbilicus
forming a chink or narrowly winding opening, 80-90 percent
concealed by columellar reflection. Background colour horn;
with diffuse to well marked, dark brown, moderately to very
thick sub-sutural and mod-whorl bands; bands may fuse with
each other concealing background colour completely; ventral
colour, outer lip and inner lip horn. Protoconch c. 2.3 mm in
diameter, comprising 2 whorls, sculptured by fine, indistinct
axial lirae. Teleoconch smooth except for axial growth lines.
Angle of aperture 30°; outer lip rounded, sharp to moderately
thick, slightly expanded, not or slightly reflected, basal and
palatal node absent. Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity deep, comprising one
whorl. Mottled pigmentation on mantle dark greyish brown.
Kidney extending about half of pallial cavity.

Genital morphology (Figs. 28-29). Penis straight, of
about same length as anterior part of oviduct. Vas deferens
forms simple loop or coils before entering penis. Penial
retractor muscle shorter than penis of same length. Penial
sheath evenly thick. Penial verge short (Vs- l A of penial
chamber), broad, with pointed tip. Penial wall covered by
small pustules, arranged in rows over entire length of inner
penial wall. Main stimulatory pilaster not differentiated.
Three pilasters are formed by rows of thickened pustules
comprising entire length of inner penial wall. Vas deferens
slightly undulated, entering penial sheath close to apical
portion. Vagina elongated, tubular. Inner vaginal wall with
smooth longitudinal pilasters. Spermathecal duct and head
not well-differentiated; duct moderately thick, internally
with smooth longitudinal pilasters; head globular to
elongately inflated, internally smooth, with delicate wall;
only tip of head connected with oviduct by connective

Kohler: New Amplirhagada species from the Kimberley

237

Figure 27. SEM photographs of Amplirhagada katerana. (A-C) Shell, AMS C463697: (A) apical whorl viewed from above (scale 200
pm); ( B ) sculpture on first three whorls viewed from above (scale 200 pm); (C) close-up of the peripheral keel and sculpture on last whorl
(scale 200 pm). ( D ) Jaw, WAM S36603 (scale 100 pm). (E-F) Radula, WAM S36603: (E) close-up of central and inner lateral teeth (scale
10 pm); (E) close-up of middle marginal teeth (scale 10 pm).

tissue. Free oviduct comprising more than half of anterior
part of oviduct, more or less straight. Spermoviduct slightly
longer than anterior part of oviduct. Talon embedded in
albumen gland close to anterior end of albumen gland.

Radular morphology (Fig. 27E-F). Rectangular. Tooth
formula C+14-20+3-4+15-22. In aerage with 126±7.0
rows of teeth, 30.7±0.1 rows per mm (n = 2). Central teeth
with bluntly pointed, triangular mesocones, shorter than
base of tooth; ectocones vestigial. Lateral teeth with bluntly
pointed, triangular mesocones, shorter than base of tooth;

ectocones and endocones vestigial. Marginal teeth with
rounded mesocones; ectocones shorter and narrower than
mesocones, simple; endocones vestigial.

Comparative remarks. Shell more conical in shape and
whorls more flattened in diameter than in any species listed
so far. Dark brown colour of most shells with pronounced
different ventral colour is very characteristic. Penial interior
with three longitudinal pilasters formed by enlarged wall
pustules is peculiar to this species. For comparison with
other species also see Solem (1981a: 198-199).

238

Records of the Australian Museum (2010) Vol. 62

Figure 28. Genitalia of Amplirhagada katerana AMS C463692 (27
July, scale 10 mm). Compare with Fig. 3 for labelling of structures.

Figure 29. Interior of penial chamber of Amplirhagada katerana
AMS C463692 (27 July, scale 5 mm) (general overview and
magnification of pustulation; drawing F.K.). Compare with Fig. 4
for labelling of structures.

Amplirhagada puescheli n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Montague Sound, unnamed island
near Prudhoe Island, 14°25’19"S 125°14’23"E; KC-075 (leg.
V. Kessner & A. Longbottom, 21 July 1988).

Type material. Holotype WAM S34608 (PI. 1.14). Paratypes AMS
C463750 (8 preserved specimens), FMNH 219290 (18 preserved
specimens), WAM S41486 (10 preserved specimens).

Etymology. Named in honour of Martin Piischel, acknow¬
ledging his indispensable help with producing most
illustrations used in this publication.

Shell (Fig. 32A-C, PL 1.14). Broadly conical with low
to medium spire; thin to solid (translucent). Periphery

slightly angulate; upper and basal sectors of whorls
rounded. Umbilicus forming a chink or narrowly winding
opening, 30-90 percent concealed by columellar reflection.
Background colour horn to yellowish brown; with diffuse
to well marked, brown sub-sutural and mid-whorl bands
being most conspicuous on last whorls; ventral colour horn;
outer lip colour same as shell; inner lip translucent, whitish.
Protoconch c. 2.5 mm in diameter, comprising 1.5 whorls,
smooth. Teleoconch smooth except of faint axial growth

Figure 30. Genitalia of Amplirhagada puescheli n.sp., paratype
AMS C463750 (21 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Figure 31. Interior of penial chamber of Amplirhagada puescheli
n.sp., paratype AMS C463750 (21 July, scale 5 mm). Compare with
Fig. 4 for labelling of stmctures.

Kohler: New Amplirhagada species from the Kimberley

239

Figure 32. SEM photographs of Amplirhagada puescheli, paratype AMS C463750. (A-C) Shell: (A) apical whorl viewed from above
(scale 200 pm); ( B ) sculpture on first three whorls viewed from above (scale 200 pm); (C) sculpture on last whorl (scale 200 pm). (D)
Jaw, broken (scale 100 pm). ( E-G ) Radula: (E) central and inner lateral teeth (scale 10 pm); (F) close-up of lateral teeth (scale 10 pm);
(G) outer lateral and inner marginal teeth (scale 20 pm).

lines. Angle of aperture 30°; outer lip rounded, sharp, slightly
expanded, slightly reflected; basal node of lip absent weak;
palatal node absent. Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending % whorl. Mantle pigmentation consist of black
spots. Kidney extending half of pallial cavity or more.

Genital morphology (Figs. 30-31). Penis curved or sharply
bent, of more or less of same length as anterior part of
oviduct. Vas deferens coils before entering penis. Penial

retractor muscle shorter than penis. Penial verge tiny (barely
visible), slender to spatulate with pointed tip. Penial wall
pustules small, elongated, arranged in rows over entire
length of inner penial wall. Main stimulatory pilaster absent.
Vas deferens entering penial sheath in upper third. Vagina
moderately wide, posteriorly inflated; inner vaginal wall
with undulating longitudinal pilasters. Spermatheca long,
clearly extending over base of spermoviduct. Spermathecal
duct internally with undulating longitudinal pilasters.
Spermathecal head globular to elongately inflated, connected
with oviduct by connective tissue, internally smooth with

240

Records of the Australian Museum (2010) Vol. 62

delicate to thin wall. Free oviduct shorter than half of
anterior part of oviduct, more or less straight. Spermoviduct
of same length as anterior part of oviduct. Talon embedded
in albumen gland close to anterior end of albumen gland.

Radular morphology (Fig. 32E-G). Rectangular. Tooth
formula C+12-15+2-3+19-20. In Average with 156±10
rows of teeth, 32.5±0.4 rows per mm (n = 2). Central teeth
with sharply pointed, triangular mesocones, shorter than
base of tooth; ectocones absent. Lateral teeth with bluntly
pointed, triangular mesocones, length equal to base of
tooth; ectocones and endocones vestigial. Marginal teeth
with triangular to rounded mesocones; ectocones shorter
and narrower than mesocones, simple, pointed; endocones
greatly reduced.

Comparative remarks. The present material has been
labelled as “Amplirhagada sp. 70” by Solem. Shell is rather
generic and similar to a number of species especially from
islands in the Admiralty Gulf. It differs by its sharply bent
penis with strongly developed pustulation of inner whorl
and absence of well-differentiated main pilaster and a rather
tiny verge.

Amplirhagada decora n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, Bigge Island, 4.3 km
north of Savage Hill, 14°35T5"S 125°11’05"E; KIS 1-41.
Vine thicket on lower and mid slopes north of small stream,
dolerite, on trees and bushes in wetter sections (leg. V.
Kessner, 17 February 2008).

Type material. Holotype WAM S34609 (PI. 1.15). Paratypes AMS
C463698 (8 preserved specimens, as holotype), WAM S36873 (17
preserved specimens, as holotype), AMS C463699 (4 preserved specimens,
14°35'13.3"S 125°11'10.5"E), WAM S36874 (9 preserved specimens,
14°35'13.3"S 125°11'10.5"E), AMS C463700 (shell, 14°35’15"S
125°H'05"E), WAM S36963 (2 shells, 14°35'15"S 125°H'05"E).

Additional, non-type material. WAM S36461-7, WAM S36652-4, WAM
S36872 (Bigge Island).

Etymology. From decora (Latin = elegant, beautiful) in
reference to its beautiful shell.

Shell (Fig. 35A-C, PL 1.15). Globose, with medium to
high spire; solid. Periphery angulate; upper sector of whorls
flattened to rounded; basal sector rounded. Umbilicus
completely concealed by columellar reflection. Background
colour brownish horn; with well defined, dark brown,
moderately broad sub-sutural and mid-whorl bands being
most conspicuous on last whorls; ventral colour horn; outer
lip colour same as shell; inner lip translucent. Protoconch
c. 2 mm in diameter, comprising 1.5 whorls, sculptured by
fine, indistinct axial lirae. Teleoconch with fine axial lirae,
rounded in cross-section, regularly spaced, spaces equal
to thickness of lirae, lirae distributed evenly across shell
surface, reduced underneath suture. Angle of aperture 30°,
outer lip rounded, sharp to moderately thick, not or slightly
expanded, not reflected; basal and palatal node absent.
Parietal wall of inner lip absent.

Figure 33. Genitalia of Amplirhagada decora n.sp., paratype AMS
C463699 (17 February, scale 10 mm) (drawing F.K.). Compare with
Fig. 3 for labelling of stmctures.

Pallial morphology. Pallial cavity deep, extending one
whorl. Mottled pigmentation on mantle dark grey. Kidney
extending about half of pallial cavity.

Genital morphology (Figs. 33-34). Penis straight, of more
or less same length as anterior pert of oviduct. Vas deferens
forms simple loop before entering penis. Penial retractor
muscle of about same length as penis. Penial sheath evenly
thin. Penial verge tiny (barely visible), with truncated tip.
Penial wall pustules of average size, densely arranged

Figure 34. Interior of penial chamber of Amplirhagada decora n.sp.,
paratype AMS C463698 (17 February, scale 5 mm) (drawing F.K.).
Compare with Fig. 4 for labelling of structures.

Kohler: New Amplirhagada species from the Kimberley

241

Figure 35. SEM photographs of Amplirhagada decora n.sp. (A-C) Shell, paratype AMS C463700: (A) apical whorl viewed from above
(scale 1 mm); ( B ) sculpture on first three whorls viewed from above (scale 1 mm); (C) sculpture on last whorl (scale 100 pm). (D) Jaw,
paratype AMS C463698 (scale 100 pm). ( E-G ) Radula, paratype AMS C463698: (E) central and lateral teeth (scale 20 pm); (F) close-up
of lateral teeth (scale 10 pm); (G) inner and middle marginal teeth (scale 20 pm).

in rows over entire length of inner penial wall; partly
fused to small elongated ridges. Main stimulatory pilaster
well-differentiated, forming a very large cone sculptured
by dense ridges comprising apical to median portion of
penial chamber; pilaster ridges support little hooks. Vas
deferens rather undulated, moderately thick, entering
penial sheath in upper third of penis. Vagina short, tubular,
posteriorly inflated; inner wall with longitudinal pilasters.
Spermathecal duct of medium thickness, internally with
smooth longitudinal pilasters. Spermathecal head globular

to elongately inflated, connected with oviduct by connective
tissue, internally smooth, with thin wall. Free oviduct
comprising more than half of anterior part of oviduct, rather
straight. Spermoviduct longer than anterior part of oviduct.
Talon embedded in albumen gland close to anterior end of
albumen gland.

Radular morphology (Fig. 35E-G). Rectangular. Tooth
formula C+16-18+0-3+20-21. In average 137.3+25.4
rows of teeth, 27.8±2.7 rows of teeth (n = 3). Central teeth

242

Records of the Australian Museum (2010) Vol. 62

D

100(jm

I-1

Figure 36. SEM photographs of Amplirhagada kessneri n.sp. (A-C) Shell, paratype AMS C463703: (A) apical whorl viewed from above
(scale 1 mm); ( B ) sculpture on second to forth whorl viewed from above (scale 1 mm); (C) detail of sculpture on mid of last whorl (scale
200 pm). (D) Jaw, paratype AMS C463701 (scale 100 pm). (E-G) Radula, paratype AMS C463701: (E) central and lateral teeth (scale
20 pm); ( F) close-up of lateral teeth (scale 10 pm); (G) inner and middle marginal teeth (scale 20 pm).

with bluntly pointed, triangular to ovate mesocones, shorter
than base of tooth; ectocones vestigial. Lateral teeth with
bluntly pointed, triangular to ovate mesocones, shorter than
base of tooth; ectocones and endocones vestigial. Marginal
teeth with broad, flattened mesocones; ectocones shorter
and narrower than mesocones, divided into two denticles,
endocones reduced in size.

Comparative remarks. Its highly elevated, dome-shaped
shell being characteristic (H/D larger than in many other
species with more conical shells). Comparatively large, cone-
shaped main pilaster, rather large pustules of inner penial
wall and tiny verge are diagnostic. This species was referred
to as “Amplirhagada sp. 23” by Solem (1991).

Amplirhagada kessneri n.sp.

Type locality (Fig. 1). Western Australia, northwestern Kim¬
berley, Bonaparte Archipelago, Bigge Island, 4.3 km north of
Savage Hill, 14°35’15"S 125°11'05"E; KIS 1-41. Vine thicket
on mid slopes north of small stream, dolerite, on trees and
bushes in drier sections (leg. V. Kessner, 17 February 2008).

Type material. Holotype WAM S34610 (PI. 1.16). Paratypes AMS
C463701 (4 preserved specimens, as holotype), WAM S36875 (10 preserved
specimens, as holotype), WAM S36477 (5 shells, 14°35'16"S 125° 11'02"E),
AMS C463702 (9 preserved specimens, 14°35'16"S 125°11'02"E),
AMS C C.463703 (2 shells, 14°35'16"S 125°11'02"E), WAM S36658
(10 preserved specimens, 14°35'16"S 125°11'02"E), WAM S36876 (16
preserved specimens, 14°35'18.5"S 125°10'54.7"E),WAMS36475 (3 shells,
14°35'16"S 125° 11’06"E), WAM S36476 (6 shells, 14°35’14"S 125°11'08"E).

Kohler: New Amplirhagada species from the Kimberley

243

Figure 37. Genitalia of Amplirhagada kessneri n.sp., paratype
WAM S36876 (17 February, scale 10 mm). Compare with Fig. 3
for labelling of structures.

Etymology. Named in honour of Vince Kessner in recognition
of his lasting achievements as one of Australia’s finest experts
in the field.

Shell (Fig. 36A-C, PL 1.16). Semi-globose to conical with
low spire; thin to solid (translucent). Periphery evenly
rounded to slightly angulate; upper sector of whorls rounded,
basal sector somewhat flattened. Umbilicus open, forming a
chink or narrowly winding opening, 30-90 percent concealed
by columellar reflection. Background colour horn to light
brown, exceptionally dark brown; sub-sutural and mid-whorl
bands absent or diffuse to well marked, brown, thin to
moderately thick, most conspicuous on last whorls; ventral
colour horn to light brown; outer lip colour lighter than shell,
horn; inner lip translucent. Protoconch c. 2 mm in diameter,
comprising 2 whorls, almost smooth. Teleoconch with
regularly spaced axial lirae becoming somewhat oblique on
last whorl. Angle of aperture 45°; outer lip rounded, sharp to
moderately thick, slightly expanded, not or slightly reflected,
basal node absent or very weak, palatal node absent. Parietal
wall of inner lip absent.

Pallial morphology. Pallial cavity deep, extending one
whorl. Mottled mantle pigmentation dark grey to black.
Kidney extending about half of pallial cavity.

Genital morphology (Figs. 37-38). Penis straight, more or
less of same length as anterior part of oviduct. Vas deferens
forms simple loop before entering penis. Penial retractor
muscle longer than penis. Penial sheath evenly thin. Penial
verge short (<'/« penial chamber), slender to spatulate, with
truncated tip. Penial wall pustules small, densely arranged in
rows over entire length of inner penial wall. Main stimulatory
pilaster well-differentiated, forming a large, elongated cone,
sculptured by smooth horizontal ridges, comprising apical
to median portion of penial chamber. Vas deferens entering
penial sheath in upper third. Vagina rather long, tubular; inner
vaginal wall densely covered by tiny pustules. Spermathecal
duct moderately thick, internally with longitudinal rows
of densely packed ciliae. Spermathecal head globular to
elongately inflated, connected with oviduct by connective
tissue, internally smooth, densely ciliated, with thin wall.
Free oviduct comprising±half of anterior part of oviduct,
straight. Spermoviduct longer to much longer than anterior

part of oviduct. Talon embedded in albumen gland close to
anterior end.

Radular morphology (Fig. 36E-G). Rectangular. Tooth
formula C+14-17+0-4+18-21. In average with 117±2.9
rows of teeth, 27.6±0.7 rows per mm (n = 3). Central teeth
with sharply pointed, triangular mesocones, shorter than
base of tooth; ectocones tiny. Lateral teeth with sharply
pointed, triangular mesocones, length equal to base of tooth;
ectocones tiny, endocones vestigial. Marginal teeth with
broad, flattened mesocones; ectocones shorter and narrower
than mesocones; endocones reduced in size.

Comparative remarks. Differs from A. decora , its congener
from Bigge Island, by flatter, uniformly coloured shell,
much smaller main pilaster and finer pustulation of inner
penial wall.

244

Records of the Australian Museum (2010) Vol. 62

Amplirhagada berthierana n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, Berthier Island,
14°29'49"S 124°59'34.1"E (leg. C. Whisson, 11 November
2006).

Type material. Holotype WAM S34611 (PL 1.17). Paratypes AMS
C463754 (8 preserved specimens), WAM S33029 (10 preserved
specimens).

Etymology. In reference to Berthier Island, where this
species occurs.

Sealing strategy. Rock sealer.

Shell (Fig. 41A-C, PI. 1.17). Globose to broadly conical with
medium high spire; thick. Whorls evenly rounded. Umbilicus
open, forming a chink, 95 percent concealed by columellar
reflection. Background and ventral colour cream; banded.
Sub-sutural band well defined, red-brown. Mid-whorl band
well marked, red brown, moderately broad, visible on entire
shell becoming more diffuse on upper whorls, blends with
sub-sutural band to cover entire surface of whorl in reddish
brown towards tip of shell; outer lip colour white; inner
lip translucent, white. Protoconch c. 2.2 mm in diameter,
comprising 1.7 whorls, with fine, indistinct axial lirae.
Teleoconch with strong axial growth lines. Angle of aperture
45°; outer lip rounded, moderately thick to thick, slightly
expanded, slightly reflected; basal node of lip weak; palatal
node absent. Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl. Pigmentation on mantle comprising
sparsely distributed, dark brown spots. Kidney extending
more than half of pallial cavity.

Genital morphology (Figs. 39-40). Penis curved, thick,
more or less of same length as anterior part of oviduct. Vas
deferens coils once before entering penis. Penial retractor
muscle stubby to short. Penial sheath evenly thick. Penial

Figure 39. Genitalia of Amplirhagada berthierana n.sp., paratype
AMS C463754 (11 November, scale 10 mm). Compare with Fig.
3 for labelling of stmctures.

verge short (<14 penial chamber), broad with rounded tip.
Penial wall pustules small, arranged in rows over entire
length of inner penial wall. Main stimulatory pilaster well-
differentiated, elongated, cone-shaped, sculptured by smooth
horizontal ridges, comprising apical two thirds of penial
chamber. Vas deferens entering penial sheath in upper third.
Vagina moderately thick, medially inflated; inner vaginal wall
with undulating longitudinal pilasters. Spermatheca long,
clearly extending over base of spermoviduct. Spermathecal
duct wide, internally with undulating longitudinal pilasters.
Spermathecal head elongately inflated, connected with
oviduct by connective tissue, internally smooth with delicate
wall. Free oviduct comprising more or less half of anterior
part of oviduct, straight. Spermoviduct longer than anterior
part of oviduct. Talon embedded in albumen gland at junction
with spermoviduct.

Radular morphology (Fig. 41E-G). Rectangular. Tooth
formula C+13-15+3-4+20-24. In average 133±5.0 rows
of teeth, 29.2+0.2 rows per mm (n = 2). Central teeth with
sharply pointed, triangular to ovate mesocones, shorter
than base of tooth; ectocones vestigial. Lateral teeth with
sharply pointed, triangular mesocones, length equal to base
of tooth; ectocones and endocones vestigial. Marginal teeth
with triangular mesocones; ectocones shorter and narrower
than mesocones; endocones reduced in size.

Comparative remarks. Distinctive species; its broadly
conical, almost dome-like shell with widely rounded, almost
depressed last whorl with pronounced spiral banding, and
the corrugated, large main pilaster are diagnostic.

Figure 40. Interior of penial chamber of Amplirhagada berthierana
n.sp., paratype AMS C463754 (11 November, scale 5 mm).
Compare with Fig. 4 for labelling of structures.

Kohler: New Amplirhagada species from the Kimberley

245

Figure 41. SEM photographs of Amplirhagada berthierana n.sp., paratype AMS C463754. (A-C) Shell: (A) apical whorl viewed from
above (scale 200 pm); ( B ) sculpture on second to forth whorl viewed from above (scale 200 pm); (C) sculpture on upper part of last whorl
(scale 200 pm). (D) Jaw (scale 100 pm). (E-G) Radula: (E) central and lateral teeth (scale 20 pm); (E) close-up of lateral teeth (scale 20
pm); (G) outer lateral and inner marginal teeth (scale 20 pm).

Amplirhagada lamarckiana n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, York Sound, Lamarck Island,
14°46'50"S 125°r23"E; KC-042 (leg. V. Kessner & A.
Longbottom, 14 July 1988).

Type material. Holotype WAM S34612 (PL 1.18). Paratypes AMS
C463755 (5 preserved specimens), FMNH 219163 (15 preserved
specimens), WAM S41485 (8 preserved specimens).

Etymology. In reference to Lamarck Island, where this
species occurs.

Shell (Fig. 42A-C, PI. 1.18). Semi-globose to broadly
conical, with medium high spire; thin (translucent). Periphery
evenly rounded to slightly angulate; upper sector of whorls
rather flattened, basal sector rounded. Umbilicus forming a
chink, 90-100 percent concealed by columellar reflection.
Background colour light brownish horn, banded. Sub-sutural
and mid-whorl bands diffuse to well defined, light brown,

246

Records of the Australian Museum (2010) Vol. 62

Figure 42. SEM photographs of Amplirhagada lamarckiana n.sp., holotype WAM S34612. (A-Q Shell: (A) apical whorl viewed from
above (scale 200 pm); ( B ) sculpture on second to forth whorl viewed from above (scale 200 pm); (C) sculpture on upper part of last whorl
(scale 200 pm). ( D ) Jaw (scale 100 pm). (E-F) Radula: (E) central and lateral teeth (scale 20 pm); (F) middle marginal teeth (scale 10 pm).

thin, visible on last whorl(s) only; ventral colour horn; outer
lip colour same as shell; inner lip translucent. Protoconch
c. 3 mm in diameter, comprising 1.7 whorls, with fine and
indistinct lirae. Teleoconch with rather strong axial growth
lines. Angle of aperture 45°; outer lip rounded, sharp to
moderately thick, slightly expanded, reflected, basal node
of lip absent or weak; palatal node absent. Parietal wall of
inner lip absent or inconspicuous.

Pallial morphology. Pallial cavity deep, extending 3 A whorl.
Pigmentation on mantle mottled, black. Kidney extending
more than half of pallial cavity.

Genital morphology (Figs. 43-44). Penis straight to
curved, of about same length as anterior part of oviduct.
Vas deferens coils before entering penis. Penial retractor
muscle shorter than penis. Penial sheath delicate. Penial
verge tiny (barely visible), slender to spatulate, with pointed
tip. Penial wall small, arranged in rows over entire length
of inner penial wall. Main stimulatory pilaster elongated,
sculptured by horizontal ridges that support little hooks,
comprising apical to median portion of penial chamber. Two
to three additional pilasters are covered by undifferentiated
pustules. Vas deferens entering penial sheath half way up.
Vagina moderate, posteriorly inflated; inner vaginal wall with

Kohler: New Amplirhagada species from the Kimberley

247

Figure 43. Genitalia of Amplirhagada lamarckiana n.sp., holotype
WAM S34612 (14 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

smooth longitudinal pilasters. Spermathecal duct moderately
thick, internally with smooth longitudinal pilasters.
Spermathecal head globular, connected with oviduct by
connective tissue, internally smooth, with delicate wall.
Free oviduct comprises more or less half of anterior part of
oviduct, straight. Spermoviduct longer than anterior part of
oviduct. Talon embedded in albumen gland at junction with
spermoviduct.

Radular morphology (Fig. 42E-F). Rectangular. Tooth
formula C+14+4+18. With 157 rows of teeth, 27.9 rows per
mm (n = 1). Central teeth with bluntly pointed, triangular
to ovate mesocones, shorter than base of tooth; ectocones
vestigial. Lateral teeth with sharply pointed, triangular
mesocones, length equal to base of tooth; ectocones
and endocones vestigial. Marginal teeth with triangular
mesocones; ectocones shorter and narrower than mesocones,
occasionally divided into two denticles, endocones reduced
in size.

Comparative remarks. Pustulation of inner penial wall
somewhat similar to A. indistincta and A. solemiana,
which lack a well-differentiated main pilaster and differ
by more conical shape of shell. Main pilaster differs in
shape and surface pattern from the two species from Bigge
Island. Material of the present species has been labelled as
“Amplirhagada sp. 68” by Solem.

Amplirhagada anderdonensis n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Prince Frederick Harbour, Anderdon
Islands, unnamed island c. 8 km ENE of Cape Torrens,
14°57'49"S 125°09'30"; KC-084 (leg. V. Kessner & A.
Longbottom, 22 July 1988).

Type material. Holotype WAM S34613 (PL 1.19). AMS C463756 (5
preserved specimens), FMNH 219330 (16 preserved specimens), WAM
S41492 (10 preserved specimens).

Etymology. In reference to Anderdon Islands, where this
species occurs.

Sealing strategy. Rock sealer.

Figure 44. Interior of penial chamber of Amplirhagada lamarckiana
n.sp., paratype FMNH 219163 (14 July, scale 5 mm). Compare with
Fig. 4 for labelling of structures.

Figure 45. Genitalia of Amplirhagada anderdonensis n.sp., paratype
FMNH 219330 (22 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

248

Records of the Australian Museum (2010) Vol. 62

Figure 46. SEM photographs of Amplirhagada anderdonensis n.sp. (A-C) Shell, paratype AMS C463756: (A) apical whorl viewed from
above (scale 200 pm); ( B ) sculpture on second and third whorl viewed from above (scale 200 pm); (C) sculpture on upper part of last
whorl (scale 100 pm). (D) Jaw (scale 100 pm). (E-G) Radula, paratype FMNH 219330: (E) central and lateral teeth (scale 20 pm); (F)
close-up of lateral teeth (scale 10 pm); (G) outer lateral and inner marginal teeth (scale 10 pm).

Shell (Fig. 46A-C, PL 1.19). Broadly conical with medium
high spire, solid. Periphery evenly rounded to slightly
angulate; upper and basal sectors of whorls rounded.
Umbilicus forming a chink, 90-100 percent concealed by
columellar reflection. Background colour greenish ochre to
yellowish brown; sub-sutural and mid-whorl band absent or
diffuse, darker than background, thin, most conspicuous on
last whorl; ventral colour cream; outer lip colour same as
shell; inner lip translucent, whitish. Protoconch c. 2 mm in
diameter, comprising 1.7 whorls, almost smooth. Teleoconch
with inconspicuous axial growth lines, last whorl smooth.

Angle of aperture 30°; outer lip rounded, moderate to thick,
slightly expanded, slightly reflected; basal node of lip
absent or weak; palatal node absent. Parietal wall of inner
lip inconspicuous.

Pallial morphology. Pallial cavity deep, extending one
whorl. Pigmentation on mantle consists of sparse greyish to
brown spots. Kidney extending about half of pallial cavity.

Genital morphology (Figs. 45, 47). Penis straight to
slightly curved, more or less of same length as anterior part

Kohler: New Amplirhagada species from the Kimberley

249

of oviduct. Vas deferens coils once before entering penis.
Penial retractor muscle shorter than penis. Penial verge
moderately long (Vs- l A penial chamber), slender to spatulate,
with pointed tip. Penial wall pustules small, arranged in
rows. Main stimulatory pilaster differentiated, relatively thin,
cone-shaped, with horizontal ridged that support little hooks,
comprising apical to median portion of penial chamber.
Vas deferens entering penial sheath in upper third. Vagina
moderately long, tubular, posteriorly inflated; inner vaginal
wall with smooth longitudinal pilasters. Spermatheca very
short, barely reaching base of spermoviduct. Spermathecal
duct normal, internally with smooth longitudinal pilasters.
Spermathecal head globular, connected with oviduct by
connective tissue, internally smooth, with delicate wall.
Free oviduct comprising more than half of anterior part of
oviduct, more or less straight. Spermoviduct longer than
anterior part of oviduct. Talon embedded in albumen gland
at junction with spermoviduct.

Radular morphology (Fig. 46E-G). Rectangular. Tooth
formula C+14-15+2+20-23. In average with 145±12.0
rows of teeth, 31.6±4.2 rows per mm (n = 2). Central teeth
with bluntly pointed, ovate mesocones, shorter than base of
tooth; ectocones vestigial. Lateral teeth with bluntly pointed,
ovate mesocones, length equal to base of tooth; ectocones
small, endocones vestigial. Marginal teeth with triangular
to rounded mesocones; ectocones shorter and narrower than
mesocones; endocones reduced in size.

Comparative remarks. Shell colour very distinctive, similar
only to A. tricenaria (see below). Penis proportionally
smaller than in other species. In addition, it differs from A.
lamarckiana by finer pustulation of inner penial wall and
smaller main pilaster. Material of the present species has
been labelled as ‘Amplirhagada sp. 75” by Solem.

Amplirhagada tricenaria n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Prince Frederick Harbour, north of
Hunter River mouth, 15 o 0F54"S 125°23’13"E; KC-080 (leg.
V. Kessner & A. Longbottom, 22 July 1988).

Type material. Holotype WAM S34614 (PL 1.20). Paratypes AMS
C463757 (2 preserved specimens), FMNH 219322 (6 preserved
specimens), WAM S41476 (4 preserved specimens).

Etymology. From tricenaria (Latin = thirty), in reference
to manuscript name assigned to this species by Solem
(“Amplirhagada n.sp. 30”).

Shell (Fig. 49A-D, PL 1.20). Semi-globose, with medium
high spire; thin to solid (translucent). Periphery slightly
angulate; upper and basal sectors of whorls rounded.
Umbilicus forming a chink, 90-100 percent concealed by
columellar reflection. Background colour dark hom to ochre;
sub-sutural and mid-whorl bands diffuse to well defined,
yellowish brown, thin to moderately broad, most conspicuous
on last whorls; ventral colour horn; outer lip colour same
as shell; inner lip yellowish white. Protoconch c. 2.6 mm in
diameter, comprising 2 whorls, almost smooth. Teleoconch
with pronounced axial growth lines, last whorl almost
smooth. Angle of aperture 30°, outer lip rounded, sharp to
moderatelv thick, slightlv expanded, slightly reflected, basal

n.sp., paratype FMNH 219332 (22 July, scale 5 mm). Compare
with Fig. 4 for labelling of stmctures.

250

Records of the Australian Museum (2010) Vol. 62

Figure 49. SEM photographs of Amplirhagada tricenaria n.sp. ( A-D ) Shell, paratype AMS C463757: (A) apical whorl viewed from above
(scale 200 pm); ( B ) sculpture on second to third whorl viewed from above (scale 200 pm); (C) suture between last and penultimate whorl
with details of sculpture (scale 100 pm); ( D ) surface of last whorl, viewed rectangularly from above (Scale 200 pm). (E) Jaw, paratype
FMNH 219322 (scale 100 pm). ( F-G ) Radula, paratype FMNH 219332: ( F) central and lateral teeth (scale 20 pm); (G) outer lateral and
inner marginal teeth (scale 20 pm).

node of lip absent or weak, palatal node absent. Parietal wall
of inner lip inconspicuous.

Pallial morphology. Pallial cavity deep, extending one
whorl. Pigmentation on mantle mottled, greyish or brownish
(probably leached in actual specimens). Kidney extending
about half of pallial cavity.

Genital morphology (Figs. 48, 50). Penis straight, more or
less of same length as anterior part of oviduct. Vas deferens
forms simple loop before entering penis. Penial retractor

muscle shorter than penis. Penial verge very short (barely
visible), slender to spatulate with pointed tip. Penial wall
pustules small, elongated, arranged in rows on apical to
median portion of penial chamber. Base of inner penial walls
with many smooth longitudinal pilasters. Main stimulatory
pilaster well-differentiated, elongate to cone-shaped,
sculptured by smooth ridges, comprising apical to median
portion of penial chamber. Vas deferens entering penial
sheath in upper third. Vagina of medium length and thickness,
tubular, posteriorly slightly inflated. Inner vaginal wall with

Kohler: New Amplirhagada species from the Kimberley

251

Figure 50. Genitalia of Amplirhagada tricenaria n.sp., paratype
AMS C463757 (22 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Figure 51. Genitalia of Amplirhagada regia n.sp., paratype WAM
S36647 (9 August; scale 10 mm). Compare with Fig. 3 for labelling
of structures.

smooth longitudinal pilasters. Spermathecal duct moderately
thick, internally with smooth longitudinal pilasters.
Spermathecal head globular to elongately inflated, connected
with oviduct by connective tissue, internally smooth, with
thin wall. Free oviduct comprising less than half of anterior
part of oviduct, more or less straight. Spermoviduct longer
than anterior part of oviduct. Talon embedded in albumen
gland at junction with spermoviduct.

Radular morphology (Fig. 49F-G). Rectangular. Tooth
formula C+15-17+3+18-23. In average with 152±3.0 rows
of teeth, 28.1±0.8 rows per mm (n = 2). Central teeth with
sharply pointed, ovate mesocones, shorter than base of
tooth; ectocones vestigial. Lateral teeth with bluntly pointed,
ovate mesocones, length equal to base of tooth; ectocones
small, endocones vestigial. Marginal teeth with triangular
mesocones; ectocones shorter and narrower than mesocones;
endocones reduced in size.

Comparative remarks. This species has a very distinctive
colour, similar only to A. anderdonensis , which differs by
smaller shell size, less globose shell shape, higher position
of peripheral angulation. Rather flattened diameter of whorls
and very broadly conical shape of shell are diagnostic.
Material of the present species has been labelled as
“Amplirhagada sp. 30” by Solem.

Amplirhagada regia n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, central section of Boongaree Island,
15°04T5"S 125°11T4"E; KIS 1-30. Dry vine thicket on west
facing sandstone scree below escarpment, under rocks (leg.
M. Shea, 9.8.2007).

Type material. Holotype WAM S34615 (PI. 1.21). Paratypes AMS
C463705 (4 preserved specimens, as holotype), WAM S36647 (3 preserved
specimens, as holotype), WAM S36449 (6 preserved specimens, 15°04'00"S
125°11'H"E), AMS C463704 (6 shells, 15°04’31"S 125°11'07"E), WAM
S36480 (12 shells, 15°04'31"S 125°H'07"E).

Additional, non-type material. WAM S36479, WAM S36481-5, WAM
S36648 (Boongaree Island).

Figure 52. Interior of penial chamber of Amplirhagada regia n.sp.,
paratype WAM S36647 (9 August; scale 5 mm). Compare with Fig.
4 for labelling of structures.

252

Records of the Australian Museum (2010) Vol. 62

Figure 53. SEM photographs of Amplirhagada regia n.sp. (A-D) Shell, paratype AMS C463704: (A) apical whorl viewed from above (scale
200 pm); ( B ) sculpture on second to forth whorl viewed from above (scale 1 mm); (C) surface of last whorl (scale 1 mm); (D) close-up
showing axial growth lines on last whorl (Scale 100 pm). ( E ) Jaw, paratype WAM S36647 (scale 100 pm). (E-G) Radula, paratype WAM
S36647: ( E ) close-up of lateral teeth (scale 10 pm); ( F) close-up of central and inner lateral teeth (scale 10 pm); (G) close-up of middle
marginal teeth (scale 10 pm).

Etymology. From regia (Latin = royal), referring to fact that
this species is among the largest of the genus.

Sealing strategy. Free sealer.

Shell (Fig. 53A-D, PI. 1.21). Globose with medium high to
high spire; solid to thick. Periphery angulated; upper sector
of whorls rather flattened, basal sector rounded. Umbilicus
forming chink or narrowly winding opening, 60-100 percent
concealed by columellar reflection. Background colour
uniform, creamish horn; outer lip purple; inner lip pale
purple. Protoconch c. 2.2 mm in diameter, comprising 1.5

whorls, sculpture almost smooth. Teleoconch smooth except
for faint axial growth lines. Angle of aperture 45; outer lip
simple rounded; moderate; slightly expanded; not reflected,
or slightly reflected; basal node of lip present; weak; palatal
node absent. Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity deep, extending one
whorl. Pigmentation on mantle comprising brownish grey
spots. Kidney extending about half of pallial cavity.

Genital morphology (Figs. 51-52). Penis straight, more or
less of same length as anterior part of oviduct. Vas deferens

Kohler: New Amplirhagada species from the Kimberley

253

forms simple loop before entering penis. Penial retractor
muscle shorter than or equal to penis length. Penial sheath
evenly thin. Penial verge short (cVs penial chamber), slender
to spatulate, with pointed tip. Penial wall pustules absent.
Main stimulatory pilaster absent. Two longitudinal pilasters
present at basal portion of penial chamber, two additional
weakly developed pilasters visible at apical portion. Vas
deferens entering penial sheath in upper third. Vagina short
to moderately long, tubular. Inner vaginal wall with smooth
longitudinal pilasters. Spermathecal duct moderately wide,
internally with smooth longitudinal pilasters. Spermathecal
head globular to elongately inflated, connected with oviduct
by connective tissue, internally entirely smooth, with delicate
wall. Free oviduct comprising about half of anterior part of
oviduct, more or less straight. Spermoviduct of same length
as anterior part of oviduct. Talon embedded in albumen gland
close to anterior end of albumen gland.

Radular morphology (Fig. 53F-H). Rectangular. Tooth
formula C+20+3+20. In average with 147±2.5 rows of
teeth, 29.6±1.7 rows per mm (n = 2). Central teeth with
sharply pointed, triangular mesocones, shorter than base
of tooth; ectocones vestigial. Lateral teeth with sharply
pointed, triangular mesocones, length equal to base of tooth;
ectocones and endocones vestigial. Marginal teeth with
elongate mesocones; ectocones shorter and narrower than
mesocones; endocones greatly reduced in size.

Comparative remarks. Diagnostic features are the broadly
conical to dome-shaped shell, smooth inner penial wall with
lack of pustulation and main pilaster.

Amplirhagada boongareensis n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Boongaree Island, central section,
15°04'36"S 125°11T2"E; KIS 1-28. Rainforest on north
facing sandstone scree below escarpment, under rocks (leg.
M. Shea, 8.8.2007).

Type material. Holotype WAM S34616 (PI. 1.22). Paratypes WAM S36655

Figure 54. Genitalia of Amplirhagada boongareensis n.sp., paratype
WAM S36655 (8 August, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

(3 preserved specimens, as holotype), AMS C463707 (8 shells, 15°04'31"S
125°11'07''E), WAM S36455 (12 shells, 15°04’31"S 125°11'07"E),
WAM S36456 (15 shells, 15°04'31"S 125 o ll'07"E), AMS C463706 (6
shells, 15°04'16"S 125°10’47"E), WAM S36458 (8 shells, 15°04'16"S
125°10'47"E), WAM S36656 (3 preserved specimens, 15°04'09"S
125°10'49"E).

Additional, non-type material. WAM S36457, WAM S36459-60
(Boongaree Island).

Etymology. In reference to Boongaree Island, where this
species occurs.

Shell (Fig. 56A-D, PI. 1.22). Semi-globose, with rather high
spire; solid. Periphery slightly to clearly angulate; upper
sector slightly shouldered, basal sector rounded. Umbilicus
forming chink or narrowly winding opening, 50-100 percent
concealed by columellar reflection. Background colour light
brown; sub-sutural and mid-whorl bands may be absent;
if present, bands are diffuse to well marked, dark reddish
brown, thin to moderately broad, most conspicuous on last
whorl; bands may blend into each other and covering entire
surface of tip of the shell with purplish brown colour; ventral
colour horn; outer lip colour same as shell; inner lip colour
blends from reddish brown near parietal wall of shell into
horn in outer sections. Protoconch c. 2.3 mm in diameter,
comprising 2 whorls, with fine, indistinct axial lirae.
Teleoconch with fine axial lirae, rounded in cross-section,
spacing regular, spaces equal to thickness of elements, evenly
distributed across shell surface, reduced underneath suture.
Angle of aperture 30°; outer lip rounded to slightly angulate,
with moderate to thick parietal notch at lower margin, slightly
expanded, not or slightly reflected, basal node of lip present,
palatal node absent.

Figure 55. Interior of penial chamber of Amplirhagada boongare¬
ensis n.sp., paratype WAM S36655 (8 August, scale 5 mm) (drawing
by F.K.). Compare with Fig. 4 for labelling of structures.

254

Records of the Australian Museum (2010) Vol. 62

Figure 56. SEM photographs of Amplirhagada boongareensis n.sp. (A-D) Shell, paratype AMS C463706: (A) apical whorl viewed from
above (scale 200 pm); (B) sculpture on second and third whorl viewed from above (scale 1 mm); (C) close-up showing axial sculpture on
last whorl (scale 100 pm); (D) close-up showing axial periphery of last whorl (Scale 1 mm). (E) Jaw, paratype WAM S36655 (scale 100
pm). ( E-G ) Radula, paratype WAM S36655: (E) central and inner lateral teeth (scale 20 pm); (E) outer lateral and inner marginal teeth
(scale 10 pm); (G) close-up of inner and middle marginal teeth (scale 10 pm).

Pallial morphology. Pallial cavity deep, extending one
whorl; mantle pigmentation mottled, dark grey. Kidney
extending about half of pallial cavity.

Genital morphology (Figs. 54-55). Penis straight to curved,
more or less of same length as anterior part of oviduct. Vas
deferens forms simple loop before entering penis. Penial
retractor muscle shorter than penis. Penial verge very short
(barely visible), spatulate, with pointed tip. Penial wall
pustules of average size, dense, randomly arranged on

entire length of inner penial wall. Main stimulatory pilaster
well-differentiated; large, cone-shaped, covered by smooth,
flattened pustules arranged in horizontal rows, comprising
apical to median portion of penial chamber. Two additional
pilasters comprise entire length of penial chamber, sup¬
porting undifferentiated pustulation. Vas deferens entering
penial sheath in upper third. Vagina moderately wide,
tubular; inner vaginal wall with smooth longitudinal pilasters.
Spermathecal duct of medium width, internally with smooth
longitudinal pilasters. Spermathecal head elongately inflated,

Kohler: New Amplirhagada species from the Kimberley

255

connected with oviduct by connective tissue, internally
entirely smooth, with delicate wall. Free oviduct comprising
more than half of anterior part of oviduct, zig-zag-folded
underneath entrance to spermoviduct. Spermoviduct longer
than anterior part of oviduct. Talon embedded in albumen
gland close to anterior end of albumen gland.

Radular morphology (Fig. 56F-G). Rectangular. Tooth
formula C+16+3+20. With 165 rows of teeth, 19.9 rows per
mm (n = 1). Central teeth with sharply pointed, triangular
mesocones, shorter than base of teeth; ectocones vestigial.
Lateral teeth with sharply pointed, triangular to ovate
mesocones; ectocones and endocones vestigial. Marginal
teeth with triangular mesocones; ectocones smaller and
narrower than mesocones; endocones smaller than ectocones.

Comparative remarks. His report that it also occurs on the
opposite mainland requires confirmation. The bee-hive shape
of the shell and its dark purplish brown colour are diagnostic.
Amplirhagada regia from Boongaree Island also differs by
absence of penial wall pustules or main pilaster. This species
was referred to as “Amplirhagada sp. 27” by Solem (1991).

Amplirhagada gibsoni n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Boongaree Island, central section,
15 o 04'15"S 125°11T4"E; KIS 1-30. Dry vine thicket on west
facing sandstone scree below escarpment, under rocks (leg.
M. Shea, 09 August 2007).

Type material. Holotype WAM S34617 (PL 2.1). Paratypes AMS
C463709 (1 preserved specimen, as holotype), WAM S36650 (2
preserved specimens, as holotype), WAM S36468 (6 shells, 15°04'15"S
125°11T4"E), AMS C463708 (4 shells, 15°04'36"S 125°11'18"E), WAM
S36469 (10 shells, 15°04'36"S 125 0 11'18"E), WAM S36651 (1 preserved
specimen, 15°04’36"S 125°11'18"E).

Etymology. Named in honour of Lesley Gibson, Western
Australian Department of Environment and Conservation,
in recognition of her support for my work.

Figure 57. Genitalia of Amplirhagada gibsoni n.sp., paratype
AMS C463709 (8 August, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Shell (Fig. 59A-C, PL 2.1-2). Semi-globose to conical with
medium high spire; thin (translucent) to solid. Periphery
evenly rounded to slightly angulate; upper and basal sectors
of whorls rounded. Umbilicus open, forming a narrowly
winding chink, about 90 percent concealed by columellar
reflection. Background colour light brownish to horn;
uniform. Outer lip colour same as shell, inner lip colour
white. Protoconch c. 2.2 mm in diameter, comprising 2
whorls, with fine and distinct axial lirae. Teleoconch with
fine axial lirae, rounded in cross-section, regularly spaced,
spaces equal to thickness of lirae, evenly distributed across
shell surface, reduced underneath suture; across whorls of
shell. Angle of aperture 30°; outer lip rounded, sharp to
moderately thick, slightly expanded, slightly reflected; basal
node absent or very weak, palatal node absent. Parietal wall
of inner lip inconspicuous.

Pallial morphology. Pallial cavity deep, extending one
whorl; mantle pigmentation mottled, black. Kidney
extending about half of pallial cavity.

Genital morphology (Figs. 57-58). Penis straight, more or
less of same length as anterior part of oviduct. Vas deferens
forms simple loop before entering penis. Penial retractor
muscle as long as penis. Penial sheath evenly thick. Penial
verge very long (c. Vs penial chamber), broad, with pointed
tip. Penial wall pustules of normal size, randomly and densely
arranged over entire length of inner penial wall. At base of
penial chamber fused rows of pustules form 3-4 smooth
longitudinal pilasters. Main stimulatory pilaster absent.
Vas deferens entering penial sheath in upper third. Vagina

256

Records of the Australian Museum (2010) Vol. 62

Figure 59. SEM photographs of Amplirhagada gibsoni n.sp. ( A-D ) Shell, paratype AMS C463708: (A) apical whorl viewed from above
(scale 1 mm); ( B ) sculpture on second to forth whorl viewed from above (scale 1 mm); (C) close-up showing axial periphery of last whorl
(Scale 200 pm); ( D) Jaw, paratype AMS C463709 (scale 100 pm). (E-F) Radula, paratype AMS C463709: (E) central and inner lateral
teeth (scale 20 pm); ( F ) outer lateral and inner marginal teeth (scale 20 pm).

tubular; inner vaginal wall with smooth longitudinal pilasters.
Spermathecal duct moderately wide, internally with smooth
longitudinal pilasters. Spermathecal head elongately inflated,
connected with oviduct by connective tissue, internally
smooth, with delicate wall. Free oviduct comprising about half
of anterior part of oviduct, more or less straight. Spermoviduct
longer than anterior part of oviduct. Talon embedded in
albumen gland close to anterior end of albumen gland.

Radular morphology (Fig. 59E-F). Rectangular. Only a
partial fragment was studied with 32 rows per mm. Central
teeth with sharply pointed, triangular mesocones, shorter

than base of teeth. Central ectocones reduced. Lateral teeth
with sharply pointed, triangular to ovate mesocones. Lateral
ectocones tiny, endocones reduced. Marginals with triangular
mesocones. Marginal ectocones smaller and narrower than
mesocones, endocones smaller than ectocones.

Comparative remarks. Shell smaller in size, not as elevated
in shape as in congeners from the same island. Large pustules
and very large penial verge are also diagnostic. A lateral
pocket off the penial chamber as mentioned by Solem
(1991) was not observed. This species was referred to as
“Amplirhagada sp. 32” by Solem (1991).

Kohler: New Amplirhagada species from the Kimberley

257

Figure 60. SEM photographs of the shell of Amplirhagada sp. AMS C463710: (A) apical whorl viewed from above (scale 1 mm); (B)
sculpture on second and third whorl viewed from above (scale 1 mm); (C) close-up showing axial sculpture and keel on periphery of
last whorl (scale 200 pm).

Amplirhagada sp.

Material studied. WAM S36953 (PI. 2.3), AMS C463710 (shells, central
section of Boongaree Island, 15°04'00"S 125°11'11"E), WAM S36740 (2
shells, 15°04’36"S 125°11'18"E).

Shell (Fig. 60A-C, PI. 2.3). Semi-globose with medium to
high spire; solid. Whorls evenly rounded in cross-section.
Umbilicus completely concealed by columellar reflection.
Background colour brownish horn (shells worn); with well
defined, chestnut brown, moderately broad sub-sutural and
mid-whorl bands, most conspicuous on last whorl(s); ventral
colour horn; outer lip colour lighter than shell. Protoconch c.
2.2 mm in diameter, comprising 1.7 whorls, almost smooth.
Teleoconch with fine axial lirae, rounded in cross-section,
regularly spaced, spaces narrower than thickness of ribs,
distributed evenly across shell surface, reduced underneath
suture. Angle of aperture 30°, outer lip rounded, moderately
thick, expanded, slightly reflected, basal node of lip weak,
palatal node absent. Parietal wall of inner lip inconspicuous.

Comparative remarks. Of this species, only five dry shells
from Boongaree island are available. Although these shells
differ clearly from those of all other species on this island,
the limited data available are considered insufficient to justify
formal description of this species. Amplirhagada regia and
A. boongareensis have much larger and more dome-shaped
shells. Amplirhagada gibsoni differs by having a flatter shell.
Adult shells of the present species are generally higher than
14.7 mm, those of A. gibsoni are lower.

Amplirhagada yorkensis n.sp.

Type locality (Fig. 1). Western Australia, northwestern
Kimberley, Bonaparte Archipelago, York Sound, Coronation
Island, southern section, 15 o 0F52"S 124°56’56"E; KIS-
3-085. Vine thicket on upper slopes of a volcanic hill behind
mangroves, under rocks (leg. V. Kessner, 30 May 2008).

Type material. Holotype WAM S34619 (PI. 2.4). Paratypes AMS C4637 11
(24 preserved specimens), WAM S36979 (42 preserved specimens).

Additional, non-type material. WAM S41477, FMNH 219151 (12
preserved specimens. Port Nelson, 5 km N of Mt. Knight, c. 3 km E of

Careening Bay; 15°06’41"S 125°01'53"E; KC-040), WAM S37376-8,
WAM S36980-6, WAM S37394-404, AMS C463712-4 (Coronation
Island, southern section).

Etymology. In reference to York Sound, where this species
occurs on islands and likely adjacent mainland.

Sealing strategy. Free sealer.

Shell (Fig. 62A-B, PI. 2.4-5). Broadly conical to almost flat
with low spire; thin (translucent). Periphery evenly rounded
to slightly angulate; upper and basal sectors of whorls
rounded. Umbilicus open, narrowly winding, 30-90 percent
concealed by columellar reflection. Background and ventral
colour brownish horn; sub-sutural band absent; mid-whorl
band absent or diffuse, light yellowish brown, thin, generally
not very pronounced, only visible on last whorl(s); outer
lip colour same as shell; inner lip translucent, whitish.
Protoconch c. 2.7 mm in diameter, comprising 2 whorls,
with fine, indistinct axial lirae. Teleoconch smooth except of
axial growth lines. Angle of aperture 30°; outer lip rounded,
sharp, expanded to largely expanded, reflected; basal and
palatal node absent. Parietal wall of inner lip inconspicuous.

Figure 61. Genitalia of Amplirhagada yorkensis n.sp., paratype
AMS C463711 (30 May, scale 5 mm). Compare with Fig. 3 for
labelling of structures.

258

Records of the Australian Museum (2010) Vol. 62

Figure 62. SEM photographs of shell, jaw and radula of Amplirhagada yorkensis n.sp., paratype AMS C463711: (A) apical whorl viewed
from above (scale 200 pm); ( B ) sculpture on second to forth whorl viewed from above (scale 200 pm); (C) jaw (scale 100 pm); (D)
close-up of central and inner lateral teeth (scale 10 pm); (E) outer lateral and inner marginal teeth (scale 10 pm); (F) close-up of middle
marginal teeth (scale 10 pm).

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl; mantle pigmentation spotted, black.
Kidney extending about half of pallial cavity.

Genital morphology (Figs. 61, 63). Penis curved to sharply
bent, more or less of same length as anterior part of oviduct.
Vas deferens coils once before entering penis. Penial retractor
muscle very short, stubby. Penial verge very short, slender to
spatulate, with pointed tip. Penial wall pustules of average
size, arranged in rows over entire length of inner penial
wall. Main stimulatory pilaster well-differentiated, large,

cone-shaped, sculptured by horizontal ridges that support
little hooks, comprising about 3 A of length of penial chamber.
Vas deferens entering penial sheath in upper third. Vagina
moderately wide, posteriorly inflated; inner vaginal wall
with smooth longitudinal pilasters. Spermathecal duct wide,
internally with smooth longitudinal pilasters. Spermathecal
head pyriform, connected with oviduct by connective tissue,
internally smooth with thing wall. Free oviduct comprising
about half of anterior part of oviduct, more or less straight.
Spermoviduct longer than anterior part of oviduct. Talon
embedded in albumen gland at junction with spermoviduct.

Kohler: New Amplirhagada species from the Kimberley

259

Figure 63. Interior of penial chamber of Amplirhagada yorkensis
n.sp., paratype AMS C463711 (30 May, scale 3 mm). Compare
with Fig. 4 for labelling of structures.

Radular morphology (Fig. 62D-F). Rectangular. Tooth
formula C+13-14+3+18. In average with 122±1.5 rows
of teeth, 32.4±2.0 rows per mm (n = 2). Central teeth with
sharply pointed, triangular mesocones, shorter than base
of tooth; ectocones vestigial. Lateral teeth with bluntly
pointed, triangular to ovate mesocones, length equal to base
of tooth; ectocones and endocones vestigial. Marginal teeth
with triangular to elongate mesocones; ectocones shorter
and narrower than mesocones; endocones reduced in size.

Comparative remarks. The shell is low spired with a
rounded periphery and light colouration; penis with a
comparatively large main pilaster. This species was referred
to as “Amplirhagada sp. 34” by Solem (1991).

Amplirhagada buffonensis n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Buffon Island, 14°54'40"S
124°44T3"S; KC-086 (leg. V. Kessner & A. Longbottom,
23 July 1988).

Type material. Holotype WAM S34620 (PI. 2.6). Paratypes AMS C463758
(2 preserved specimens, as holotype), FMNH 219337 (8 preserved
specimens, same as holotype), WAM S41482 (5 preserved specimens,
as holotype), FMNH 219365 (8 preserved specimens, unnamed islet W
of Buffon Island, 14°54’29"S 124°43'28"E), AMS C463759 (2 preserved
specimens, as FMNH 219365), WAM S41483 (5 preserved specimens,
same as FMNH 219365).

Additional, non-type material. FMNH 219374, WAM S41484 (preserved
specimens, unnamed island S of Buffon Island, 14°57'07"S 124°44'50"E).

Etymology. In reference to Buffon Island, where this species
occurs.

Shell (Fig. 65A-C, PL 2.6). Broadly conical to almost
flat with low spire; thin (translucent). Periphery evenly
rounded to slightly angulate; upper and basal sectors of
whorls rounded. Umbilicus open, forming narrowly winding
opening, 10-20 percent concealed by columellar reflection.
Background and ventral colour greyish horn to slightly beige,
whitish cracks and growth lines visible; sub-sutural band
absent; mid-whorl band absent or diffuse, yellowish brown,
thin, only visible on last whorl; outer and inner lip whitish.
Protoconch c. 2.7 mm in diameter, comprising 1.7 whorls,
smooth. Teleoconch with faint axial growth lines only. Angle
of aperture 45°, outer lip rounded, sharp to moderately thick,
expanded, slightly reflected, basal node of lip absent to weak,
palatal node absent. Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl; mantle pigmentation comprises densely
packed black spots or patches. Kidney extending about half
of pallial cavity.

Genital morphology (Figs. 64, 66). Penis straight to
slightly curved, more or less of same length as anterior part
of oviduct. Vas deferens forms simple loop or coils before

Figure 64. Genitalia of Amplirhagada buffonensis n.sp., paratype
AMS C463758 (23 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

260

Records of the Australian Museum (2010) Vol. 62

Figure 65. SEM photographs of shell, jaw and radula of Amplirhagada buffonensis n.sp., paratype AMS C463759: (A) apical whorl
viewed from above (scale 200 pm); ( B ) sculpture on first three whorls viewed from above (scale 200 pm); (C) close-up of periphery of
last whorl (scale 200 pm); ( D ) jaw (scale 100 pm); (E) central and inner lateral teeth (scale 10 pm); (F) outer lateral and inner marginal
teeth (scale 10 pm).

entering penis. Penial retractor muscle shorter than penis.
Penial verge very short (barely visible), slender to spatulate,
with pointed to rounded tip. Penial wall pustules very small,
densely arranged in rows on apical to median portion of
penial chamber, becoming oblique towards base of penial
chamber. Main stimulatory pilaster well-differentiated, large,
cone-shaped, sculptured by horizontal ridges that support
little hooks, comprising almost entire length of penial
chamber. Vas deferens entering penial sheath in upper third
of penis. Vagina moderately wide, posteriorly inflated; inner
vaginal wall with smooth longitudinal pilasters. Spermatheca
of long, clearly extending over base of spermoviduct.

Spermathecal duct rather wide, internally with smooth
longitudinal pilasters. Spermathecal head elongately inflated
with extended tubular or pyriform tip, connected with oviduct
by connective tissue, internally smooth, with delicate wall.
Free oviduct comprising less than half of anterior part of
oviduct, more or less straight. Spermoviduct longer than
anterior part of oviduct. Talon embedded in albumen gland
close to anterior end of albumen gland.

Radular morphology (Fig. 65E-F). Rectangular. Tooth
formula C+14-15+3-4+18-22. In average 153±17.5 rows
of teeth, 27.4±0.1 rows per mm (n = 2). Central teeth with

Kohler: New Amplirhagada species from the Kimberley 261

bluntly pointed, triangular mesocones, shorter than base of
tooth; ectocones vestigial. Lateral teeth with sharply pointed,
triangular to ovate mesocones, length equal to base of tooth;
ectocones well developed, endocones vestigial. Marginal
teeth with triangular mesocones; ectocones shorter and
narrower than mesocones; endocones reduced in size.

Figure 67. Genitalia of Amplirhagada uwinsensis n.sp., paratype
AMS C463715 (7 August, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Comparative remarks. It has a low spired shell with
reduced banding pattern. Differs from its sister species in
the phylogeny, A. yorkensis, by angulate periphery, slightly
larger shell size (Table 1), and smaller verge. Material of
this species has been labelled as “Amplirhagada sp. 67”
by Solem.

Amplirhagada uwinsensis n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Hanover Bay, Uwins Island,
15°15'32"S 124°46'08"E; KIS-3-106. Vine thicket and large
fig trees on upper slopes, piles of sandstone rocks, under
large slabs, (leg. V. Kessner, 09 June 2006).

Type material. Holotype WAM S34621 (PL 2.7). Paratypes AMS
C463715 (15 preserved specimens, as holotype), WAM S36991 (35
preserved specimens, as holotype), AMS C463716 (15 shells, 15°15'25"S
124°48'04"E), WAM S37441 (30 shells, 15°15’25"S 124°48'04"E).

Additional, non-type material. WAM S36987-90, WAM S37440, WAM
S37442^1, AMS C463717 (Uwins Island).

Etymology. In reference to Uwins Island, where this species
occurs.

Figure 68. Interior of penial chamber of Amplirhagada uwinsensis
n.sp., paratype AMS C463715 (7 August, scale 5 mm). Compare
with Fig. 4 for labelling of structures.

262

Records of the Australian Museum (2010) Vol. 62

Figure 69. SEM photographs of Amplirhagada uwinsensis n.sp. (A-C) Shell, paratype AMS C463716: (A) apical whorl viewed from
above (scale 200 pm); ( B ) sculpture penultimate whorl viewed rectangularly from above (scale 100 pm); (C) close-up of periphery of last
whorl (scale 200 pm). ( D ) Jaw, paratype AMS C463715 (scale 100 pm). (E-F) Radula, paratype AMS C463715: (E) close-up of central
and inner lateral teeth (scale 10 pm); ( F) close-up of inner to middle marginal teeth (scale 10 pm).

Shell (Fig. 69A-C, PI. 2.7-9). Semi-globose with medium
high spire; solid. Periphery angulate; upper and basal sectors
of whorls rounded. Umbilicus completely concealed by
columellar reflection. Background colour light reddish
brown; sub-sutural band diffuse to well defined, light brown
to chestnut brown, varies greatly in thickness and contrast;
mid-whorl band diffuse, brownish to chestnut brown, thin,
varies greatly in contrast and thickness, visible on last
whorl(s) only; bands may blend into each other across whorl
surface; ventral colour yellowish-greenish brown to ochre;
outer lip colour blending into chestnut brown; inner lip
blending into dark ochre to chestnut brown. Protoconch c. 2.7
mm in diameter, comprising 1.7 whorls, with fine, indistinct
axial lirae. Teleoconch with marked growth lines, becoming
oblique towards base of shell. Angle of aperture 45°; outer lip
rounded, slightly expanded, not reflected, basal node of lip
weak, palatal node absent. Parietal wall of inner lip absent.

Pallial morphology. Pallial cavity short, extending Vi to
3 A whorls; mantle pigmentation consists of indistinctive,
sparsely distributed brown spots. Kidney extending about
half or more than half of pallial cavity.

Genital morphology (Figs. 67-68). Penis rather straight,
more or less of same length as anterior part of oviduct. Vas
deferens coils once before entering penis. Penial retractor

muscle very short to stubby. Penial verge very short (barely
visible), slender to spatulate, with pointed tip. Penial wall
pustules comparatively large, elongated, arranged in rows
on median and basal portion of penial chamber, forming
densely packed, narrow longitudinal pilasters at apical
end of penial chamber. Main stimulatory pilaster well-
differentiated, large, cone-shaped, covered with flattened
pustules, comprising anterior to median portion of penial
chamber. Two additional pilasters are formed that are covered
by flattened pustules or smooth narrow ridges. Vas deferens
entering penial sheath in upper third. Vagina moderately
wide, tubular to posteriorly inflated; inner vaginal wall
with weakly developed longitudinal pilasters. Spermathecal
duct wide, internally with smooth longitudinal pilasters.
Spermathecal head globular, connected with oviduct by
connective tissue, internally smooth, with delicate wall. Free
oviduct comprising less than half to about half of anterior
part of oviduct, more or less straight. Spermoviduct of same
length as anterior part of oviduct or longer. Talon embedded
in albumen gland close to anterior end of albumen gland.

Radular morphology (Fig. 69E-F). Rectangular. Tooth
formula C+14-16+4+22. In average 149±16.0 rows of
teeth, 29.5±1.0 rows per mm (n = 2). Central teeth with
sharply pointed, triangular mesocones, shorter than base
of teeth; ectocones vestigial. Lateral teeth with sharply

Kohler: New Amplirhagada species from the Kimberley

263

Figure 70. SEM photographs of Amplirhagada sphaeroidea n.sp. (A-B) Shell, paratype AMS C463719: (A) apical whorl viewed from
above (scale 200 pm); ( B ) close-up of periphery of last whorl (scale 1 mm); (C) jaw, paratype WAM S37027 (scale 100 pm). ( D-E )
Radula, paratype WAM S37027: ( D ) close-up of inner lateral teeth (scale 20 pm); (E) Close-up of outer lateral and inner marginal teeth
(scale 20 pm).

pointed, triangular mesocones, length equal to base of teeth;
ectocones and endocones vestigial. Marginal teeth with
elongate mesocones; ectocones smaller and narrower than
mesocones; endocones shorter than ectocones.

Comparative remarks. One of the few species with large,
dome-shaped shells. Differs from A. tricenaria and A. regia
by conspicuously dark brown outer lip colour and by huge,
cone-shaped main pilaster, presence of additional pilasters
and characteristically elongated pustules of inner penial wall.

Amplirhagada sphaeroidea n.sp.

Type locality (Fig. 1). Western Australia, Kimberley, Saint
George Basin, St. Andrews Island, 15 0 21'24"S 124°59’46"E;
KIS-2-40. Volcanic boulder scree, vine thicket, under rocks
(leg. R. Teale, 25 May 2008).

Type material. Holotype WAM S34622 (PL 2.10). Paratypes AMS
C463718 (2 preserved specimens, same as holotype), WAM S37029 (7
preserved specimens, same as holotype), AMS C463719 (shell, 15°21'34"S
125°00’07"E), WAM S37027 (3 preserved specimens, 15°21'34"S
125°00'07"E), WAM S37411 (1 dry shell, 15°21'34"S 125°00'07"E).

Figure 71. Genitalia of Amplirhagada sphaeroidea n.sp., paratype
WAM S37027 (24 May, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Etymology. From sphaeroeides (Greek = globular,
spherical), referring to globular shape of shell.

Additional, non-type material. WAM S37028, WAM S37030, WAM

S37410, wam S37412 (St. Andrews island). Sealing strategy. Free sealer.

264

Records of the Australian Museum (2010) Vol. 62

Shell (Fig. 70A-B, PL 2.10). Globose to semi-globose,
with medium high spire; solid to thick. Periphery angulate;
upper sector of whorls rather flattened, basal sector rounded.
Umbilicus forming a chink, 90-100 percent concealed by
columellar reflection. Background colour blends from horn
at base to lightly brownish at top of shell; sub-sutural band
absent or diffuse, indistinct, brownish; mid-whorl band
absent or diffuse, brownish, thin, indistinct, visible on last
whorl(s) only; ventral colour cream; outer lip colour differs
from shell, dark brownish-purple; inner lip blends from dark
pink outside to horn deeper inside. Protoconch c. 2.4 mm
in diameter, comprising 1.7 whorls, smooth. Teleoconch
smooth, except of faint growth lines. Angle of aperture 45°;
outer lip rounded, thick, slightly expanded to expanded,
slightly reflected, basal node of lip weak, palatal node absent.
Parietal wall of inner lip absent or inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending± 3 /4 whorl; mantle pigmentation mottled, diffuse
dark grey. Kidney extending about half of pallial cavity.

Figure 72. Interior of penial chamber of Amplirhagada sphaeroidea
n.sp., paratype WAM S37027 (24 May, scale 3 mm). Compare with
Fig. 4 for labelling of structures.

Genital morphology (Figs. 71-72). Penis straight, more or
less of same length as anterior part of oviduct. Vas deferens
forms simple loop before entering penis. Penial retractor
muscle very short, stubby. Penial sheath evenly thick. Penial
verge short (<Vs penial chamber), slender to spatulate, with
pointed tip. Penial wall pustules fused to regular, smooth,
diagonal and delicate lamellae on apical portion, becoming
more and more oblique towards base of penial chamber.
Main stimulatory pilaster well-differentiated; elongated,
large, cone-shaped, corrugated, comprising mainly the
apical portion of penial chamber and gradually decreasing in
size towards base. Three additional, smooth, narrow, rather
indistinct pilasters are formed at base of penial chamber.
Vas deferens entering penial sheath apically. Vagina rather
short, posteriorly inflated; inner vaginal wall with smooth
longitudinal pilasters. Spermathecal duct moderately wide,
internally with smooth longitudinal pilasters. Spermathecal
head globular, connected with oviduct by connective tissue,
internally entirely smooth, wall delicate. Free oviduct
comprising about half of anterior part of oviduct, or more
than half of anterior part of oviduct; more or less straight.
Spermoviduct much longer than anterior part of oviduct.
Talon embedded in albumen gland anteriorly.

Radular morphology (Fig. 70D-E). Rectangular. Tooth
formula C+15+4+18. With 118 rows of teeth, 26.0 rows
per mm (n = 1). Central teeth with bluntly pointed, broadly
elongate mesocone, length equal to base of teeth; ectocones
vestigial. Lateral teeth with bluntly pointed, broadly elongate
mesocone, length equal to base of teeth; ectocones small,
endocones vestigial. Marginal teeth with broadly elongate
mesocones; ectocones smaller and narrower than mesocones;
endocones greatly reduced.

Comparative remarks. One of the few species with large,
dome-shaped shells. Differs from A. tricenaria and A. regia
by dark outer lip and from A. uwinsensis purplish colour
of outer lip. Covering of inner penial wall with lamellae,
elongate-conical, corrugated main pilaster, and relatively
large verge are diagnostic.

Amplirhagada basilica n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Augustus Island, southern section,
Brecknock Harbour, 15°23’53"S 124°036’03 , 'E; KIS 2-51.
Weathered south facing sandstone scree in deep gorge, under
rocks (leg. R. Teale, 1 June 2008).

Type material. Holotype WAM S34623 (PI. 2.11). Paratypes AMS C463720
(4 preserved specimens, same as holotype), WAM S37004 (11 preserved
specimens, same as holotype), AMS C463721 (3 shells, 15°22'54"S
124°35’24"E), WAM S37414 (4 shells, 15°22'54"S 124°35'24"E).

Additional, non-type material. WAM S37000-3, WAM S37413 (Augustus
Island).

Etymology. From basilica (Latin = royal, splendid,
magnificent), referring to its large shell. Amplirhagada regia
is among the largest species of the genus.

Sealing strategy. Free sealer.

Kohler: New Amplirhagada species from the Kimberley

265

Figure 73. SEM photographs of Amplirhagada basilica n.sp. (A-C) Shell, paratype AMS C463721: (A) apical whorl viewed from above
(scale 200 pm); ( B ) close-up of periphery of penultimate whorl (scale 1 mm); (C) close-up of periphery of last whorl (scale 1 mm). (D)
Jaw, paratype AMS C463720 (scale 100 pm). (E-F) Radula, paratype AMS C463720: (E) central and inner lateral teeth (scale 10 pm);
(F) outer lateral and inner marginal teeth (scale 10 pm).

Shell (Fig. 73A-C, PI. 2.11). Semi-globose, with medium
high spire; solid to thick. Periphery slightly angulate; upper
and basal sectors of whorls rounded. Umbilicus open,
forming a chink to narrowly winding opening, 80-95 percent
concealed by columellar reflection. Background colour
blends from cream into brownish (upper sector of lower
whorls darker); sub-sutural band diffuse, light brownish;
mid-whorl band diffuse, thin, brownish, most pronounced,
visible on last whorls only, blurring towards tip of shell;
ventral colour horn; outer lip colour differs from shell,
brownish purple; inner lip colour pinkish. Protoconch 2.4
mm in diameter, comprising 1.7 whorls, smooth. Teleoconch
smooth except of indistinct growth lines. Angle of aperture
45°; outer lip rounded, moderate to thick, expanded, slightly
reflected, basal node of lip weak, palatal node absent. Parietal
wall of inner lip absent.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl; mantle pigmentation mottled, dark grey.
Kidney extending about half of pallial cavity.

Genital morphology (Figs. 74-75). Penis straight, shorter
than anterior part of oviduct, very thin. Vas deferens forms
simple loop before entering penis. Penial retractor muscle
of same length as penis. Penial sheath evenly thick. Penial
verge comprising Vs- l A of penial chamber, slender to
spatulate, with rounded tip. Penial wall pustules absent. Main

stimulatory pilaster absent. Two smooth, narrow longitudinal
pilasters comprise entire length of penial chamber; two
additional smooth, narrow pilasters develop at base of
penial chamber. Vas deferens rather thick, entering penial
sheath apically. Vagina rather short, posteriorly inflated;
inner vaginal wall densely covered by tiny pustules and
smooth longitudinal pilasters. Spermatheca long, extending
over base of spermoviduct. Spermathecal duct moderately

Figure 74. Genitalia of Amplirhagada basilica n.sp., paratype
AMS C463720 (1 June, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

266

Records of the Australian Museum (2010) Vol. 62

wide, internally with pustules. Spermathecal head globular,
connected with oviduct by connective tissue, internally
smooth, with thin wall. Free oviduct comprising half of
anterior part of oviduct or more; more or less straight.
Spermoviduct much longer than anterior part of oviduct.
Talon embedded in albumen gland anteriorly.

Radular morphology (Fig. 73E-F). Rectangular. Tooth
formula C+14+4+22. In average 119±19.0 rows of teeth,
28.0±0.4 rows per mm (n = 2). Central teeth with bluntly
pointedovate mesocone, shorter than base of teeth;
ectocones vestigial. Lateral teeth with bluntly pointed,
ovate mesocones, shorter than base of teeth; ectocones and
endocones vestigial. Marginal teeth with broadly rounded
mesocones; ectocones pointed, much smaller and narrower
than mesocones; endocones vestigial.

Comparative remarks. One of the few species with large,
dome-shaped shells. Differs from A. tricenaria and A. regia
by dark outer lip. Purplish colour of outer lip si mil ar to A.
sphaeroidea. Absence of penial wall pustules is characteristic
for this species.

Amplirhagada camdenensis n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Camden Sound, Augustus Island,
northwestern section, 15°20T6"S 124 0 3F16"E; KIS 1-49.
Vine thicket amongst boulders at base of sandstone cliffs,
under rocks (leg. M. Shea, 25 May 2008).

Type material. Holotype WAM S34624 (PL 2.12). AMS C463722 (3
preserved specimens, as holotype), WAM S37006 (5 preserved specimens,
as holotype), WAM S37005 (3 preserved specimens, 15°20'59"S
124°31'30"E).

Additional, non-type material. WAM S37007-8, WAM S37445-50, WAM
S41481, FMNH 219063 (Augustus Island).

Etymology. In reference to Camden Sound, in which
Augustus Island is situated.

Sealing strategy. Free sealer.

Shell (Fig. 77A-B, PL 2.12). Globose, with medium to high
spire; solid. Periphery evenly rounded to slightly angulate;
upper sector of whorl flattened to slightly shouldered, basal
sector rounded. Umbilicus forming a chink to narrowly
winding opening, 40-100 percent concealed by columellar
reflection. Background colour yellowish brown to horn;
sub-sutural band diffuse, dark to chestnut brown, becomes
more indistinct or blurs towards tip of whorl, may fuse
with mid-whorl band to cover shell in light brown colour;

Kohler: New Amplirhagada species from the Kimberley

267

Figure 77. SEM photographs of shell and radula of Amplirhagada camdenensis n.sp., paratype AMS C463722: (A) apical whorl viewed
from above (scale 200 pm); ( B ) sculpture on first three whorls viewed from above (scale 1 mm); (C) close-up showing central and inner
lateral teeth (scale 10 pm); (D) close-up showing inner and middle marginal teeth (scale 10 pm).

mid-whorl band diffuse, dark to chestnut brown, thin to
moderately broad, visible on last whorl(s) only; ventral
colour whitish cream to yellowish brown; outer and inner lip
colour differs from shell, pinkish brown. Protoconch c. 3.5
mm in diameter, comprising 2 whorls, smooth. Teleoconch
smooth except of faint axial growth lines. Angle of aperture
30°; outer lip rounded, moderately thick, expanded, not or
slightly reflected, basal and palatal node absent. Parietal wall
of inner lip absent.

Figure 78. Genitalia of Amplirhagada camdenensis n.sp., paratype
AMS C463722 (25 May, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Pallial morphology. Pigmentation on mantle mottled, dark
grey. Kidney extending about half of pallial cavity.

Genital morphology (Figs. 76, 78). Penis straight, shorter
than anterior part of oviduct, very thin. Vas deferens forms
simple loop before entering penis. Penial retractor muscle
as long as penis. Penial verge rather short (Vs-Vt of length
of penial chamber), broad, with pointed to rounded tip.
Penial wall pustules absent. Main stimulatory pilaster not
well differentiated, narrow, smooth, on apical to median
portion of penial chamber. One to three additional, smooth,
narrow pilasters on apical portion. Vas deferens entering
penial sheath in upper third. Vagina rather short, medially
to posteriorly inflated; inner vaginal wall predominantly
smooth, with few weakly developed longitudinal pilasters.
Spermathecal duct moderately wide, internally smooth.
Spermathecal head globular to elongately inflated, connected
with oviduct by connective tissue, internally smooth, with
thin wall. Free oviduct comprising about half of anterior part
of oviduct; more or less straight. Spermoviduct much longer
than anterior part of oviduct. Talon embedded in albumen
gland at junction with spermoviduct.

Radular morphology (Fig. 77C-D). Rectangular. Tooth
formula C+16+4+?. With 122 rows of teeth, 27.4 rows per
mm (n = 1). Central teeth with sharply pointed, triangular
mesocones, shorter than base of tooth; ectocones vestigial.
Lateral teeth with sharply pointed, triangular to ovate

268

Records of the Australian Museum (2010) Vol. 62

Figure 79. SEM photographs of Amplirhagada kimberleyana n.sp. ( A-D ) Shell, paratype FMNH 219055: (A) apical whorl viewed from
above (scale 200 pm); ( B ) sculpture on first three whorls viewed from above (scale 1 mm); (C) close-up showing sculpture on penultimate
whorl (scale 200 pm); ( D ) close-up showing surface of last whorl (scale 200 pm). (E) Jaw, paratype AMS C463760 (scale 100 pm). (F-G)
Radula, paratype AMS C463760: (F) central and lateral teeth (scale 20 pm); (G) outer lateral and inner marginal teeth (scale 20 pm).

mesocones, length equal to base of tooth; ectocones and
endocones vestigial. Marginal teeth with triangular to broadly
flattened mesocones; ectocones shorter and narrower than
mesocones; endocones vestigial.

Comparative remarks. Material of this species has been
labelled as “Amplirhagada sp. 65” by Solem. It has the
smallest of all dome-shaped shells with a significantly higher
H/D ratio as most other species (Table 1). Absence of penial
wall pustules and pattern of smooth, longitudinal pilasters
are diagnostic.

Amplirhagada kimberleyana n.sp.

Type locality (Fig. 1). Western Australia, Kimberley, Bona¬
parte Archipelago, Camden Sound, northern Slate Islands,
2.4 km NW of Wilson Point, 15°32T0"S 124°23’56"E;
KC-005 (leg. V. Kessner & A. Longbottom, 09 July 1988).

Type material. Holotype WAM S34625 (PL 2.13). Paratypes AMS
C463760 (5 preserved specimens), FMNH 219055 (21 preserved
specimens), WAM S41480 (15 preserved specimens).

Kohler: New Amplirhagada species from the Kimberley 269

Figure 80. Genitalia of Amplirhagada kimberleyana n.sp., paratype
AMS C463760 (9 July, scale 10 mm). Compare with Fig. 3 for
labelling of structures.

Etymology. In reference to the Kimberley region.

Shell (Fig. 79A-D, PL 2.13). Semi-globose to broadly
conical, with medium high spire; thin to solid (translucent).
Periphery evenly rounded to slightly angulate; upper sector of
whorl and basal sector rounded. Umbilicus forming a chink
to narrowly winding opening, 60-100 percent concealed by
columellar reflection. Background colour beige-brown; sub-
sutural band absent or diffuse, brown; mid-whorl band absent
or diffuse, brown, thin, visible on last whorl only; ventral
colour lighter, horn or whitish; outer lip colour differs from
shell, light brown to whitish; inner lip whitish. Protoconch c.
3 mm in diameter, comprising 2 whorls, smooth. Teleoconch
smooth except of axial growth lin es. Angle of aperture 45°;
outer lip rounded, moderately thick, slightly expanded, not
or slightly reflected, basal and palatal node absent. Parietal
wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl. Pigmentation on mantle mottled, black.
Kidney extending about half of pallial cavity.

Genital morphology (Figs. 80-81). Penis straight, more or
less of same length as anterior part of oviduct or longer. Vas
deferens coils once before entering penis. Penial retractor
muscle shorter than penis. Penial verge moderately long
(Vs-V4 penial chamber), slender to spatulate, with pointed
tip. Penial wall pustules large, arranged in rows on apical
to median portion of the penial chamber. Main stimulatory
pilaster undifferentiated, formed by few rows of slightly
enlarged, hooked pustules. Base of penial wall smooth,
supporting two smooth and narrow pilasters. Vas deferens
entering penial sheath in upper third. Vagina moderately
long, tubular to posteriorly slightly inflated; inner vaginal
wall with smooth longitudinal pilasters. Spermathecal duct
wide, internally with corrugated longitudinal pilasters.
Spermathecal head globular, connected with oviduct by
connective tissue, internally smooth, with thin wall. Free
oviduct comprising about half of anterior part of oviduct,
more or less straight. Spermoviduct of same length as anterior
part of oviduct. Talon embedded in albumen gland at junction
with spermoviduct.

Radular morphology (Fig. 79F-G). Rectangular. Tooth
formula C+16+3+22. With 147 rows of teeth, 31.9 rows per
mm (n = 1). Central teeth with sharply pointed, triangular
mesocones, shorter than base of tooth; ectocones vestigial.
Lateral teeth with sharply pointed, triangular mesocones,
length equal to base of tooth; ectocones small, endocones
vestigial. Marginal teeth with elongate to triangular
mesocones; ectocones shorter and narrower than mesocones,
divided into two denticles; endocones greatly reduced.

Comparative remarks. Shell broadly conical to dome¬
shaped but smaller than species with similar shell shape.
Inner penial wall with strong developed pustulation and
undifferentiated main pilaster is diagnostic. Material of the
present species has been labelled as “Amplirhagada sp. 64”
by Solem.

270

Records of the Australian Museum (2010) Vol. 62

Figure 82. SEM photographs of Amplirhagada gemina n.sp. (A-B) Shell, holotype WAM S34626: (A) apical whorl viewed from above
(scale 200 pm); ( B ) sculpture on second whorl viewed from above (scale 100 pm); (C) close-up showing sculpture on penultimate whorl
(scale 200 pm). ( D) Jaw (scale 100 pm), paratype FMNH 219049. (E-F) Radula, paratype FMNH 219049: (E) central and lateral teeth
(scale 20 pm); ( F) middle marginal teeth (scale 10 pm).

Amplirhagada gemina n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Camden Sound, mainland 4.5 km
N of Prior Point, 1.5 km SE of Hall Point; KC-005 (leg. V.
Kessner & A. Longbottom, 09 July 1988).

Type material. Holotype WAM S34626 (PI. 2.14). Paratypes AMS
C463761 (preserved specimen), FMNH 219049 (5 preserved specimens),
WAM S41479 (3 preserved specimens).

Etymology. From gemina (Latin = twin-born), in reference
to its close overall similarity with its sibling species A.
kimberleyana.

Shell (Fig. 82A-C, PI. 2.14). Semi-globose to broadly
conical, with medium spire, thin to solid (translucent).
Periphery evenly rounded to slightly angulate; upper and
basal sectors of whorls rounded. Umbilicus forming a chink
to narrowly winding opening, 60-100 percent concealed
by columellar reflection. Background colour beige brown
sub-sutural band absent or diffuse, brown; mid-whorl band
absent or diffuse, brown, thin, visible on last whorl only;
ventral colour lighter, horn or whitish; outer lip colour
differs from shell, light brown to whitish; inner lip whitish.
Angle of aperture 45°, outer lip moderately thick, slightly
expanded, not or slightly reflected, basal and palatal node
absent. Teleoconch with pronounced axial growth lines

Kohler: New Amplirhagada species from the Kimberley

271

Figure 83. Genitalia of Amplirhagada gemina n.sp., holotype WAM
S34626 (9 July, scale 10 mm). Compare with Fig. 3 for labelling
of structures.

which become more oblique towards the base of the shell.
Parietal wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity moderately deep,
extending 3 A whorl; mantle pigmentation mottled, black.
Kidney extending about half of pallial cavity.

Genital morphology (Figs. 83-84). Penis straight, more
or less of same length as anterior part of oviduct. Vas
deferens coils once before entering penis. Penial retractor
muscle shorter than penis. Penial verge short (<Vs penial
chamber), slender to spatulate, with pointed tip. Penial
wall pustules rather large, arranged in rows on entire wall
of penial chamber, becoming more oblique towards base.
Main stimulatory pilaster not well differentiated, covered
by undifferentiated pustules, comprising apical to median
portion of penial chamber. Vas deferens entering penial
sheath in upper third. Vagina moderately long, tubular
to posteriorly inflated. Inner vaginal wall with smooth
longitudinal pilasters. Spermathecal duct wide, inflated,
internally with smooth longitudinal pilasters. Spermathecal
head elongately inflated, connected with oviduct by
connective tissue, internally smooth, with delicate wall. Free
oviduct comprising about half of anterior part of oviduct,
more or less straight. Posterior parts of genital system
unknown.

Radular morphology (Fig. 82E-F). Rectangular. Tooth
formula C+16+4+20. With 150 rows of teeth, 34.2 rows per
mm (n = 1). Central teeth with sharply pointed, triangular
mesocones, shorter than base of tooth; ectocones vestigial.
Lateral teeth with sharply pointed, triangular mesocones,
length equal to base of tooth; ectocones tiny, endocones
vestigial. Marginal teeth with elongate to triangular
mesocones; ectocones shorter and narrower than mesocones;
endocones smaller than ectocones.

Comparative remarks. Smaller than other species with
broadly conical to dome-shaped. Inner penial wall pustules

Figure 84. Interior of penial chamber of Amplirhagada gemina
n.sp., holotype WAM S34626 (9 July, scale 3 mm). Compare with
Fig. 4 for labelling of structures.

larger and denser than in A. kimberleyana. Although
morphologically similar, both species are genetically well-
differentiated. Material of the present species has been
labelled as “Amplirhagada sp. 64” by Solem together with
material of A. kimberleyana, which has a si mil ar shell.

Amplirhagada dubitabile n.sp.

Type locality (Fig. 1). Western Australia, Kimberley,
Bonaparte Archipelago, Doubtful Bay, Steep Island, 2.3 km
NE of Raft Point, 16°03'36"S 124°22’03"E; KC-002 (leg.V.
Kessner & A. Longbottom, 08 July 1988).

Type material. Holotype WAM S34627 (PL 2.15). Paratypes AMS
C463762 (3 preserved specimens), FMNH 219047 (8 preserved
specimens), WAM S41478 (5 preserved specimens).

Etymology. From dubitabile (Latin = doubtful), in reference
to Doubtful Bay, where this species occurs on Steep Island.

Shell (Fig. 85A-C, PI. 2.15). Semi-globose to broadly
conical, with medium high spire; solid. Periphery slightly
angulate; upper and basal sectors of whorls rounded.
Umbilicus open, forming narrowly winding opening, 30-70

272

Records of the Australian Museum (2010) Vol. 62

Figure 85. SEM photographs of shell, jaw and radula of Amplirhagada dubitabile n.sp., paratype FMNH 219047: (A) apical whorl viewed
from above (scale 200 pm); (B) sculpture on penultimate whorl viewed rectangular from above (scale 200 pm); (C) close-up showing
growth lines on last whorl (scale 200 pm); ( D ) jaw (scale 100 pm); (E) central and inner lateral teeth (scale 20 pm); (F) outer lateral and
inner marginal teeth (scale 20 pm).

percent concealed by columellar reflection. Background
colour brownish beige, may display irregular pattern of
hornish growth lines; sub-sutural band absent; mid-whorl
band absent or diffuse, yellowish brown, thin, visible on
last whorl only; ventral colour horn; outer lip colour differs
from shell, horn; inner lip whitish. Protoconch c. 2.8 mm
in diameter, comprising 2 whorls, smooth. Teleoconch
with pronounced axial growth lines. Angle of aperture 45°;
outer lip rounded, moderately thick, slightly expanded, not
or slightly reflected, basal and palatal node absent. Parietal
wall of inner lip inconspicuous.

Pallial morphology. Pallial cavity short, extending half a
whorl; mantle pigmentation mottled, greyish black. Kidney
extending about half of pallial cavity.

Genital morphology (Figs. 86-87). Penis straight, more
or less of same length as anterior part of oviduct. Vas
deferens coils once before entering penis. Penial retractor
muscle shorter than penis. Penial verge short (<Vs penial
chamber), broad, pointed. Penial wall pustules absent; wall
covered by delicate, narrow, longitudinal pilasters. Main
stimulatory pilaster well-differentiated; smooth undulating,
well developed at apical portion, decreasing in size in
median portion, largely reduced at basal portion of penial
chamber. Vas deferens entering penial sheath in upper third.

Vagina rather short, posteriorly inflated; inner vaginal wall
with weakly developed longitudinal pilasters. Spermatheca
slightly extending over base of spermoviduct. Spermathecal
duct wide, internally with smooth longitudinal pilasters.
Spermathecal head globular, connected with oviduct by
connective tissue, internally smooth, with delicate wall. Free
oviduct comprising about half of anterior part of oviduct,
more or less straight. Spermoviduct longer than anterior
part of oviduct. Talon embedded in albumen gland close to
anterior end of albumen gland.

Figure 86. Genitalia of Amplirhagada dubitabile n.sp., paratype
FMNH 219047 (8 July, scale 10 mm). Compare with Fig. 3 for
labelling of stmctures.

Kohler: New Amplirhagada species from the Kimberley

273

Figure 87. Interior of penial chamber of Amplirhagada dubitabile
n.sp., paratype FMNH 219047 (8 July, scale 3 mm). Compare with
Fig. 4 for labelling of structures.

Radular morphology (Fig. 85E-F). Rectangular. Tooth
formula C+16+3+22. With 139 rows of teeth, 32.3 rows
per mm (n = 1). Central teeth with sharply pointed,
triangular mesocones, shorter than base of tooth; ectocones
vestigial. Lateral teeth with sharply pointed, triangular to
ovate mesocones, length equal to base of tooth; ectocones
small; endocones vestigial. Marginal teeth with elongate to
triangular mesocones; ectocones shorter and narrower than
mesocones; endocones smaller than ectocones.

Comparative remarks. Material of this species has been
labelled as “ Amplirhagada sp. 63” by Solem. Shell similar
to A. kimberleyana and A. gemina in shape, size, and
colouration. Inner penial wall structure (lamellae, no pustules
with long, undulating main pilaster) very characteristic.

Amplirhagada napierana Solem, 1981

Amplirhagada napierana Solem, 1981a: 225-232, pi. 14d,
figs. 36b, 48a-b, 50, 51a (Barker Gorge, Napier Range).

Material examined. AMS C150457 (Western Australia, Napier Range, E
side, 1.2 km N of Napier Downs, 17°19.5'S 124°48.0'E) (Fig. 1, PI. 2.16).

Remarks. This species was described from various localities
in the Napier Range, where it occurs in three disjunct areas
in close proximity but not sympatrically with A. percita.
The anatomy was described by Solem (1981a). The penial
morphology of the present sample corresponds with the
original description. This species is represented in the
molecular tree shown below.

Amplirhagada percita (Iredale, 1939)

Tenuigada percita Iredale, 1939: 68, pi. V, fig. 14 (northern
end of Napier Range), Solem, 1981a: 211-225, pi. 14e-f,
figs. 47a-j, 50c, 51b-c.

Tenuigada ignara Iredale, 1939: 68, pi. V, fig. 13 (north end
of Napier Range).

Nomenclatural remarks. Iredale (1939) described the two
species, A. percita and A. ignara, based on series of dry
shells. Subsequently, the two names were synonymized by
Solem (1981a), who revised the genus, on the grounds that
no significant difference in the shells were found, which
would justify treatment as distinct species. However, Solem
(1981a) also stated that he found distinct differences between
allopatric populations of this species. Hence, it remains to
be critically tested whether the taxon as currently delimited
by Solem (1981a) encompasses more than a single species.

Material examined. AMS C144028 (Western Australia, Napier Range, 24
km N of Wagon Pass, 17°04.5"S 124°34’E) (Fig. 1, PI. 2.17).

Remarks. A sample of this species is represented in the
molecular tree shown below.

Amplirhagada varia Solem, 1981

Amplirhagada varia varia Solem, 1981a: 294-300, pi. 13c,
figs. 36e, 65b, 66a, 67a-b, 69a-d (Mitchell Plateau,
AM AX port, Warrender Road).

Material examined. AMS C460963 (track to Crystal Creek, 1.5 km from
Walsh Point, 14.446°S 125.792°E) (Fig. 1, PI. 2.18).

Remarks. Solem (1981a) described two subspecies, A. varia
varia and A. varia depressa, which essentially differ in the
length of the penial verge. Both taxa are found in very close
geographical proximity. They were apparently treated as
subspecies because of their otherwise very close anatomy.
However, as they do not meet the criteria of geographical
races that replace each other, I suggest elevating both to the
rank of distinct species. Consequently, herein I refer to the
species name A. varia for the so-called nominate form. The
penis anatomy of the present sample corresponds with the
figures shown by Solem (1981a). This species is represented
in the molecular tree shown below.

274

Records of the Australian Museum (2010) Vol. 62

Amplirhagada mitchelliana Solem, 1981

Amplirhagada mitchelliana Solem, 1981a: 272-280, pi. 13b,
figs. 37i, 55c-d, 61c-e, 62a-b, 64a (Mitchell Plateau,
Warrender Road).

Material examined. WAM S41462 (lookout on track between Mitchell
Plateau camp and Walsh Pool) (Fig. 1, PI. 2.19), AMS C144039 (Mitchell
Plateau, Mitchell Plateau Road, 43.6 km W of Gibb Road-Kalamburu
turnoff).

Remarks. Found in the same area as A. varia, this species
differs most significantly from the former by a distinct
anatomy of the interior penial wall. The penis anatomy of
the present sample corresponds with that shown by Solem
(1981a). This species is represented in the molecular tree
shown below. The two samples sequenced herein do not form
a monophyletic cluster but are separated by a considerable
uncorrected pair-wise genetic distance of 17% and are shown
as non-monophyletic with respect to A. varia. This indicates
problems with the species recognition and probably the
presence of an unidentified species.

Amplirhagada castra Solem, 1981

Amplirhagada castra Solem, 1981a: 286-293, pi. 13d, figs.
37h, 63a-b, 64b, 67e-f, 68 (Mitchell Plateau, Camp
Creek).

Material examined. AMS C460966 (track to Mitchell Plateau ranger
Station, 14.821 S 125.721 E) (Fig. 1, PI. 2.20).

Remarks. Found in the same area as A. varia and A.
mitchelliana this species differs most significantly from the
former two by a distinct anatomy of the interior penial wall.
The penis anatomy of the present sample corresponds with
that shown by Solem (1981a). This species is represented in
the molecular tree shown below.

Molecular phylogeny

Phylogenetic trees were reconstructed for a sequence data
set with a length of 655 bp that comprised 53 sequences
representing 28 species of Amplirhagada and two out-group
sequences of Carinotrachia. DNA was successfully extracted
from up to 20 years old museum samples but it was not
possible to amplify the mitochondrial COI fragment from
all studied Amplirhagada species because DNA quality
was sometimes low. Nevertheless, the phylogeny contains a
representative subsample of the species treated in this paper.

Rates of transitions and transversions were plotted against
sequence divergence (corrected after Tamura & Nei, 1993)
by use of DAMBE (Xia & Xie, 2001) in order to test for
substitution saturation. Both curves showed nearly linear
relationships indicating no significant levels of saturation. A
hierarchical likelihood ratio test implemented in Treefinder
revealed the generally time reversible model of sequence
evolution (GTR+I+r) as the best-fit model. Settings were
adjusted accordingly in the following ML and BI analyses.

Trees obtained by the three different analytical techniques
(MP, ML, BI) revealed widely congruent topologies that
differed only in minor aspects. The maximum parsimony
tree found by the MP Ratchet (length 2052 steps, Ci = 25,
Ri = 66) showed A. napierana as the most basal offshoot
within the genus (Pig. 88). By contrast, the BI tree (Pig. 89)
and the ML trees (not depicted) showed this species in an
unresolved relationship with respect to the out-group and
all other Amplirhagada species. Next to A. napierana, all
trees reveal essentially five species clusters (Pigs. 88-89).
The basal relationships between these clusters are not
well supported in terms of branch support values and are
unresolved in the Bayesian tree (Pig. 89). Nodal support
within the clusters, however, is usually high. The sister pair
of A. bujfonensis and A. yorkensis (Cluster 1) is consistently
shown in a position basal to all other clades. Monophyly
of this species pair receives high nodal support. The four
other clusters contain species that do not always restricted
to certain geographical areas. Cluster 2 (A. kimberleyana
and other species) encompasses taxa from Boongaree Island
and all coastal areas south of it. Most of these species have
rather dome-shaped shells (except for A. kimberleyana )
and are free sealers; the branching order is consistently
revealed by all three trees. Cluster 3 contains species that
are distributed to areas between the Napier Ranges in the
hinterland (A. percita) to islands of the central Kimberley
coast (A. katerana, A. berthierana). The MP cladogram
indicates that species of Cluster 4 from the Mitchell Plateau
(A. castra, A. mitchelliana ) and A. kessneri from Bigge
Island are more closely related to Cluster 2, while the BI
shows the relationships between them and A. tricenaria
as unresolved. Cluster 5 contains species from islands in
the Admiralty Gulf and the Mitchell Plateau (A. varia and
others).

Pair-wise genetic distances within species did not exceed
the maximum of 4% found between the two sequences
of A. combeana. Average pair-wise distances between
species usually range between 26 and 35% (corrected
sequence divergence, Tamura & Nei, 1993) or 16 and 23%
of uncorrected p-distance. Average interspecific distances
between some species, which are very close to each other in
the phylogenetic tree were not lower than 10% (Tamura-Nei
distances) or 8% (p-distances).

Kohler: New Amplirhagada species from the Kimberley

275

Discussion

Species delimitation by means of morphology
and molecules

Solem (1981a) established that the most informative and
convenient source of information for identifying species
of Amplirhagada is the genital anatomy, particularly the
structure of the inner penial wall and development of the
main stimulatory pilaster. Naturally, the development and
relative size of some genital structures depends on the
maturity of the animal and on its actual reproductive state.
Among fully mature specimens (with adult shell features and
fully developed albumen gland), a simple pattern of seasonal
variation is found with the genitalia being inactive and
reduced in size during the early to middle dry season between
May and August (Solem & Christensen, 1984). As figures and
descriptions of genital features herein are based exclusively
on the examination of adult specimens collected at the end
of the dry or during the wet season (November to April), this
phenomenon should not seriously affect the assessment of
these features with respect to the delimitation of taxa.

The marked interspecific differentiation in the repro¬
ductive anatomy and the observation that differences are
greater when species occur in sympatry led Solem (1981 a) to
argue that the structure of penial and vaginal walls may have
a significant function in mate recognition. Recent findings of
reproductive character displacement in genital morphology
of Asian camaenids ( Satsuma ) seem to provide evidence in
support of this idea (Kameda et al., 2009).

Contrasting the significance of reproductive anatomy,
Solem (1981a) found that shell features often are of limited
value for the recognition of species. Species that occur
under allopatric conditions may exhibit quite similar shells
while conversely species found in sympatry are usually
well recognisable by their shells. Radulae and jaws have
been considered rather useless for the differentiation of taxa
at species level by Solem (1981a). Similar assessments of
anatomical characters have been made also with regard to other
Australian (Willan et al., 2009; Kohler, 2010), Asian (e.g.,
Chiba, 1999a; Kameda et al., 2007; Chiba & Davison, 2008)
and New Guinean Camaenidae (e.g., Jordaens et al., 2009).

In the present study, the observations and conclusions
of Solem (1981a) with regard to the significance of
morphological characters for the recognition of species are
generally confirmed. The considerable genetic differentiation
of species supports the appraisal of reproductive features.
Species delimited by their anatomy are differentiated on
average by Tamura-Nei distances of about 30% in the
mitochondrial marker COI. In the few exceptions to this
general rule, species were found to be separated by as little
as 10% sequence divergence (such as A. comheana and
A. montesquieuana). However, even these rates of genetic
differentiation do not seriously contradict the supposed status
as distinct species. Corresponding with Solem’s (1981a)
statements on Amplirhagada (see also Solem, 1979, 1981b,
1984, 1985, 1988, 1993; 1997 for other camaenid groups
from Western Australia), the anatomy of the penis (i.e.,
the structure and development of inner penial wall, main
stimulatory pilaster, and penial verge) is most informative
on the species level. Differentiation within populations was
generally negligible compared to the well-marked differences
between species. Also, the differences in the genital anatomy

between sympatric species (e.g., A. kessneri and A. decora
on Bigge Is., A. boongareensis, A. gibsoni, A. regia on
Boongaree Is) are often pronounced - well in agreement
with the statement of Solem (1981a). Only in one allopatric
species pair, A. solemiana and A. indistincta, were virtually
no differences observed in the genital anatomy. Both species
occur on neighbouring islands and form a sister pair. They
are genetically well-differentiated and can be distinguished
by their shell size (Fig. 8).

Shell characters have otherwise been revealed as being
of limited value for the discrimination of Amplirhagada
species. Shells of most species are within a size range of
15 to 25 mm in diameter with few exceptions from this rule
(Table 1). The general shape of the shell (i.e., broadly conical
or dome-shaped) is a feature often found in a clade with
little differentiation among the constituent species. This is
illustrated by the example of the clade of dome-shaped species
(A. uwinsensis, A. sphaeroidea, A. regia, and A. camdenensis),
all of which have very similar shells. It is correct, though, that
sympatric species tend to show more pronounced differences
in the shell, which may facilitate species recognition even
without examination of the genitalia. This is exemplified
by the sympatric (an in part closely related) species on
Boongaree and Bigge islands, which are readily recognisable
by their remarkably different shells.

Patterns of endemism and diversity among
Kimberley camaenids

In the Kimberley Rainforest Survey (1987-1988) large
numbers of camaenid species were found only in one or two
rainforest patches (Solem, 1991; Solem & McKenzie, 1991).
These patches and similarly suitable habitats are usually
surrounded by areas that are uninhabitable for the snails.
Most camaenid land snails have limited dispersal abilities.
For snails, these patches function as islands just as the “real
islands” of the Bonaparte Archipelago off the mainland coast,
which are also inhabited by these camaenids. If islands are
simply seen as areas surrounded by hostile environments
(Gittenberger, 2007), from a biogeographical point of view
the entire Kimberley region is best understood as a huge
island archipelago. Camaenid land snails in the Kimberley
therefore exhibit distributions that are similar with to those
found in snails on oceanic islands. Most camaenid species in
the Kimberley have very short ranges with a calculated median
range of only 20 km This holds true not only for species
of Amplirhagada, but also for many other genera with few
notable exceptions, such as species of Xanthomelon inhabiting
open woodland (Solem, 1991). This highly insular distribution
is inevitably connected with narrow-range endemism in many
camaenids groups in the Kimberley. Documenting these
patterns will provide the basis for a better understanding of
the biogeography of the region and the evolution of its biota
and facilitate the development of appropriate conservation
strategies (e.g., Willan et al., 2009; Kohler, 2010).

Solem & McKenzie (1991) concluded that camaenid land
snails reveal geographic patterns, which reflect their low
dispersal abilities and multiple centres of endemism; the
phylogeographic relationships within Amplirhagada are fully
consistent with this. The mitochondrial phylogeny reveals a
basal polytomy, which is not caused by sequence saturation
and hence considered as a “hard polytomy”. The branching
pattern of the tree corresponds closely with geographical

276

Records of the Australian Museum (2010) Vol. 62

Carinotrachia sp. WAM S36600
Carinotrachia sp. AM C.463606
A. napierana AM C. 150457
A. yorkensis FMNH 219151
A. yorkensis WAM S36976
A. yorkensis AM C.463711
A, buffonensis WAM S41484
A. buffonensis WAM S41483
A, buffonensis FMNH 219365
A. kimberieyana AM C.463760
A. gemina WAM S41479
A. gemina FMNH 219049
A. basilica AM C.463720
A. boongareensis AM C.463706
A. gibsoni WAM S36650
A. gibsoni WAM S36651
A. camdenensis WAM S37005
A. camdenensis AM C.463722
A. regia WAM S36647
A. regia WAM S36648
A. uwinsensis WAM S36991
A. uwinsensis AM C.463715
A. sphaeroidea AM C.463718
A. sphaeroidea WAM S37030
A. percita AM C. 144028
A. berthierana AM C.463754
A. berthierana WAM S33029
A. katerana AM C.463692
A. katerana WAM S36879
A. katerana WAM S36603
A. katerana AM C.463696
A. euroa AM C.463680
A. euroa WAM S37083
A. tricenaria AM C.463757
A, kessneri WAM S36876a
A. kessneri WAM S36876b
A. kessneri AM C.463701
A. mitchelliana WAM S41462
A. mitchelliana AM C. 144039
A. castra AM C.460966
A. montesquieuana AM C.463748
A. descartesana WAM S41491
A. descartesana FMNH 219276
A. mckenziei FMNH 219250
A. combeana AM C.463725
A. combeana WAM S41455
A. ponderi FMNH 219268
A. varia AM C.460963
A. solemiana WAM S36595
A solemiana AM C.463681
A. indistincta AM C.463724
A. indistincta AM C.463686
A. indistincta WAM S36864

Kohler: New Amplirhagada species from the Kimberley

277

100 I

” I

100

95

0.1

I—I

Carinotrachia sp. WAM S36600
Cahnotrachia sp. AM C.463606

A. napierana AM C. 150457

100

m~ a

hi

93

100

96

72

59

99

95

A. buffonensis WAM S41484
buffonensis WAM S41483
A. buffonensis FMNH 219365
A. yorkensis WAM S36976
A. yorkensis AM C.463711
A. yorkensis FMNH 219151
— A. tricenaria AM C.463757

A. percita AM C. 144028

100

67

100

100

100

100

n

94

92

100

100

57-72

100

100 /
91-99
100-97'
72-77

100

100

A. berthierana AM C.463754
A. berthierana WAM S33029
A. katerana AM C.463692
A. katerana WAM S36879
A. katerana WAM S36603
A. katerana AM C.463696

A. mitcheliiana WAM S41462
— A. castra AM C.460966
— A. mitcheiliana AM C. 144039
A, kessneri WAM S36876a
A. kessneri WAM S36876b
A. kessneri AM C.463701

- A. combeana AM C.463725

- A. combeana WAM S41455
A. mckenziei FMNH 219250
A. montesquieuana AM C.463748
A. descartesana WAM S41491
A. descartesana FMNH 219276

i— A. varia AM C.460963
- A. ponderi FMNH 219268

76-691.1 od" ' nc ^' s ^ ncta AM C.463724
^ 33 A indistincta AM C.463686

95-75L| 1 A. indistincta WAM S36864

A. solemiana WAM S36595
A. soiemiana AM C.463681
A, euroa AM C.463680
A. euroa WAM S37083

A. kimberleyana AM C.463760
ioo r gemina WAM S41479
A. gemina FMNH 219049
ioo r A. camdenensis WAM S37005
A. camdenensis AM C.463722
A. sphaeroidea AM C.463718
A. sphaeroidea WAM S37030
A. uwinsensis WAM S36991
A. uwinsensis AM C.463715
A. regia WAM S36647
A. regia WAM S36648

Figure 89. Bayesian phylogram for COI. Numbers on
branches indicate support of the shown topology by Bayesian
posterior clade probabilities/expected likelihood weights of
local rearrangements of tree topology (LR-ELW).

o |- A. basilica AM C.463720

85 I

7fl 100 |- A. boongareensis AM C.463706

97 ~ j 100 r A. gibsoni WAM S36650
9^1 A. gibsoni WAM S36651

Table 1 . Shell parameters of Amplirhagada species, given in the form maximum-minimum (mean±standard deviation) measures of shells for n specimens.

278

Records of the Australian Museum (2010) Vol. 62

of

rn

in'

ip

CP

s

of

s

rn

NO

rn

NO

rn

s

Ip

rn

in'

CP

S

rn

rn

rn

of

o

O

o

o

o

o

O

o

o

o

O

o

O

o

o

o

O

o

o

O

o

O

o

o

o

o

o

O

O

o

O

o

O

o

o

O

o

o

o

O

o

O

o

+i

+1

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

ti

tl

tl

tl

tl

tl

o

NO

co

OO

it

CO

t-H

ON

if

it

04

04

ON

in

04

t-H

04

NO

t-H

O'

it

It

it

ON

\a

VO

NO

NO

O'

O'

O'

NO

VO

O'

O'

NO

OO

NO

OO

oo

O'

O'

O'

ON

O'

oo

in

in

8

o

O

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

Q

o

On

ON

^H

o

NO

it

o

T— H

iO

04

it

T— H

O'

On

NO

m

On

04

04

On

o

04

PC

o

NO

O'

oo

oo

O'

O'

O'

oo

OO

O'

o

O'

oo

OO

O'

oo

O'

OO

OO

NO

NO

<?

<?

<?

<?

<?

<?

?

?

7

?

?

?

?

?

?

7

cp

?

?

<?

(N

oo

ON

On

oo

it

oo

in

in

in

it

in

On

in

in

NO

oo

o

O'

in

On

iT

NO

It

04

<n

ON

NO

m

ON

NO

it

oo

O

ON

©

in

it

On

CO

in

NO

vo

vo

in

in

in

NO

NO

NO

NO

NO

NO

NO

NO

in

NO

NO

O'

NO

>n

NO

O'

oo

NO

NO

O'

OO

S'-

O'

NO

o-

NO

C-'

NO

O'

oo

ON

O'

o-

oo

oo

it

in

>n

in

o

o

o

o

o

o

O

o

o

o

o

o

O

o

o

O

o

o

o

o

o

o

o

o

o

o

o

o

o

o

O

o

o

o

O

o

o

o

o

o

O

o

o

o

£

oT

of

of

CO

d"

CO

CO

04

04

<N

CO

it

CO

CO

O)

04

04

04

T—H

NO

CO

3

T—H

OI

o

o

d

tl

o

d

tl

d

tl

o

O

O

O

o

o

o

o

O

O

O

o

o

O

o

o

o

o

+i

+1

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

43

oo

NO

NO

o

in

m

CO

CO

O'

o

in

CO

NO

NO

On

CO

CO

o

04

NO

CO

in

£

3

it

'it

in

it

it

it

it

it

in

it

it

it

in

it

it

in

in

NO

in

in

it

it

O

on

o

CO

oo

oo

in

m

NO

o

O'

o

in

04

NO

m

OO

NO

ON

in

o

U

<D

T

in

1

in

1

in

1

T

T

T

T

in

1

in

1

in

1

T

m

1

in

1

in

1

in

1

in

1

m

1

NO

1

in

1

in

1

T

m

1

X i

VO

oo

o

it

04

in

in

04

CO

CO

ON

NO

NO

o

in

CO

oo

CO

o

in

in

it

o

o

O'

04

04

oo

NO

NO

it

o

in

in

CO

it

it

o

oo

it

CO

NO

8

a

c

3

it

it

'it

'it

'it

it

it

it

it

CO

it

it

it

it

it

it

it

it

it

CO

it

it

m

it

m

it

it

it

it

m

in

in

m

it

m

it

in

m

m

it

it

it

3

44

44

£

o

£

o

d

44

44

44

44

44

of

NO

H

S3

S3

a

a

a

a

a

cb

43

43

hd

3

3

3

3

3

v

v

S3

Y

Y

Y

Y

Y

in

oo

_o

M

M

o

1

dJ

<D

as

1

<D

in

M

1

<D

M

44

m

44

44

o

44

CD

(D

m

1

(D

44

1

<D

as

1

dJ

dJ

44

44

1

dJ

44

1

<D

as

44

44

as

as

P 4

a

a

C/5

C/3

a

C/)

e

a

2

a

a

2

a

O)

V)

2

<y>

a

05

05

a

a

05

a

a

a

a

1

-e

3

3

a

o

o

o

o

o

a

3

o

o

in

3

3

3

3

3

o

o

in

o

3

o

o

o

o

3

3

o

3

o

o

3

3

o

o

o

04

in

8

o

cs

o

a

o

o

cs

o

04

04

o

o

o

o

o

o

3

04

o

o

o

u

o

o

o

o

o

o

o

as

o

cs

u

o

04

04

04

04

o—\

NO

it

Co

o

ti

o

NO

o

ti

oo

On

oo

it

CO

t>

it

o

+1

3 s

o

CO

o

tl

O'

in

<?

?

oo

in'

o

Co

ti

ON

o

tl

NO

Co

O

tl

04

o

tl

NO

o

tl

NO

o

tl

ON

o

tl

o

of

Ip

CP

P

>n

43

£

it

d

+i

CO

4

04

o

d

tl

CO

o

ti

m

o

o

d

04

o

ti

NO

it

04

cb

cb

tl

It

tl

o

ti

o

ti

o

ti

ON

On

x — b

xx

ON

^^

^^

^^

s — ^

ON

3

O'

^'

S -

O'

o

oo

^^

04

^^

it

0^

O'

CO

t-H

CO

NO

^^

3

oo

OO

_a

o

o

00

04

CO

CO

in

it

it

04

it

in

in

It

oi

3

3

°5

NO

On

q

t-h

o

1

t-H

t-H

t-H

o

t-H

t-H

t-H

t-H

04

o

It

o

t-H

—H

1

t—H

a

On

t-H

a

t-H

1

oo

1

1

1

1

i

oo

oc

ON

1

NO

i

1

CO

ON

1

o

1

OO

NO

1

1

oo

1

ON

it

O'

1

o

in

oo

oo

T^

oo

T—H

in

T^

it

T—H

1

ON

i

in

1

1

VO

o

CO

CO

ON

o

o

04

o

oo

ON

it

CO

o

o

O

o

o

04

O'

o

04

it

o

3

o

04

cb

3

rn

04

it

o

NO

q

04

oo

it

bX)

ON

oo

ON

d

On

oo

On

On

On

NO

3

NO

3

3

3

OO

3

On

3

NO

O'

O'

OO

oo

oo

O'

m

6

ON

OO

O'

t-H

oo

m

t-H

oo

O'

NO

ON

6

o

O

o

o

d

ti

d

tl

o

o

o

T—H

T—H

o

T—H

o

T—H

o

o

t—h

o

o

o

O

T—H

+1

+1

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

tl

Q

vo

NO

O'

ON

>n

O'

O'

CO

ON

t-H

CO

ON

o

CO

It

It

ON

it

NO

It

CO

04

ON

O'

On

On

O'

oo

NO

NO

O'

04

On

On

ON

04

O

o

CO

CO

o

OO

OO

to

OO

04

04

04

04

04

04

04

<D

Id

ro

'it

04

O

ON

oo

04

OO

oo

O

OO

O

NO

04

it

NO

in

O

ON

04

OO

oo

CO

a

o

oo

t— H

,-H

OO

ON

oo

O'

oo

in

t^H

04

t-H

CO

CO

T— H

it

NO

t-H

ON

OO

oo

On

04

T^

04

04

t-H

T— H

T^

T— i

04

04

04

04

04

04

04

04

04

04

t-H

t— H

T^

H

3

1

m

04

1

04

in

1

in

o

1

in

O

o

o

1

in

O

CO

o

OO

1

m

1

CO

1

OO

o

1

04

o

a

o

a

1

NO

04

a

04

1

04

1

OO

it

1

ON

04

1

CO

1

CO

m

1

in

ON

1

On

oo

ON

NO

oo

oo

o

oo

NO

oo

ON

>n

o

T—H

in

O'

NO

O'

ON

O'

o

OO

o

>n

o

r—H

CO

OO

04

ON

O

CO

it

T—H

CO

OO

T—H

O'

oo

O'

oo

in

O'

NO

OO

’ 1

04

’ 1

04

04

04

04

04

04

04

04

04

04

04

04

04

04

in'

©

PI

o7

ixi

S

S'

00

s

Ip

rn

o7

s

rn

o

s

p

00

o

VO

1—H

t—H

o

o

o

so

o

T^

o

T—4

it

t-H

T— (

o

T^

t-H

T—|

o

o

S'

o

+i

o

tl

tl

tl

tl

tl

o

tl

tl

tl

tl

t—H

tl

tl

ti

tl

tl

tl

tl

tl

tl

o

+i

in

oo

oo

It

NO

ti

CO

ON

o

oo

tl

CO

>n

NO

NO

o

CO

>n

o

Ph'

cb

(N

CO

'it

04

CO

T^

it

CO

in

04

O'

it

oo

NO

It

OO

NO

o

CO

in

+1

T^

d

04

04

d

£

On

oo

NO

On

it

o

in

On

CO

NO

oo

o

T—H

oo

04

OO

On

o-

OO

M

cb

04

'it

NO

cb

it

cb

q

it

O

cb

o

O)

ON

ON

NO

ON

OO

04

3

NO

it

T—H

CD

04

04

04

it

04

o

43

1

co

o

7

o

O'

C^4

1

in

1

CO

CO

1

in

OO

oo

7

NO

nA

oo

1

OO

a

O

1

oo

OO

7

04

1

in

o

1

OO

NO

1

O'

OO

a

m

1

in

04

O

a

O'

a

NO

7

1

m

m

oi

co

ON

T^

t-H

04

CO

T^

04

04

o

CO

CO

it

T^

T^

CO

04

T^

CO

it

NO

O';

CO

in

ON

It

o-

CO

It

NO

oo

it

OO

m

t-H

04

CO

it

>n

it

It

o

o

ON

On

ON

04

3

ON

co

o

in

CO

O'

CO

in

OO

CO

NO

NO

it

o

oo

NO

in

04

04

04

O'

CM

H-

04

ON

O'

04

04

it

/ lfl

a

$h

o

C/3

u

(D

C/3

TO

C/5

So

\ri

S

a

a

o

1)

S3

$h

oj

Vh

a

<D

<D

s

a

00

T3

3

>

o

Id

Cu

>>

o

a

3

s

3

-

5a

d

cn

o

a

PQ

&

CS

a

3

OS

©

S3

3

a

o

<D

Q-

a

|

a

a

a

<D

O-

3

a

a

1

a

1

a

a

w

Cu

§

a

a

"3

c/a

^3

a

.69

c?

3

a

po

a

a

rT*

3

a

.3

So .3

So

c^

So

po

3

>0

So

>0

g

e

cs

>■.

§

fo,

So

po

Si

S',

fo,

3

a

So

So

O

a

o

s

o

c

<3

K?

o

2

o

t;

o

a

o

3

o

o

3

o

o

1

o

3

o

o

3

bo

3

o

o

^0

_o

>3

o

JO

o

3

o

i

o

+-*

Co

o

<o

a

•a

o

c3

o

g

o

a

3

o

o

a

o

-a

o

o

o

3

.3

a

o

regia

o

o

3

o

-a

Q-

>p

JO

o

.3

o

s

a

<4S

43

"o

43

•S

43

§

a

IS

5

43

1

a

a

a.

O-

§

S

43

c^

“3

43

£

3

a

cx,

43

3

"3

43

Co

43

a

3

■a

43

Q

43

"3

|

43

43

43

a

43

sS

So

43

43

S'

-3

43

3

3

3

3 3 3

q

q

q

3

3

3

3

3

3

3

3

Table 1 (continued). Shell parameters of Amplirhagada species, given in the form maximum-minimum (mean±standard deviation) measures of shells for n specimens.

Kohler: New Amplirhagada species from the Kimberley

279

cm

cm

<77

C<7

(7T

o

o

O

O

o

O

o

o

?

?

?

?

o

oo

CM

oo

NO

in

CM

oo

q

m

NO

NO

oo

C-

NO

NO

o

o

o

o

o

o

o

o

S-H

n

NO

CM

CO

On

oo

On

NO

i—i

no

q

r-

OO

r-

NO

NO

£

cp

cp

?

?

?

Cp

in

'O

o

(SI

NO

oo

OO

r-

in

oo

oo

oo

NO

m

in

NO

r-

NO

O

in

NO

l>

oo

NO

l>

NO

NO

m

NO

o

o

o

o

o

o

o

o

o

o

o

o

o

o

o

£

C/3

44 s

cm

77 s

47 s

47 s

CM

47 s

47 s

o

o

o

o

o

o

o

o

d

+i

+i

+i

+i

+i

+i

+i

+i

£

q

cq

<N

<S)

'Ct

q

cq

q

d

d

in

in

■ci

q

q"

<-l—1

o

m

q

in

'it

CO

oo

NO

NO

CD

o

T

in

1

in

1

in

1

in

1

T

T

T

£

co

oo

CM

CM

in

^t

oo

CO

oo

CO

NO

CM

q"

q"

in

in

in

in

in

, ci

in

''cl-

q

q

q

q"

q"

a

CM

CM

s

o

+i

o

+i

&

q

44

=3

CM

C

CM

O

'—'

4=

v —'

=3

r-~

y

m

cm

<u

o

03

44

44

44

in

44

in

44

44

44

CM

1

i/3

c/3

c/3

C

C

C

2

C

C

C

C

o

CM

s

=s

CM

CM

o

To

To

£

To

4=

CO

CO

CO

o

2

4=

o

d

o

4=

O

o

CM

S'

q'

in

rn

w '

o

o

o

o

i>

NO

Th

cf

+i

+i

+1

+i

o

o

q

o

CM

On

NO

NO

+i

+1

o

A

%

<si

q

<N

q

o

NO

+1

CM

£

o

o

C/D

oo

CM

ON

CO

d

in

co

CM

CO

<N

in

';

NO

<-w

oo

%

%

%

%

-H

ON

O

1

co

CM

ON

NO

1

in

i

o

4

c*

oo

<si

q

<N

CO

q

OO

O'

CM

t>

'"H

'"H

oo

'"H

ON

'"H

oo

ON

^ — N

^ — N

s

^ — N

. — N

>— N

^ N

^— N

43

m

oo

I>

(N

^1-

o

I>

ON

o

o

o

__ ;

o

__ ;

o

o

+i

+i

+i

+i

+i

+i

+1

+1

Q

oo

q

o

NO

(N

NO

CM

ON

d

_4

<N

CO

ON

U

ON

U

u

CM

<n

(N

<D

a)

cl

co

CM

o

O

OO

NO

OO

i>

U

co

co

U

ON

ON

ON

On

£h

q

(SI

C±

CM

(SI

q

q

q

'd

l

ON

ON

q

I

(N

(N

1

CM

NO

1

CM

NO

NO

m

d

o

,-H

o

CM

(N

'Ct

oo

On

d

ON

OO

ON

d

U

1—1

CM

CM

(SI

CM

(N

<N

,—l

1—1

,—l

' —l

' —l

1—1

1—1

,—l

r-7

t-d

00

4d

NO

in'

o

o

o

o

o

o

+i

+i

+i

+1

+1

+i

+i

£

•

no

ON

1";

(N

q

q

o

+1

in

in

d

CM*

CO

q

r-H

—-h

—,

i—H

. ,

t"

s—^

s—'

s—✓

'--

'w'

^^

s-✓

43

On

OO

<N

CO

CO

q

o

top

oo

l>

in

u

vn

CO

CM

<D

43

°\

1

cq

q

1

in

OO

3

'Ct

1

oo

2

CO

1

CO

CO

2

q

m

q"

in

cn

in

'Ct

d

'ci

d

in

cm"

CO

o

d

On

co

CO

CO

S3

m

(N

oo

1—1

1—1

1—1

CO

ON

'"m

c

,o

d

3

pq

5h

S3

<D

C/3

w

a

•2

a

a

q

,52

a

;a

<D

S3

00

S3

C?

Ri

a

Pn

a

a

S3

a

a

s

d 1

>p

>N

ia

>>

a

a

a

Co

o

a

a

.a

O

d

a

a

d

a

a

d

d

4

.a

d

a

’o

*a

'o

o

-a

d

d

a

d

-3

<

4=

CO

4=

a

4=

S 4=

4=

43

a 5

S3

a

a

a

a

-a

a

Co

-a

a

<50

"a

q;

distributions, with clades comprising species from regional
island groups plus the adjacent mainland. This is consistent
with several clades having colonized islands more or less
simultaneously from different centres of origin along the
mainland coast. Rates of genetic differentiation between
species are generally high, suggesting that species became
separated a long time ago. Moreover, genetic distances
between the mainland species are not significantly different
from those between the island species or between species
from islands and mainland. This seems to indicate that for
the evolution of these snails it is not relevant whether their
insular habitats are isolated by means of surrounding sea or
other hostile environments.

It would be helpful to know when exactly the islands of
the Bonaparte Archipelago were colonized by Amplirhagada
snails in order to understand whether dispersal or vicariance
played a major role. However, attempting to deduce the age
of the group from estimated rates of sequence evolution,
would be an arbitrary endeavour. Wilke et al., (2009) have
suggested the applicability of a local molecular clock for
small invertebrates. However, it has also been demonstrated
that evolutionary rates on islands may be accelerated (Chiba,
1999b; Millien, 2006). In addition, it remains unclear whether
the clock suggested by Wilke et al. (2009) for freshwater
caenogastropods would be applicable to pulmonates, which
were stated to show accelerated rates of molecular evolution
(Thomaz etal., 1996; Chiba, 1999b; Watanabe & Chiba, 2001;
Pinceel et al., 2005). The latter two phenomena undermine
the reliability of a molecular clock approach by assuming
rates similar to those in caenogastropods. When a very coarse
general estimate of 1 to 2% substitution rates per million years
in the COI gene of invertebrates was considered (Wilke et al.,
2009), observed substitution rates of around 30% were taken
to indicate divergence times for most species of several million
years and that sea level changes during the Pleistocene, which
may have periodically re-connected islands with the mainland,
did not have a great impact on the patterns that may well have
evolved prior to these events.

Johnson et al. (2010) studied Amplirhagada species
on 16 Kimberley islands that are only partly covered by
the present study on a finer spatial scale and came to very
similar conclusions. They showed that the mitochondrial
variation across the populations on the 16 islands correlates
tightly with the geographical patterns. Genetic divergence
was generally found to be low within local populations but
much higher when populations from different islands were
compared. Unfortunately, this study remained inconclusive
with respect to the number of species involved. The authors
found 16-27% sequence divergence in a small fragment of
the 16S rRNA gene between major clades of Amplirhagada
that are separated by 10 to 160 km It is demonstrated herein
that such distances are equivalent to the differentiation at the
species level, which implies that Johnson et al. (2010) dealt
indeed with several distinct, yet mostly unrecognized species.
The phylogeographic patterns further revealed that species are
restricted to single islands or smaller groups of closely adjacent
islands. This finding corresponds well with the results of the
present study. Johnson et al. (2010) concluded that even when
accelerated evolutionary rates of 10 to 12.9 % per million
years were considered, as reported from some land snails
(Thomaz et al., 1996; Chiba, 1999b; Thacker & Hadfield,
2000), the amount of differentiation found in Amplirhagada
would indicate divergence times of species of at least 1.5 to

280

Records of the Australian Museum (2010) Vol. 62

Plate 1. [Shells of Amplirhagada species (natural size, scale bar = 10 mm). Caption given on page 282].

Kohler: New Amplirhagada species from the Kimberley

281

Plate 2. [Shells of Amplirhagada species (natural size, scale bar = 10 mm). Caption given on page 282].

282

Records of the Australian Museum (2010) Vol. 62

Plate 1 (page 280). Shells of Amplirhagada species (natural size, scale bar = 10 mm). (1.1) A. euroa n.sp. holotype
WAM S34601. (1.2) A. euroa n.sp. paratype WAM S37392. (1.3) A. solemiana n.sp. holotype WAM S34602. (1.4) A.
solemiana n.sp. paratype WAM S36572. (1.5) A. indistincta n.sp. holotype WAM S34603. (1.6) A. indistincta n.sp.
paratype WAM S36499. (1.7) A. combeana WAM S41450. (1.8)A. mckenziei n.sp. holotype WAM S34604. (1.9) A.
mckenziei n.sp. paratype FMNH 219253. (1.10) A. ponderi n.sp. holotype WAM S34605. (1.11) A. montesquieuana
n.sp. holotype WAM S34606. (1.12) A. descartesana holotype WAM S34607. (1.13) A. katerana WAM S36580.
(1.14)A. puescheli n.sp. holotype WAM S34608. (1.15) A. decora n.sp. holotype WAM S34609. (1.16) A. kessneri
n.sp. holotype WAM S34610. (1.17) A. berthierana n.sp. holotype WAM S34611. (1.18) A. lamarckiana n.sp.
holotype WAM S34612. (1.19) A. anderdonensis n.sp. holotype WAM S34613. (1.20) A. tricenaria n.sp. holotype
WAM S34614. (1.21) A. regia n.sp. holotype WAM S34615. (1.22) A. boongareensis n.sp. holotype WAM S34616.
Note that foot protrudes from shell in some cases.

Plate 2 (page 281). Shells of Amplirhagada species (natural size, scale bar =10 mm). (2.1) A. gibsoni n.sp. holotype
WAM S34617. (2.2) A. gibsoni n.sp. paratype AMS C463708. (2.3) Amplirhagada sp. Bigge Island, WAM S36953.
(2.4) A. yorkensis n.sp. holotype WAM S34619. (2.5) A. yorkensis n.sp. paratype WAM S37401. (2.6) A. buffonensis
n.sp. holotype WAM S34620. (2.7) A. uwinsensis n.sp. holotype WAM S34621. (2.8) A. uwinsensis n.sp. paratype
WAM S37441. (2.9) A. uwinsensis n.sp. paratype AMS C463716. (2.10) A. sphaeroidea n.sp. holotype WAM
S34622. (2.11) A. basilica n.sp. holotype WAM S34623. (2.12) A. camdenensis n.sp. holotype WAM S34624. (2.13)
A. kimberleyana n.sp. holotype WAM S34625. (2.14) A. gemina n.sp. holotype WAM S34626. (2.15) A. dubitabile
n.sp. holotype WAM S34627. (2.16) A. napierana AMS C150457. (2.17) A. percita AMS C144028. (2.18) A. varia
AMSC460963. (2.19)A. mitchelliana WAM S41462. (2.20)A. castraAMS C460966. Note that foot protrudes from
shell in some cases.

2 million years. In this period of time, particularly during the
Pleistocene, the studied islands were repeatedly interconnected
due to periodically lowered sea levels. However, the species
have maintained distinctiveness on a small geographical scale
over this period of time, which implies that fluctuating sea
levels during Pleistocene did not influence the processes of
genetic differentiation or allopatric speciation.

In general, narrow-range endemics, such as the Kimberley
camaenids, are promising candidates for the study of
speciation and adaptation. While the isolated occurrence
of most species may suggest allopatric speciation as the
predominant mode, Schilthuizen & Scott (2004) have
stressed that for a number of model cases including Western
Australian camaenids, a strong ecological component may
also be involved. Observed cases of character displacement
in sympatric species also points towards the relevance of
ecological factors and sexual selection in these species.
Hence, further studies are needed that address aspects of
speciation, adaptation and radiation in Amplirhagada in
more detail. Markedly different patterns of morphological
versus genetic differentiation were found in the camaenid
snail Rhagada in Dampierland, Western Australia. In
these snails, high levels of morphological differentiation
were accompanied by generally very low levels of genetic
differentiation (Johnson et al., 2004). A comparative study
of the contrasting patterns in these two Western Australian
camaenid genera may help to better understand the factors
that drive the evolution of Australian Camaenidae.

With respect to overall species diversity, the Kimberley
with its approximately 180 known camaenid species (Solem,
1998) is comparable with many textbook cases of mega-

diverse island faunas (Whittaker, 1998). Moreover, given
the large number of still unsurveyed islands off the mainland
coast and other equally inaccessible regions on the mainland,
the number of yet undiscovered species in the Kimberley is
without doubt considerable. Amplirhagada may be taken as a
prime example to estimate the proportion of yet undescribed
species in relation to what is already known. Thirty species
are currently considered as valid (Solem, 1981a, 1988).
However, museum material examined by Solem contains
some further 35 Amplirhagada species to be described, which
includes the 25 unnamed species reported by Solem (1991).
Here, only eleven of these species are described. Additional
species descriptions are based on materials collected during
the ongoing Kimberley Island Survey in 2007-2008 on eight
larger islands. During survey work conducted in 2009 we
found probably another six currently unknown species of
Amplirhagada, which are not included here. Consequently,
at this stage there are 57 named plus 30 yet to be described
Amplirhagada species. These numbers include species
collected on approximately 25 islands of the Bonaparte
Archipelago, which includes the 19 largest islands of this
archipelago. However, the snail fauna of at least 50 larger
islands in this region remains entirely unknown. If only every
second island supports an endemic Amplirhagada species,
which is probably a conservative estimate, than there might
be 25 undiscovered species on top of the numbers given
above. It can only be speculated, how many species await
discovery on the mainland but less than half of the species
diversity in Amplirhagada is likely documented so far. There
is no reason to assume that these figures are different for
other camaenid genera in the area.

Kohler: New Amplirhagada species from the Kimberley

283

Acknowledgments. This work has been conducted as part of the
Kimberley Island Survey of the Western Australian Department
of Environment and Conservation (DEC), a project jointly funded
by the Commonwealth of Australia and the Western Australian
Government. I owe a debt of gratitude to Marlene Vial, Sue Lindsay
and Martin Piischel for their help with sample processing, SEM
work, anatomical drawings and photography. I further wish to thank
Shirley Slack-Smith and Corey Whisson (WAM), Jochen Gerber
(FMNH) as well as Alison Miller and Janet Waterhouse (AM) for
most effectively handling extensive loans from or to their respective
museums. Special thanks are due to Winston Ponder and Norm
McKenzie for encouraging me to work with Kimberley camaenids.
Thanks are also due to everybody who contributed to the successful
implementation of the survey work on the Kimberley Islands. This
includes traditional landowners of the Malaya, Dambimangari, and
Bardi-Jawi people, the Kimberley Land Council and staff of the
DEC and the Australian Museum. Last but not least, I am indebted
to two anonymous reviewers for their constructive and helpful
comments on the submitted version of the manuscript.

References

Barker, G.M., 1999. Naturalised terrestrial Stylommatophora
(Mollusca: Gastropoda). Fauna of New Zealand , vol. 38.
Canterbury, New Zealand: Lincoln.

Cameron, R.A.D., B.M. Pokryszko & F.E. Wells, 2005. Alan
Solem’s work on the diversity of Australasian land snails:
an unfinished project of global significance. Records of the
Western Australian Museum, Supplement 68: 1-10.

Chiba, S., 1999a. Character displacement, frequency-dependent
selection, and divergence of shell colour in land snails
Mandarina (Pulmonata). Biological Journal of the Linnean
Society 66: 465-479.

Chiba, S., 1999b. Accelerated evolution of land snails Mandarina
in the oceanic Bonin Islands: Evidence from mitochondrial
DNA sequences. Evolution 53: 460-471.

http://dx.doi.org/10.2307/2640782

Chiba, S., & A. Davison, 2008. Anatomical and molecular studies
reveal several cryptic species of the endemic genus Mandarina
(Pulmonata: Helicoidea) in the Ogasawara Islands. Journal of
Molluscan Studies 74: 373-382.

http ://dx.doi.org/10.1093/mollus/eyn029

Felsenstein, J., 1985. Confidence limits on phylogenies: an
approach using the bootstrap. Evolution 39: 783-791.

http://dx.doi.org/10.2307/2408678

Folmer, O., M. Black, W. Hoeh, R. Lutz & R. Vrijenhoek, 1994.
DNA primers for amplification of mitochondrial cytochrome
c oxidase subunit I from diverse metazoan invertebrates.
Molecular Marine Biology and Biotechnology 3: 294-299.
Gittenberger, E., 2007. Islands from a snail’s perspective. In
Biogeography, Time, and Place: Distributions, Barriers,
and Islands , ed. W. Renema, pp. 347-363. Heidelberg:
Springer-Verlag.

http://dx.doi.org/10.1007/978-1 -4020-6374-9_11

Iredale, T., 1933. Systematic notes on Australian land shells.
Records of the Australian Museum 19: 37-59.

http ://dx.doi.org/10.3853/j.0067-1975.19.1933.690

Iredale, T., 1938. A basic list of the land Mollusca of Australia—
Part III. Australian Zoologist 9: 83-124.

Iredale, T., 1939. A review of the land Mollusca of Western
Australia. Journal of the Royal Society of Western Australia
25: 1-88.

Jobb, G., A. v. Haeseler & K. Strimmer, 2004. Treefinder:
a powerful graphical analysis environment for molecular
phylogenetics. BMC Evolutionary Biology 4: 18.

http://dx.doi.org/10.1186/1471-2148-4-18

Johnson, M.S., Z.R. Hamilton, C.E. Murphy, C.A. MacLeay, B.
Roberts & PG. Kendrick, 2004. Evolutionary genetics of island
and mainland species of Rhagada (Gastropoda: Pulmonata) in
the Pilbara Region, Western Australia. Australian Journal of
Zoology 52: 341-355.

http://dx.doi.org/10.1071/Z004027

Johnson, M.S., E.K. O’Brien & J.J. Fitzpatrick, 2010. Deep,
hierarchical divergence of mitochondrial DNA in Amplirhagada
land snails (Gastropoda: Camaenidae) from the Bonaparte
Archipelago, Western Australia. Biological Journal of the
Linnean Society 100: 141-153.

http://dx.doi.org/10.1111/j. 1095-8312.2010.01407.x

Jordaens, K., L. Bruyndoncx, J. Van Goethem & T. Backeljau,
2009. Morphological and anatomical differentiation of three land
snails of the genus Rhynchotrochus (Gastropoda: Pulmonata:
Camaenidae). Journal of Molluscan Studies 75: 1-8.

http://dx.doi.org/10.1093/mollus/eyn035

Kameda, Y.C., A. Kawakita & M. Kato, 2007. Cryptic genetic
divergence and associated morphological differentiation in
the arboreal land snail Satsuma (Luchuhadra) largillierti
(Camaenidae) endemic to the Ryukyu Archipelago, Japan.
Molecular Phylogenetics and Evolution 45: 519-533.

http://dx.doi.org/10.1016/j.ympev.2007.03.021

Kameda, Y.C., A. Kawakita & M. Kato, 2009. Reproductive
character displacement in genital morphology in Satsuma land
snails. The American Naturalist 173: 689-697.

http://dx.doi.org/10.1086/597607

Kohler, F., 2010. Three new species and two new genera of land
snails from the Bonaparte Archipelago in the Kimberley,
Western Australia (Pulmonata, Camaenidae). Molluscan
Research 30: 1-16.

Larget, B., & D.L. Simon, 1999. Markov chain Monte Carlo
algorithms for the Bayesian analysis of phylogenetic trees.
Molecular Biology and Evolution 16: 750-759.

Millien, V., 2006. Morphological evolution is accelerated among
island mammals. PLoS Biology 4: 1863-1868.

Nixon, K.C., 1999. The Parsimony Ratchet, a new method for
rapid parsimony analysis. Cladistics 15: 407-414.

http://dx.doi.org/10.1111/j.1096-0031.1999.tb00277.x

Pinceel, J., K. Jordaens & T. Backeljau, 2005. Extreme mtDNA
divergences in a terrestrial slug (Gastropoda, Pulmonata,
Arionidae): accelerated evolution, allopatric divergence
and secondary contact. Journal of Evolutionary Biology 18:
1264-1280.

http://dx.doi.org/10.1111/j.1420-9101,2005.00932.x

Ronquist, F., & J.P Huelsenbeck, 2003. MRBAYES 3: Bayesian
phylogenetic inference under mixed models. Bioinformatics
19: 1572-1574.

http://dx.doi.org/10.1093/bioinformatics/btg180

Smith, E.A., 1894. On the land-shells of Western Australia.

Proceedings of the Malacological Society of London 1: 84-99.
Solem, A., 1979. Camaenid land snails from Western and central
Australia (Mollusca: Pulmonata: Camaenidae). I. Taxa with
trans- Australian distribution. Records of the Western Australian
Museum Supplement 10: 5-142.

Solem, A., 1981a. Camaenid land snails from Western and central
Australia (Mollusca: Pulmonata: Camaenidae). II. Taxa from
the Kimberley, Amplirhagada Iredale 1933. Records of the
Western Australian Museum Supplement 11: 147-320.

Solem, A., 1981b. Camaenid land snails from Western and central
Australia (Mollusca: Pulmonata: Camaenidae). III. Taxa from
the Ningbing Ranges and nearby areas. Records of the Western
Australian Museum, Supplement 11: 321-425.

Solem, A., 1984. Camaenid land snails from Western and central
Australia (Mollusca: Pulmonata: Camaenidae). IV. Taxa from
the Kimberley, Westraltrachia Iredale, 1933 and related genera.
Records of the Western Australian Museum, Supplement 17:
427-705.

284

Records of the Australian Museum (2010) Vol. 62

Solem, A., 1985. Camaenid land snails from Western and central
Australia (Mollusca: Pulmonata: Camaenidae). V. Remaining
Kimberley genera and addenda to the Kimberley. Records of
the Western Australian Museum, Supplement 20: 707-981.

Solem, A., 1988. New camaenid land snails from the northeast
Kimberley, Western Australia. Journal of the Malacological
Society of Australia 9: 27-58.

Solem, A., 1991. Land snails of Kimberley rainforest patches
and biogeography of all Kimberley landsnails. In Kimberley
rainforests of Australia, ed. N.L. McKenzie, R.B. Johnston,
& RG. Kendrick, pp. 145-246. Canberra: Surrey Beatty &
Sons and Department of Conservation and Land Management
Western Australia.

Solem, A., 1993. Camaenid land snails from Western and central
Australia (Mollusca: Pulmonata: Camaenidae). VI. Taxa from
the Red Centre. Records of the Western Australian Museum,
Supplement 43: 983-1489.

Solem, A., 1997. Camaenid land snails from Western and central
Australia (Mollusca: Pulmonata: Camaenidae). VII. Taxa from
Dampierland trough the Nullabor. Records of the Western
Australian Museum, Supplement 50: 1461-1906.

Solem, A., 1998. Family Camaenidae. In Mollusca: The Southern
Synthesis, ed. PL. Beesley, G.J.B. Ross, & A. Wells, pp.
1112-1113. Melbourne: CSIRO Publishing.

Solem, A., & C. Christensen, 1984. Camaenid land snail
reproductive cycle and growth patterns in semiarid area of
north-western Australia. Australian Journal of Zoology 32:
471-491.

http://dx.doi.org/10.1071/Z09840471

Solem, A., & N.L. McKenzie, 1991. The composition of land
snail assemblages in Kimberley rainforests. In Kimberley
Raonforests of Australia, ed. N.L. McKenzie, R.B. Johnston
& PG. Kendrick, pp. 247-263. Canberra: Surrey Beatty &
Sons and Department of Conservation and Land Management
Western Australia.

Strimmer, K., & A. Rambaut, 2002. Inferring confidence sets of
possibly misspecified gene trees. Proceedings of the Royal
Society of London B Biological Sciences 269: 137-142.

http://dx.doi.org/10.1098/rspb.2001.1862

Tamura, K., & M. Nei, 1993. Estimation of the number of
nucleotide substitutions in the control region of mitochondrial
DNA in humans and chimpanzees. Molecular Biology and
Evolution 10: 512-526.

Thacker, R.W., & M.G. Hadfield, 2000. Mitochondrial phylogeny
of extant Hawaiian tree snails (Achatinellidae). Molecular
Phylogenetics and Evolution 16: 263-270.

http://dx.doi.org/10.1006/mpev.2000.0793

Thomaz, D., A. Guiller & B. Clarke, 1996. Extreme divergence of
mitochondrial DNA within species of pulmonate land snails.
Proceedings of the Royal Society of London B Biological
Sciences 263: 363-368.

http://dx.doi.org/10.1098/rspb. 1996.0056

Watanabe, Y., & S. Chiba, 2001. High within-population mito¬
chondrial DNA variation due to microvicariance and population
mixing in the land snail Euhadra quaesita (Pulmonata:
Bradybaenidae). Molecular Ecology 10: 2635-2645.

http://dx.doi.org/10.1046/j.0962-1083.2001.01388.x

Whittaker, R.J., 1998. Island Biogeography. Ecology, Evolution
and Conservation. Oxford, NY: Oxford University Press.
Wilke, T., R. SchultheiB & C. Albrecht, 2009. As time goes
by: A simple fool’s guide to molecular clock approaches in
invertebrates. American Malacological Bulletin 27: 25-45.

http://dx.doi.org/10.4003/006.027.0203

Willan, R.C., F. Kohler, V., Kessner & M.F. Braby. 2009.
Description of four new species of limestone associated
Torresitrachia land snails (Mollusca: Pulmonata: Camaenidae)
from the Katherine District of the Northern Territory, with
comments on their conservation. The Beagle. Records of the
Museums and Art Galleries of the Northern Territory 25:
85-98.

Xia, X., & Z. Xie, 2001. DAMBE: Data analysis in molecular biology
and evolution. Journal of Heredity 92: 371-373.

http://dx.doi.org/10.1093/jhered/92.4.371

Manuscript submitted 3 August 2009, revised 30 December 2009, and
accepted 31 December 2009.

© The Author, 2010. Journal compilation © Australian Museum, Sydney, 2010
Records of the Australian Museum (2010) Vol. 62: 285-392. ISSN 0067-1975
doi:10.3853/j.0067-1975.62.2010.1556

A Revision of the Australian Funnel-web Spiders
(Hexathelidae: Atracinae)

Michael R. Gray

Australian Museum, 6 College Street, Sydney NSW 2010, Australia
mikegray7@optusnet.com.au

Abstract. The Australian funnel-web spiders of the subfamily Atracinae are revised. The Atracinae are
divided into three genera— Atrax O.P.-Cambridge with three species (two new), Hadronyche L. Koch
with 31 species (18 new), and Illawarra n.gen. with one new species. Hadronyche is divided into four
species groups—the lamingtonensis, adelaidensis, infensa and cerberea groups. Fourteen previously
described species are redescribed or diagnosed. Twenty one new species are described: Atrax sutherlandi
n.sp., A. yorkmainorum n.sp., Illawarra wisharti n.sp., Hadronyche alpina n.sp., H. emmalizae n.sp., H.
marracoonda n.sp., H. monaro n.sp., H. tambo n.sp., H. nimoola n.sp., H. mascordi n.sp., H. jensenae
n.sp., H. orana n.sp., H. lynabrae n.sp., H. kaputarensis n.sp., H. levittgreggae n.sp., H. macquariensis
n.sp., H. walkeri n.sp., H. raveni n.sp., H. lamingtonensis n.sp., H. annachristiae n.sp. and H. monteithi
n.sp. Two species described from Melanesia, H. hirsuta Rainbow and Styphlopis insularis Rainbow, are
regarded as nomen dubia.

Gray, Michael R., 2010. A revision of the Australian funnel-web spiders (Hexathelidae: Atracinae). Records of the
Australian Museum 62(3): 285-392.

Contents

Taxonomic history.288

Material and methods.289

Notes on descriptions.289

Abbreviations.290

Systematics.290

Family Hexathelidae Simon.290

Subfamily Atracinae Hogg.290

Comments on morphology.292

Keys to Atracinae.294

Key to genera.294

Key to Atrax species.294

Key to Hadronyche species.294

Atrax O.P.-Cambridge.297

Atrax robustus O.P.-Cambridge.299

Atrax yorkmainorum n.sp.302

Atrax sutherlandi n.sp.304

Illawarra n.gen.308

Illawarra wisharti n.sp.308

286

Records of the Australian Museum (2010) Vol. 62

Hadronyche L. Koch.311

cerberea species group.312

Hadronyche cerberea L. Koch.312

Hadronyche versuta (Rainbow).316

Hadronyche emmalizae n.sp.319

Hadronyche formidabilis (Rainbow).322

Hadronyche alpina n.sp.326

Hadronyche venenata (Hickman).329

Hadronyche marracoonda n.sp.332

Hadronyche tarnbo n.sp.335

Hadronyche monaro n.sp.337

Hadronyche mascordi n.sp.339

Hadronyche nimoola n.sp.342

Hadronyche modesta (Simon).346

Hadronyche meridiana Hogg.348

Hadronyche jensenae n.sp.351

Hadronyche pulvinator (Hickman).354

infensa species group.355

Hadronyche infensa (Hickman).355

Hadronyche orana n.sp.358

Hadronyche valida (Rainbow & Pulleine).361

Hadronyche lynabrae n.sp.364

Hadronyche kaputarensis n.sp.367

Hadronyche macquariensis n.sp.368

Hadronyche walkeri n.sp.371

Hadronyche levittgreggae n.sp.374

lamingtonensis species group.377

Hadronyche raveni n.sp.378

Hadronyche lamingtonensis n.sp.379

Hadronyche annachristiae n.sp.381

Hadronyche monteithi n.sp.383

Hadronyche anzses Raven.384

adelaidensis species group.385

Hadronyche flindersi (Gray).385

Hadronyche adelaidensis (Gray).386

Hadronyche eyrei (Gray).387

References.390

Appendix 1.391

Appendix 2.392

The atracine spiders include some of the most venomous
animals in the world. They have been responsible for many
serious envenomations and at least 13 deaths in Australia
(Sutherland & Tibballs, 2001). Much of their notoriety can be
attributed to a single species, Atrax robustus O.P.-Cambridge,
1877, better known as the Sydney funnel-web spider.

Fourteen valid atracine species have been described
previously (Table 1)—the first by L. Koch (1873) as
Hadronyche cerberea. Work by Gray (1986) resulted in
preliminary reports on atracine spider relationships and
distribution (Gray, 1987, 1988). This publication provides
the first taxonomic revision of the group. It includes re¬
descriptions of valid genera and species, and descriptions of
a new genus ( Illawarra n.gen.) and twenty-one new species
(Atrax sutherlandi n.sp., A. yorkmainorum n.sp., Illawarra
wisharti n.sp., Hadronyche alpina n.sp., H. emmalizae n.sp.,
H. marracoonda n.sp., H. monaro n.sp., H. tambo n.sp.,
H. nimoola n.sp., H. mascordi n.sp., H. jensenae n.sp., H.
orana n.sp., H. lynabrae n.sp., H. kaputarensis n.sp., H.
levittgreggae n.sp., H. macquariensis n.sp., H. walkeri n.sp.,
H. raveni n.sp., H. lamingtonensis n.sp., H. annachristiae
n.sp., H. monteithi n.sp.). A full list of atracine species is
given in Appendix 1. Recent genetic studies (e.g., Beavis &

Rowell, 2006) and morphological observations suggest that
many more species await characterization.

Valid distribution records of atracine spiders are limited
to eastern Australia, including Tasmania and the Gulf
region of South Australia—the Eyre Peninsula marks the
current western limit of the group (Fig. 2; Gray, 1987).
The description of Hadronyche anzses Raven, 2000 from
the Mosman region in north Queensland greatly extended
knowledge of the northern distribution of the group from its
former known limits in southeastern Queensland.

The Atracinae are typically a moist-adapted forest
dwelling group, but they can be found in habitats ranging
from montane herbland and open woodland to closed forest.
Most species are ground dwellers that occupy burrow retreats
in sheltered microhabitats—under rocks and logs, inside
rotting logs and stumps, and on stable, vegetated or mossy
soil ha nks (Fig. 1C). Ground burrowers often make use of
natural soil crevices such as rotted root channels. At least
two species (H. cerberea and H. formidabilis [Rainbow,
1914]) are associated with standing trees, their retreats
occupying trunk holes resulting from branch fractures or
borer damage, as well as the rotted internal pipes of older
trees (Fig. IE). Most atracine spider retreats are characterized

The original caption has been corrected by the author on this replacement page. The original
PDF of the entire work (dated 25 November 2010) has been replaced by the present PDF
(dated 7 August 2012) which has the replacement page inserted [Editor, 7 August 2012].

Gray: Australian funnel-web spiders

287

Figure 1. (A)Atrax robustus, female; (B) A. robustus, male; (C) Hadronyche macquariensis n.sp., burrow with two entrances on soil bank;
(D) H. versuta, female; (E) H. cerberea, single entrance retreat in rot-hole on tree trunk ( Casuarina sp.); (F) H. formidabilis, mating
(photo: P. Walker).

by the presence of silk trip-lines radiating out from a silk
entrance sheet (Fig. IE). The entrance tunnel (typically one
to three) usually has a crescentic opening with the surface
part of the silk lumen more or less collapsed when not in
use—perhaps as a predator deterrent. A few, more specialized
atracine species have burrow entrances opening within the
litter layer, and they lack any trip-lines. These species include
Illawarra wisharti n.sp., and the “adelaidensis group”
species of Hadronyche —the latter species are unique in
having an internal burrow chamber with a trapdoor (Main,

1967; Gray, 1984). Biological data on atracine spiders are
mostly anecdotal (e.g., McKeown, 1963) or associated with
field collecting observations (Main, 1976; Gray, 1986),
apart from a few studies by Levitt (1961), Bradley (1993)
and Wishart (1993). Of particular interest are atracine
eco-evolutionary studies by Cooley (1989) on diet related
character displacement in sympatric species, Woodman
et al. (2006) on climatic impacts on saproxylic habitats
and demography of associated atracines, and a genetic
phylogeographic study by Beavis & Rowell (2006).

Gray: Australian funnel-web spiders

287

Figure 1. (A) Atrax robustus, female; (B) Hadronyche versuta, female; (C) A. robustus, male; (D) H. cerberea, single entrance retreat in
rot-hole on tree trunk ( Casuarina sp.); (E) H. macquariensis n.sp., burrow with two entrances on soil bank; (F) H. formidabilis, mating
(photo: P. Walker).

by the presence of silk trip-lines radiating out from a silk
entrance sheet (Fig. IE). The entrance tunnel (typically one
to three) usually has a crescentic opening with the surface
part of the silk lumen more or less collapsed when not in
use—perhaps as a predator deterrent. A few, more specialized
atracine species have burrow entrances opening within the
litter layer, and they lack any trip-lines. These species include
Illawarra wisharti n.sp., and the “adelaidensis group”
species of Hadronyche —the latter species are unique in
having an internal burrow chamber with a trapdoor (Main,

1967; Gray, 1984). Biological data on atracine spiders are
mostly anecdotal (e.g., McKeown, 1963) or associated with
field collecting observations (Main, 1976; Gray, 1986),
apart from a few studies by Levitt (1961), Bradley (1993)
and Wishart (1993). Of particular interest are atracine
eco-evolutionary studies by Cooley (1989) on diet related
character displacement in sympatric species, Woodman
et al. (2006) on climatic impacts on saproxylic habitats
and demography of associated atracines, and a genetic
phylogeographic study by Beavis & Rowell (2006).

288

Records of the Australian Museum (2010) Vol. 62

Table 1. Atracinae—list of valid taxa, synonyms and transfers.

valid species—current placement

(transfers from At rax to Hadronyche by Gray, 1988)

original name, synonyms and transfers

Hadronyche cerherea L. Koch, 1873
Atrax robustus O.P.-Cambridge, 1877

Hadronyche modesta (Simon, 1891)
Hadronyche meridiana Hogg, 1902
Hadronyche versuta (Rainbow, 1914)

Hadronyche formidabilis (Rainbow, 1914)
Hadronyche valida (Rainbow & Pulleine, 1918)

Hadronyche pulvinator (Hickman, 1927)
Hadronyche venenata (Hickman, 1927)
Hadronyche infensa (Hickman, 1964)
Hadronyche adelaidensis (Gray, 1984)
Hadronyche flindersi (Gray, 1984)
Hadronyche eyrei (Gray, 1984)
Hadronyche anzses Raven, 2000

Hadronyche cerberea L. Koch, 1873

Atrax robustus O.P.-Cambridge, 1877 Euctimena tibialis Rainbow,
1914 (synonymy with A. robustus by Musgrave, 1927).
Poikilomorpha montana Rainbow, 1914 (transfer to Atrax by
Main, 1985 after Gray, 1978. Synonymy with A. robustus by
Gray, 1988).

Atrax modesta Simon, 1891
Hadronyche meridiana Hogg, 1902

Atrax versutus Rainbow, 1914 Pseudatrax moreaui Rainbow, 1914
(transfer to Atrax by Main, 1985 after Gray, 1978. Synonymy
with H. versuta by Gray, 1988). Aname bicolor Rainbow, 1914
(transfer to Atrax by Rainbow & Pulleine, 1918. Synonymy with
H. versuta by Gray, 1988).

Atrax formidabilis Rainbow, 1914

Atrax validus Rainbow & Pulleine, 1918. Transfer to Hadronyche by
Gray, 1988). Anepsiada ventricosa Rainbow & Pulleine, 1918.
Synonymized with A. validus (Gray, in Main, 1985).

Atrax pulvinator Hickman, 1927
Atrax venenatus Hickman, 1927
Atrax infensus Hickman, 1964
Atrax adelaidensis Gray, 1984
Atrax flindersi Gray, 1984
Atrax eyrei Gray, 1984
Hadronyche anzses Raven, 2000

nomen dubia

Styphlopis insularis Rainbow, 1913. Solomon Islands.

Hadronyche hirsuta Rainbow, 1920. Papua [= Papua New Guinea]. (Listed with comments in Main, 1985).

Taxonomic history

Atracine spiders were first described by L. Koch (1873)
who erected the genus Hadronyche for H. cerberea L.
Koch, 1873 from Sydney, New South Wales. Four years
later O.P.-Cambridge erected a second genus, Atrax, for A.
robustus O.P.-Cambridge, 1877 (locality “Australia”). Both
species were described from females. The female syntypes
of H. cerberea were destroyed during World War II (Main,
1985). The resultant uncertain identity of this species was
discussed by Gray (1981, 1986) and H. cerberea was fixed
as the southern tree funnel-web spider, a species common
in the Sydney region.

By 1920,14 species had been described, some more than
once, in no less than eight genera. Of these taxa, only seven
species and two genera are now recognized as valid (Table
1). Most species described during that period were based
only upon females or, even more uninformatively, juveniles,
i.e. generally character poor and taxonomically difficult
specimens—and several had suspect locality data. The only
species described from male spiders were H. meridiana
Hogg, 1902 and Euctimena tibialis Rainbow, 1914—the
latter spider proved to be a male of Atrax robustus (Musgrave,
1927). Rainbow (1914) also described the female of Atrax
robustus as Poikilomorpha montana Rainbow (synonymized
by Gray, 1988—Table 1).

After 1920, the older generic name, Hadronyche, largely
fell into disuse (although still listed in catalogues) and
Atracinae were associated almost exclusively with the
genus Atrax (Table 1). This process was facilitated both by

uncertainty about the taxonomic identity of the type species,
H. cerberea, and the increasing medical notoriety of A.
robustus. Hadronyche was listed in synonymy with Atrax
by Raven (1980), from which it was implicitly removed by
Main (1985) when she listed 11 species in Atrax and three in
Hadronyche. Most of the species listed under Atrax in Main
(1985) were subsequently transferred to Hadronyche (Gray,
1988). All atracine species now regarded as valid have been
described from Australia. These species and their synonyms
are listed in Table 1. Collection data for two species from
Melanesia are regarded as unreliable (Main 1982, 1985).

The relationships of the Atracinae remain uncertain.
Simon (1892) placed Atrax and Hadronyche within his
Macrotheleae in the family Dipluridae. His key grouped
them with the genera Macrothele and Porrhothele Simon,
1892 by the presence of labial cuspules. In 1901, Hogg
erected the group Atraceae to distinguish these genera from
the remaining members of Simon’s Macrotheleae. This
distinction was based upon the sternal sigilla (large size and
marginal position), presence of three rows of cheliceral teeth
and the relatively short apical segment of the PLS.

Raven (1980, 1985) removed Simon’s Hexatheleae from
the Dipluridae, elevating it to family status. Within his
Hexathelidae he included part of Simon’s Macrotheleae,
i.e., those quadritheline genera possessing labial cuspules
(Atrax, Porrhothele and Macrothele Ausserer, 1871), making
the possession of numerous labial cuspules the unifying
character for the Hexathelidae. Gray (1988) re-instated the
Atracinae, comprising Hadronyche and Atrax (Atraceae of
Hogg, 1901), additionally distinguishing these taxa from

Gray: Australian funnel-web spiders

289

Figure 2. Distribution of atracine spiders in eastern Australia arrow
points to locality of Hadronyche anzses ).

Porrhothele and Macrothele by the presence of a wider
embolus, strong retromarginal cheliceral teeth, maxillary
lobe and procurved fovea.

Raven (1985) acknowledged difficulties with the place¬
ment of At rax in the Hexathelidae and noted several characters
that suggested possible (but less parsimonious) affinity with
the Cyrtaucheniidae. Goloboff (1993), in his morphologically
based reanalysis of mygalomorph relationships, found weak
support for a monophyletic Hexathelidae. A molecular study
of mygalomorph relationships (Hedin & Bond, 2006) did not
find support for the monophyly of the Hexathelidae— Atrax
and Hadronyche were always recovered as sister taxa but
were not associated with other hexathelid taxa. However, as
similar anomalies were recorded for several well established
families, interpretation of these results is problematic.

Material and methods

Observations, measurements and illustrations were made
using a Wild M5 stereomicroscope with measuring graticle
and drawing attachment. All measurements are given in
millimetres. Appendage illustrations are of the left hand side
unless otherwise stated. Genitalic preparations were made
using lactic acid or 4% KOH solution. SEM micrographs
were made from ethanol dehydrated specimens, air dried
and gold coated.

Notes on descriptions

Keys are given to atracine genera and species of Hadronyche
and Atrax', Illawarra is monotypic. Identification to species
requires male specimens.

In the species descriptions given below, measurement/
count data given are for the holotype or specified paratype
specimens. Additional range, mean and ratio data (which
includes the paratype specimens) are given in tables
accompanying the descriptions (males) or in Table 34
(females). Qualitative data given in descriptions refers to both
holotype and paratype specimens. On a few occasions when
type material was not available for re-description, non-type
material is described and illustrated. Types, non-type figured
or described material and Other material examined are listed
with the descriptions.

Undescribed species dealt with in Gray (1986, 1987,
1988) were then identified by number codes. These codes
are given in Appendix 2 with the corresponding species
names allocated here.

Measurements and counts. Body, leg and palpal measure¬
ment positions are illustrated in Appendix 1. Measurements,
counts and ratios given in keys and diagnoses refer to male
spiders unless specifically noted as data from females.

Palpal organ (bulb). “Tegular area” refers to tegulum/
subtegulum region of bulb. Palpal organ measurements, viz.,
bulb length; bulb width; embolus length and mid-width,
plus other bulbal characters, viz. “basal embolus offset from
tegulum” and “twisting of distal embolus”, are illustrated in
Appendix 1. (Note: palpal orientation should be as shown
in Appendix 1 “A”)

Colour. Atracine spiders are typically glossy black on
the carapace and mat black, often with a plum tinge, on
the dorsolateral abdomen. They rarely possess distinctive
colour patterning that can be used to characterize species,
and melanic pigmentation intensity and distribution may
vary intraspecifically (pigment intensity also fades with
preservation). In species descriptions any consistent
variations in colour pattern are noted.

Leg spination. In descriptions, the total spine count per
leg segment is given first. This count includes the ventral/
retroventral spines (usually the most numerous spine
category on the tibia, metatarsus and tarsus), plus any dorsal,
prolateral and retrolateral spines present. Counts for the
dorsal (d), prolateral (p) and retrolateral (r) spines are given in
brackets after the total count. Total spine counts for selected
leg segments are given in tables with the descriptions. Patellal
counts include total ventral and prolateral dorsal spines, with
the latter also numbered in brackets (pd) if present. Tibial and
metatarsal counts refer to total ventral or retrolateral ventral
(retroventral) spines, including apical ventrolateral spines
and prolateral spines; the latter are also given separately in
brackets when present.

290

Records of the Australian Museum (2010) Vol. 62

Abbreviations

Morphology. BulbL—palpal bulb length; BulbW—palpal
bulb width; CL—carapace length; CW—carapace width;
CFW—carapace anterior width; CH—carapace height;
ChGL—cheliceral groove length; ChGW—cheliceral groove
width; ChGCT—cheliceral groove central row teeth number;
CUSP—labium cuspule number; EmbL—embolus length;
EmbmidW—width of embolus at middle; FelS—femur
I spine number; LL—labium length; LW—labium width;
MtlS—metatarsus I spine number; Mt2S—metatarsus II
spine number; PalS—patella I spine number; PalpTibL—
male palpal tibia length; PalpTibW—male palpal tibia width;
PalpFemS—male palpal femur spine number; PalpPatS—
male palpal patella spine number; PalpTibS—male palpal
tibia spine number; PLSAPL—posterior lateral spinneret
apical segment length; PLSAPW—posterior lateral spinneret
apical segment width; SL—sternum length; SW—sternum
width; TalS—tarsus I spine number; TilS—tibia I spine
number; Ti2S—tibia II spine number; STC—superior tarsal
claws; STC2teeth—number of teeth on tarsus II superior
claws; Pa3proS—patella III prolateral dorsal spine number.

Australian political boundaries (states). Qld—Queensland;
NSW—New South Wales; VIC—Victoria; SA, South
Australia.

Repository institutions. AMS—Australian Museum,
Sydney; MNHN—Museum national d’Histoire naturelle,
Paris; NHM—Natural History Museum, London; QMB—
Queensland Museum, Brisbane; QVM—Queen Victoria
Museum, Launceston; MV—Museum Victoria, Melbourne;
SAM—South Australian Museum, Adelaide; TM—
Tasmanian Museum and Art Gallery, Hobart; ANIC—
Australian National Insect Collection.

Specimen registration codes. AMS KS—prefix of Australian
Museum, Sydney register numbers; S—prefix of Queensland
Museum, Brisbane, register numbers.

Systematics

Family Hexathelidae Simon

Subfamily Atracinae Hogg

Macrotheleae (part): Simon, 1892: 182.

Atraceae Hogg, 1901: 250; Simon, 1903:961,968; Rainbow,
1913:4, 1914: 252.

Pseudatraceae Rainbow, 1914: 258, 259.

Poikilomorphiae Rainbow, 1914: 260, 162.

Anaepsiadiae Rainbow & Pulleine, 1918: 166; Roewer,
1942: 208.

Atraxeae Roewer, 1942: 207.

Pseudoatraxeae Roewer, 1942: 208.

Atraxini Bucherl, 1971:211.

Macrothelinae (part): Musgrave, 1948:77; Gray, 1978: 125;

Raven, 1980: 254, 1985: 71, 441.

Atracinae.-Gray, 1988: 115.

Diagnosis. Atracinae differs from Hexathelinae in having
only 2 pairs of spinnerets and from Macrothelinae in having
a relatively broad embolic shaft (not spiniform), cheliceral

retromargin with row of large teeth, posterior sternal sigilla
large, maxillae with coniform anterior lobe, PLS relatively
short, and fovea a transverse groove.

Description. Basic colour pattern: carapace, chelicerae and
legs dark brown to black, sometimes a lighter reddish brown;
carapace with a polished appearance; Abdomen brown to
black often with a maroon tint; anterodorsal abdomen with a
pair of unpigmented sigilla (often subdivided) which flank a
weakly pigmented mid-dorsal patch of cuticle. Posterior to the
sigilla are 3-5 pairs of more or less distinct, narrow chevron
markings made up of small unpigmented sigilla-like spots.
Abdomen with thin cover of dark hairs and bristles. Carapace
almost glabrous, with strial, midline and marginal setae.
Carapace with caput arched and weakly to strongly raised
(Fig. 1A,D). Thoracic margins grooved. Fovea a transverse
groove, straight to strongly procurved. Eye tubercle low
or absent. Cheliceral groove with promarginal (few in H.
anzses ) and retromarginal tooth rows plus smaller teeth in a
central irregular row(s) of variable length of (Figs. 3D, 5F);
paturon robust, more so in females, with a distally widening
dorsal band of strong dark hairs and bristles and a medial
dorsofrontal swelling developed into boss-like protuberance
in many Hadronyche spp. No intercheliceral tumnescence.
Fangs with ventrolateral longitudinal keels. Maxillae longer
than wide with a strong coniform lobe anteromedially (Figs.
5A, 2IE); serrula usually present (absent in adelaidensis
group), serrula teeth variably clustered on apicodorsal
face of maxillary lobe (Fig. 3H); numerous cuspules along
medioventral surface of maxilla, dense basally, diminishing
toward lobe (Fig. 3H). Labium large, more or less rectangular
and rounded anteriorly, varying from clearly wider than long
(i.e., relatively short) (Fig. 5A) to about as long as wide
(i.e., relatively long) (Fig. 21E). Labial cuspules usually
numerous, occasionally relatively few (c. 50-500); cuspules
clustered over anteroventral half to two-thirds of labium
(Fig. 3E). Labiostemal sigilla typically a wide transverse
groove, rarely reduced to a pair of small, lateral sigilla (. H.
mascordi). Sternum ovoid, wide to moderately narrow, six
submarginal to central sigilla, anterior pair small, circular,
posterior pair large, ovoid. Cuticle with scaliform patterning.
Legs moderately to weakly hirsute with longitudinal glabrous
strips on dorsal and lateral surfaces. Leg tarsi ventrally
spinose, spines usually confined in two lateral rows (Fig.
3A), sometimes with isolated central spines (1-3), rarely
with a full central spine row (Fig. 3B). Three tarsal claws,
each STC with a sigmoid row of 7-15 strong teeth, ITC with
few slender to short teeth; numerous long fimbriated hairs
arise at base of claws (Fig. 3F). Tarsal scopulae absent in
females, weak to moderate in males, sometimes extending
onto distal metatarsus; scopula hairs short, thick, tapering
(Fig. 3C). Metatarsal preening combs absent. Male tibiae I
and metatarsi I often strongly spinose with spines placed on
retroventral and ventral surfaces respectively. Male tibiae II
either unmodified with clustered or scattered ventral spines,
or weakly sinuous (distoventrally concave) with a ventral
spined apophysis or apophyseal swelling. Male metatarsus
II either unmodified, or sinuous (proximoventrally concave)
with small mid-ventral apophysis. Trichobothria filiform
with weakly collariform bothria; in zig-zag row on tarsus,
linear dorsal row on metatarsus, double row on tibia. Tarsal
organ dome-like, with a few weak concentric ridges. (Fig.
II). Male palpal patella and tibia more or less swollen (least

Gray: Australian funnel-web spiders

291

Figure 3. (A) Hadronyche infensa, ventral tarsus II, scopula and lateral spine rows; ( B ) Illawarra
wisharti n.gen., n.sp., ventral tarsus II with weak divided scopula and medial and lateral spine
rows; (C) H. nimoola n.sp., scopula setae; (D) H. versuta, fang and cheliceral groove teeth; (E)
H. infensa, labium and cuspules; (F) H. infensa, tarsus I, claws, teeth and fimbriated hairs; G, H.
annachristiae n.sp., male leg tarsi: I, II unmodified (left), III, IV enlarged (right); H, H.formidabilis,
maxillary serrula; I, H. tambo n.sp., tarsus II, distal trichobothrium and tarsal organ.

in some Atrax spp.), patella about as wide or wider than the
femur; (Figs. 10A, 77A,100A). Cymbium short, equally
bilobed anteriorly, without spines. Bulb with ovoid-pyriform
tegular area, a deep, longitudinal groove separating tegulum
and subtegulum; groove occasionally broadly open, exposing
middle haematodocha (Fig. 102D). Embolus a relatively
wide, flattened rod, variable in taper, length and curvature,
more or less twisted distally; apical embolus with a broad or

narrow ejaculatory groove with a flange-like lower margin
(Figs. 6B, 29B, 59B). Female genitalia with a pair of simple,
unilobate spermathecae, often slightly constricted subapically
(Figs. 5G, 31H). Four spinnerets. PMS separated by width of
basal segment; PLS with long or short digitiform terminal
segment, spigots present on all segments; spigot shafts with
scaliform patterning. Mating involves leg II embrace—male
tibia I I/metatarsus II clasping base of female leg II.

292

Records of the Australian Museum (2010) Vol. 62

Included genera. Hadronyche L. Koch, 1873; Atrax O.P.-
Cambridge, 1877; Illawarra new genus

Distribution. Eastern Australia, including Tasmania (Fig. 2).

Melanesian species of uncertain status. Two atracine
species have been described from Melanesia— Styphlopis
insularis Rainbow 1913 from the Solomon Islands (AMS
KS993); and Hadronyche hirsuta Rainbow 1920 from
Papua (Chevert Expedition) (KS992). These species
are regarded here as nomen dubia (Table 1). Both were
described from large female specimens that are attributable
to the genus Hadronyche. Female atracines mostly lack
defining specific characters, but the presence of a relatively
long labium suggests that both species could be placed in
either the infensa or cerberea species groups. At present,
these species groups are known only from southeastern
Australia. Although atracine spiders were recently found
in northeastern Australia (Mosman region), they belong
to a different, highly distinctive species group. Neither
melanesian species has been validated by subsequent
collecting and no other atracine species have been recorded
outside Australia. Consequently, both species are regarded
here as mislocated, possibly Australian taxa of uncertain
species status. Main (1982, 1985) had reached the same
conclusions regarding H. hirsuta.

Comments on morphology

Body size (carapace length ). Some 14 species of Atracinae
have a relatively small body size (male CL 4.5-7.5). These
include the lamingtonensis and adelaidensis group species,
plus H. jensenae, H. mascordi, H. meridiana, H. modesta, H.
monaro and H. nimoola (currently members of the cerberea
group). Most of the remaining species fit, with some overlap,
into the medium (CL 7.5-10.0) or large (CL 10-12) male
size classes.

Carapace height and cheliceral robustness. The arched
cephalic area or caput is weakly to strongly raised in
Atracinae. It is lowest in Atrax and Illawarra spp. (male
CH/CW 0.35-0.37) (Figs. 4B, 16B). It is more strongly
raised in Hadronyche spp. (male CH/CW 0.40-0.53). (Fig.
30A). The higher caput in Hadronyche spp. is associated
with broadening of the carapace and a deeper, more robust
cheliceral paturon, both in males and especially females (cf.
Fig. 9B,F and Fig. 21B,C).

Cheliceral groove and teeth. The cheliceral groove is
usually widest distally and clearly tapering proximally,
but the margins of narrower grooves may be subparallel
or only weakly tapered. The groove is narrow in Atrax
and Illawarra (Figs. 4C, 16C), and in the lamingtonensis
group of Hadronyche. In most other Hadronyche spp. the
groove is relatively wider, particularly in females (Fig. 3D).
Three tooth rows are present—two more or less full (long)
rows of large teeth on the prolateral and retrolateral groove
margins, plus a mid-groove, central row of much smaller
teeth. This central row is always short and basal in Atrax,
Illawarra (Figs. 4C, 16C) and the lamingtonensis group spp.

of Hadronyche. In most Hadronyche spp. the central teeth
typically occupy the full groove length, with one (males) to
several (females) irregular rows of small teeth (Figs. 20G,
2ID). In H. meridiana a few mid row or distal teeth may
be missing, making some individuals difficult to key out.
Hadronyche anzses is unusual in having the prolateral tooth
row reduced to a few basal teeth (Raven, 2000).

Labium and cuspules. The labium is typically wider
than long, but varies considerably in relative length. It is
consistently short and wide in both Atrax and Illawarra (Figs.
5 A, 16D) and in the lamingtonensis and adelaidensis species
groups of Hadronyche (0.700.80). By contrast, the labium is
relatively longer (occasionally about as long as wide) in the
cerberea and infensa species groups of Hadronyche (Figs.
27A, 72A), except in four cerberea group species (see group
diagnosis) including H. modesta (Fig. 57E). A short labium
is also seen in H. mascordi (Fig. 50A). In this species the
labiostemal sigilla is uniquely divided into 2 lateral sigilla
(usually an entire groove-like sigilla), with fusion of the
medial labium and sternum between them—the short labium
is regarded as a probable apomorphic condition related to
these modifications.

The short, coniform to bluntly pointed labial cuspules
(similar in males and females) are mostly clustered on the
anterior part of the ventral labium (Fig. 3E). They are usually
numerous (200-500), but may be relatively few in some
species (e.g., 50-125 in most lamingtonensis group spp).
Cuspule numbers in males are categorized as follows: high,
>250; moderate, 125-250; low, <125.

Maxillary serrula. A serrula is usually present on the
maxillary apical lobe (absent in adelaidensis group spp.).
The serrula field consists of many minute pointed teeth
spreading from the anterior lobe surface onto the dorsomedial
face (Fig. 3H). There is considerable variation in serrula size
and number of teeth.

Anterolateral sternal bristles. This is a group of bristles,
stronger and thicker than those generally found on the
sternum, placed at the sternal angles on each side of the
labium (Fig. 20H). They are best developed in H. cerberea
(both sexes) and variably present in a few related species
(e.g., H. emmalizae ), but absent in H. versuta.

Coxal setae. Setae on the anterobasal surface of coxae 1, 2
are usually hair-like. In a few species some of these may be
shortened and basally thickened. This is best developed in
H. cerberea, resulting in thom-like setae (Fig. 21H) in both
sexes (absent in H. versuta).

Posterior lateral spinnerets. The PLS are much shorter than
in Macrothele and Porrhothele. However, PLS length still
varies considerably in the Atracine, especially the apical
segments—from relatively long digitiform (c. length 4X
mid-width) to short, wide digitiform (c. length 1.5X mid¬
width). The longest spinnerets are found in Atrax spp. (Fig.
4G), while shorter states are found in Illawarra (Fig. 17F)
and in Hadronyche, notably in the lamingtonensis species
group (Fig. 97A). Many Hadronyche spp. have spinnerets of
intermediate length but a few species (e.g., H. formidabilis)
have PLS almost as long as Atrax spp.

Gray: Australian funnel-web spiders

293

Male tarsi III, IV shape. Typically, Atracinae have more or less
“cylindrical” tarsi. Occasionally, tarsi III and IV are swollen
and lengthened (tarsus IV is usually most affected) compared
with tarsi I and II, giving them a “boat-shaped” appearance
(Fig. 3G). This character is associated with several species in
the lamingtonensis species group of Hadronyche, including
the north Queensland species, H. anzses. Comparable swelling
of tarsi III, IV is seen in H. lynahrae, and weaker swelling of
these tarsi has been observed in H. flindersi and H. jensenae,
and of tarsus IV in H. mascordi and H. monaro all species
outside the lamingtonensis group.

Tarsal scopulae. Scopulae are present only in males.
Scopula setae are numerous to relatively few in number
(Fig. 3A,B). The setae are short, thick and curved (rather
vermiform), and set in ovoid bases (Fig. 3C). The shafts
have finely circumferentially ribbed or annulate patterning
while the tapering distal region may be more or less flattened
apically. The structure of these scopula setae is quite
different from that seen in the dense scopulae of barychelids
and theraphosids, but I am unfamiliar with scopula setal
morphology in other mygalomorph groups.

Male legs I, II spination.

Femur. The presence of mid-dorsal spines (one to row
of several), with or without some distad prolateral dorsal
spines, is characteristic of all Atrax and Illawarra spp. In
Hadronyche these spines are absent in both the infensa
and the lamingtonensis species groups. They are present in
cerberea group species, but a few species have individuals
that lack dorsal spines.

Patella. Both prolateral dorsal spines (sometimes with
additional prolateral spines) and distal ventral spines are
often present, sometimes absent.

Tibia. Tibia I spines may be numerous and distributed
ventrally and retrolaterally (retroventrally) (Fig. 22D,E),
or less numerous and more ventrally placed, with one to
several lateroventral apical spines; prolateral surface with
0-4 spines. Tibia II spines are mostly placed ventrally, either
scattered or more or less midventrally grouped, with 0-5
spines placed prolaterally and one to several lateroventral
apical spines. Both scattered and clustered spine patterns are
associated with species lacking tibial modifications. (Figs.
55H,I; 93H,I). Tibial apophyses and swellings are always
associated with clustered spines (Fig. 32H,I).

Metatarsus. Ventrally spinose, metatarsus I with spines
often denser proximomidventrally, sometimes associated
with proximal-midventral metatarsal thickening (Fig.
32F). Metatarsus II with ventral spines mostly placed
midventrally to distally (Fig. 55J,K), sometimes associated
with a midventral apophysis/swelling (Fig. 32J,K). Prolateral
surface with 0-1 spines.

Tarsus. Typically with two rows of small spines ventro-
laterally (Fig. 3A). An additional complete row of mid-
ventral spines characterizes males of the genus Illawarra
(Fig. 3B).

Male leg II apophyses. Apophyses, and less prominent

apophyseal swellings, are found on both the second tibia and
metatarsus of some atracine species. They are any ventral
cuticular projections or swellings, placed in the middle to
proximal part of the segment and clustered with spines.
These leg II structures are found in all Atrax spp. and several
species currently placed in the cerberea species group of
Hadronyche. All other atracines, comprising Illawarra and
most Hadronyche spp., lack such structures. Tibial apophyses
are best developed in Atrax spp., as a prominent co nif orm
apophysis with short, stubby spines (Figs. 4F; 6H,I). In
Hadronyche , a less specialized structure occurs in the form
of a broad, ovoid to rounded ventral swelling, placed mid¬
ventrally to proximally and clustered with strong, often
somewhat shortened spines. This type of apophysis is most
strongly developed in H. formidabilis (Fig. 32H,I). In other
species it is represented as a weaker apophysis or apophyseal
swelling (Figs. 22J, 361, 401). The latter term refers to a low
ventral tibial thickening upon which spines are clustered—
however tibial apophyses and apophyseal swellings tend to
intergrade into each other. Tibial apophyses and apophyseal
swellings are associated with a variably developed disto-
ventral concavity that adjoins a similar anteroventral
concavity on the metatarsus. Metatarsal apophyses are
strongly to weakly developed, spinose, and typically
associated with a variable sinuosity of the metatarsus, (Figs.
22K, 29K). However, in H. venenata the apophyseal swelling
is very weak and there is no metatarsal apophysis (Fig. 38E);
and in H. monaro the tibia is unmodified but the metatarsus
has a small apophysis (Fig. 47G).

Male palp (Figs. 6A,B,C; 43A,B,C; 65A,B,C) (Appendix
1). The palpal bulb is rather simple in structure, comprising
an ovoid-pyriform tegular area and a rod-like embolus. The
tegular area consists of an ovoid tegulum separated by a deep,
curved groove from the smaller subtegulum (Fig. 48B). The
tegulum and subtegulum are sometimes widely separated in
lamingtonensis group spp. (and often in Illawarra wisharti
also), exposing the membranous median haematodocha
spanning the widened space (Figs. 97H, 100B). The relatively
wide embolic shaft is quite different from the slender spiniform
embolus of macrotheline and hexatheline spiders. It is variably
basally offset from the tegulum and varies in width, length,
curvature, taper and degree of distad axial twisting. On the
distal part of the embolus the ejaculatory duct opens into a
broad or narrow distal groove, the lower margin of the groove
appearing as a more or less flange-like structure.

The male palpal patella and tibia are enlarged in many
atracine species. Raven (2000) suggested that the greater
width of the palpal patella compared to the femur represents
a synapomorphy for the genus Hadronyche —in most
other mygalomorph spiders (as in Atrax ) the palpal patella
may be as wide but not wider than the femur. A limited
survey confirms Raven’s observation in both smaller-sized
Hadronyche species— lamingtonensis and adelaidensis
group species and most smaller cerberea group species
(Figs. 64G, 100A)—and several larger species examined,
e.g., H. venenata (Fig. 40A). While this character state is
present in many Hadronyche species, it does not seem to be
universal (e.g., in H. levittgreggae the femur is as wide as
the patella). Interestingly, the wider patella state is present
in Illawarra wisharti. A fuller survey of atracine species is
needed to determine both the distribution of this character
and its status in Hadronyche.

294

Records of the Australian Museum (2010) Vol. 62

Keys to Atracinae

The information given in these keys relies upon characters
taken from male spiders—the keys require male specimens.
Measurements, counts and ratios given in these keys refer
to male spiders unless specifically noted as female data.
Leg and palp spine counts represent totals for the segment,
unless otherwise noted.

Species attributed to the diverse cerberea group show
intra-specific variation in dorsal femoral I, II spination
(spines present in most spp., but absent in some individuals
of a few spp.). This means that some species have two (part)

key outcomes. The use of ratio data in some parts of the key
make it less user friendly than one would like, especially for
inexperienced users.

No species key is given for the genus Illawarra
(monotypic). Males of the Tasmanian species, H. pulvinator
(Hickman, 1927) and the South Australian species, H. eyrei
(Gray, 1984), are unknown and these species are not included
in keys—their species diagnoses and apparently limited
distributions should assist with their identification.

Key to genera

1 Male tibia II with large, prominent, conical apophysis (Fig. 61).

Caput weakly raised (Figs. 4B, 5B). Central cheliceral tooth row

short, basal (Fig. 5F). Labium short (Fig. 4A). Atrax O.P.-Cambridge

-Male tibia II not with a prominent conical apophysis—tibia II

either unmodified or with bluntly rounded apophysis or apophyseal

swelling. Caput weakly to strongly raised. 2

2 Male tibia II unmodified (Fig. 16F). Caput weakly raised and
frontally narrow, cheliceral paturon relatively weak (Figs. 16A,B;
17B,D). Male tarsi I, II with a midventral spine row as well as two
lateral rows (Fig. 3B). Central cheliceral tooth row short, basal
(Fig. 16C). Labium short. PLS short (Figs. 16E, 17F) .

-Caput moderately to strongly raised and frontally broad,

cheliceral paturon robust (Figs. 20B,E; 21B,C). Male tarsi
I, II lacking a full midventral spine row. Male tibia II either
unmodified (Fig. 68F), or with rounded apophysis/apophyseal
swelling (Figs. 30C, 34G). Central cheliceral tooth row long
or short. Fabium long or short .

Illawarra n.gen.

Hadronyche F. Koch

Genus Atrax —Key to species

1 Palpal tibia with 4-11 spines, dorsal spines present. A. robustus

-Palpal tibia with few or no spines (0-4), dorsal spines absent . 2

2 Embolus long and slender, strongly curved; base strongly offset

from tegulum (Fig. 10B) .A. yorkmainorum

-Embolus relatively shorter, wider and more weakly curved (Fig.

14B) . A. Sutherlandi

Genus Hadronyche —Key to species

1 Farge spiders (male CF 10.0-12.0). Tibia II with a large,
protuberant, rounded apophysis (Fig. 30C). PFS relatively long

(PFSASF > 3xW) (Figs. IF, 30D) . H. formidable

-- Spiders usually smaller. Tibia II apophysis less prominent or

absent. PFS often shorter. 2

2 Tibia I and metatarsus I incrassate (Fig. 106F) . H. adelaidensis

-Tibia I and metatarsus I not incrassate. 3

3 Palpal patella with many spines (12-16) (Fig. 103F). Feg II

unmodified . H.flindersi

-- Palpal patella spines absent or fewer (0-9). Feg II modified or

unmodified . 4

4 Fabium and sternum fused medially, labiostemal sigilla divided

(Fig. 50A). H. mascordi

-Fabium and sternum not fused, labiostemal sigilla entire . 5

Gray: Australian funnel-web spiders

295

5 Caput high and wide frontally, almost bulbous, rising relatively

steeply from fovea (CH > 0.5xCW) (Figs. 53A,C; 54B). Meta¬
tarsus II unmodified (Fig. 53E). H. nimoola

-Not in above combination. 6

6 Chelicerae with central tooth row short, basal (Fig. 82D). Labium
almost as long as wide (LL/LW 0.96) (Fig. 82A)

. H. kaputarensis

-Not in above combination. 7

7 Embolus short, with distal flanged part set at distinct angle to

shaft (Fig. 65B,C) . H.jensenae

-Distal embolus not as above . 8

8 Chelicerae with central tooth row short, basal. (Fig. 97E). Palpal
middle haematodocha widely exposed (Figs. 97H, 100B). Caput
strongly raised (Fig. 97B). PMS apical segment short (Fig. 97A)

(.lamingtonensis species group). 9

-Chelicerae with central tooth row typically long (Fig. 20G), rarely

short. Palpal middle haematodocha not or rarely exposed (Fig. 48B).

Caput height and PMS length variable . 13

9 Cuspules on labium relatively numerous (200-225). Cheliceral

margins diverge distally (Fig. 94C) . H. raveni

-Cuspules on labium relatively sparse (40-100). Cheliceral margins

subparallel (Fig. 97E) . 10

10 Male tarsi III and IV unmodified . H. lamingtonensis

-Male tarsi III and IV swollen (“boat-shaped”) than, tarsi I and II

(Fig. 3G). 11

11 Cheliceral promargin with few teeth (c. 3), in short basal row.

Embolus straight . H. anzses

-Cheliceral promargin with several teeth (c. 6-12) in longer row

(Figs. 99D, 102C). Embolus strongly curved (Figs. 100A,B; 102D). 12

12 Male tarsus I strongly spinose (24-43). H. annachristiae

-Male tarsus I weakly spinose (5-7) . H. monteithi

13 Labium relatively short (LL/LW 0.70-0.80) (Figs. 57E, 60D). Tibia

II and metatarsus II with apophyseal swellings . 14

-Labium relatively long (LL/LW 0.85-1.05) (Figs. 23 A, 68A). Tibia

II and metatarsus II apophyses/apophyseal swellings present or

absent . 15

14 Embolus short (BulbW/EmbL 0.80), not apically twisted (Fig.

59B,C). H. modesta

-Embolus longer (BulbW/EmbL 0.60), twisted apically (Fig. 62B,C)

. H. meridiana

15 Dorsal femur I and/or II with spines. 16

-Dorsal femur I, II with no spines . 23

16 Tibia II with weak, proximad, spinose apophyseal swelling (Figs.

38F; 40H,I); metatarsus II lacks apophyseal swelling (Fig. 40K).

Carapace relatively low (CH/CL 0.40) (Fig. 38B) . H. venenata (Tasmania)

-Not as above. 17

17 Tibia II unmodified, with few spines (7-11); metatarsus II with
small apophyseal swelling (Figs. 47G; 48H,I). Carapace relatively

high (CH/CL 0.49) (Fig. 47B) . H. monaro

-Not as above. 18

296

Records of the Australian Museum (2010) Vol. 62

18

19

20

21

22

23

24

25

26

27

28

29

30

31

Tibia II and metatarsus II with apophyses/apophyseal swellings.

Tibia II and metatarsus II lacking apophyses/apophyseal swellings.

. 19

. 22

Embolus with strongly curved shaft (Fig. 29B).

Embolus shaft weakly curved-straight (Fig. 25B) .

. H. emmalizae

. 20

Male (and female) coxae I, II with thorn-like setae anterobasally
(Fig. 21H), and anterolateral sternal angles with tuft of bristle-like
hair (Fig. 20C,H).

Not as above.

. H. cerberea

. 21

Tibia II with spines distributed over apophysis and distoventral
tibia (Figs. 23C; 25H,I). PLS relatively short (PLSAPW/ L 0.49)

(Fig. 23E) . H. versuta (part)

Tibia II with spines grouped on apophyseal swelling, few or
none on distoventral tibia (Figs. 34G; 36H,I). PLS relatively long

(PLSAPW/ L 0.33) (Pig. 34F) . H. alpina (part)

Tibia II and metatarsus II ventrally concave (bowed) (Pig. 431,K).

Embolus relatively wide (EmbmidW/ L 0.12) (Pig. 43B). H. marracoonda

Tibia II and metatarsus II more or less straight, not bowed (Fig. 461,

K). Embolus narrower (EmbmidW/ L 0.08) (Fig. 46B) . H. tambo (part)

Tibia II and metatarsus II with apophyses/apophyseal swellings. 24

Tibia II and metatarsus II lacking apophyses/apophyseal swellings. 25

Tibia II with spines distributed over apophysis and distoventral
tibia (Figs. 23C; 25H,I). PLS relatively short (PLSAPW/ L 0.49)

(Fig. 23E) . H. versuta (part)

Tibia II with spines grouped on apophyseal swelling, few or
none distributed onto distoventral tibia (Figs. 34G; 36H,I). PLS

longer (PLSAPW/ L 0.33) (Fig. 34F) . H. alpina (part)

Embolus relatively broad (EmbmidW/L 0.12-0.16); moderately to

strongly twisted distally (Fig. 86B,C). 26

Embolus relatively narrow (EmbmidW/L 0.06-0.09); weakly to

moderately twisted distally (Fig. 74B,C). 29

Embolus base strongly offset from tegulum (Figs. 70B, 8IB) . 27

Embolus base weakly offset from tegulum (Figs. 86B, 89B). 28

Embolus shaft strongly curved proximally, distal third strongly

twisted (Fig. 81B,C). Tarsi III, IV swollen, “boat-shaped”. H. lynabrae

Embolus shaft curvature and distal twisting less strongly developed

(Fig. 70B,C). Tarsi III, IV unmodified . H. infensa

Tibia II sinuous ventrally, with ventral spines clustered in proximal

half, (Fig. 89H,I). Tarsus I spines 8-38 . H. walkeri

Tibia II not sinuous, ventral spines more scattered (Fig. 86H,I).

Tarsus I spines often more numerous (30-63). H. macquariensis

Embolus slender, elongate (EmbL 3.53-4.15) (Fig. 93B). H. levittgreggae

Embolus not as long (EmbL 2.18-3.06) . 30

Labial cuspule number moderate (149-184). Bulb shorter than

palpal tibia (BulbL/TibL 0.77) (Fig. 46A). H. tambo (part) (Gippsland, Victoria)

Labial cuspules numerous (246-393) Bulb almost as long as palpal

tibia (BulbL/TibL 0.86-0.89) (Fig. 77A) . 31

Leg spines relatively few: tarsus I with 4-10, metatarsus I with 10-

16, tibia II with 3-7 spines. H. or ana

Leg spines more numerous: tarsus I with 23-46, metatarsus I with

31—56, tibia II with 13-25 spines. H. valida

Gray: Australian funnel-web spiders

297

Generic and Species Descriptions

Atrax O.P.-Cambridge

Atrax O.P.-Cambridge 1877: 26. Type species Atrax robustus
O.P.-Cambridge 1877 by monotypy. Simon, 1891: 302;
1892: 175, 182, 186; 1903: 967-969. Hogg, 1901: 272;
Rainbow, 1911: 121; 1913: 4; 1914: 252. Rainbow &
Pulleine, 1918: 165. Hickman, 1927: 63; 1964: 107.
Musgrave, 1927: 33; 1948: 77. Roewer, 1942: 207.
Bonnet, 1955: 772. Main, 1976: 70; 1981: 839; 1985:

40. Gray, 1978: 121,125. Raven, 1980: 255; 1985: 55,71;
Brignoli, 1983: 122. Gray, 1984: 441; Gray, 1988: 114.
Goloboff, 1993:6. Hedin & Bond, 2006: 467. Platnick,
2010 .

Euctimena- Rainbow 1914: 248. Type species Euctimena
tibialis Rainbow 1914 by monotypy. First synonymized
by Musgrave, 1927: 33. Hickman, 1964: 107. Gray, 1978:
125. Raven, 1980: 255. Main, 1985:40. Gray, 1988: 114.
Poikilomorpha. -Rainbow 1914: 264. Type species Poikilo-
morpha montana Rainbow 1914 by monotypy. First
synonymized by Gray, 1978: 125. Raven, 1980: 255.
Main, 1985: 40. Gray, 1988: 114.

Diagnosis. Differs from other atracine genera by presence
of a large co nif orm apophysis on male tibia II (Fig. 6H,I);
and from Hadronyche by lower caput height.

Figure 4. Atrax robustus, male: (A) sternum, labium and maxilla; (B)
cephalothorax and chelicerae, lateral; (C) cheliceral groove teeth;
(D) cephalothorax and chelicerae, dorsal; (E) palp, prolateral; (E)
leg II, prolateral; (G) spinnerets. Scale lines 1 mm.

Description. With characters of Atracinae. Medium to large
sized Atracinae. Carapace longer than wide, caput only
weakly raised (CH/CW 0.35-0.36) and narrow frontally
(CFW/CL 0.55-0.62). Cheliceral paturon relatively less
robust and narrower than in most Hadronyche spp. (Figs.
4B,D; 5B,E). Cheliceral groove narrow, V-shaped; central
teeth in single short row confined to the basal half of the
groove, lateral marginal tooth rows run full length of groove.
(Fig. 4F). Labium relatively short (LL/LW 0.70-0.75)
(Fig. 4A). Sternum ovoid to narrow ovoid. Apical segment
of posterior lateral spinnerets relatively long, digitiform
(PLSAPW 0.25-0.28) (Fig. 4G). Male femora I, II with
dorsal spines (Fig. 4F). Male palp with patella width less than
or equal to femur width. Bulb with a long, slender, curved
embolus (EmbmidW/L 0.06-0.08) (Fig. 6B). Male tibia II
with a large mid-ventral conical apophysis, the narrow apex
surmounted by short, peg-like spines (Fig. 6H,I); metatarsus
II sinuous (proximoventrally concave) with a small mid-
ventral apophysis (Fig. 6J,K). Spermathecal sacs narrow,
elongate (Fig. 5G).

Included species. Atrax robustus O.P.-Cambridge, A.
sutherlandi n.sp., A. yorkmainorum n.sp.

Distribution. Southeastern coast and highlands of Australia,
from the Hunter River region, New South Wales, to eastern
Victoria.

Comments. Burrows often with a weak or incomplete silk
lining, typically sited under rocks and logs

Figure 5. Atrax robustus, female: (A) sternum, labium and maxilla;
( B ) cephalothorax and chelicerae, lateral; (C) leg III, prolateral,
patella, tibia, metatarsus; ( D ) spinnerets; (E) cephalothorax and
chelicerae, dorsal; (F) cheliceral groove teeth; (G) spermathecae.
Scale lines 1 mm.

298

Records of the Australian Museum (2010) Vol. 62

Figure 6. Atrax robustus, male ( A,D-K , AMS KS3173; B,C, AMS
KS5170): (A) palp, prolateral; ( B ) bulb; (C) embolus, distal; (D,E)
distal patella and tibia I— D, ventral, E prolateral; (EG) metatarsus
I— F, ventral, G, prolateral; (//,/) distal patella and tibia II— H,
ventral, /, prolateral; (J,K) metatarsus II— J, ventral, K, prolateral.
Scale lines: 0.3 mm, except A,B, G, 1.0 mm.

Gray: Australian funnel-web spiders

299

Atrax robustus O.P.-Cambridge

Figures 4-7; Tables 2, 34

Atrax robustus O.P.-Cambridge, 1877: 26. Hogg, 1901: 273.
Musgrave, 1927: 33. Nishikawa, 1976: 179. Main, 1985:

41. Gray, 1978: 122. Gray, 1987: 314. Gray, 1988: 114.
Euctimena tibialis- Rainbow, 1914: 249. First synonymized
by Musgrave, 1927: 33.

Poikilomorpha montana- Rainbow, 1914: 265.

Atrax montana. -Main, 1985: 41 (Transferred from
Poikilomorpha ). First synonymized by Gray, 1988; 114.

Types. Holotype female: BMNH 71 (NMH). Type dry, pinned, labelled
“New Holland”, no other data (not seen).

Other material examined. New South Wales (males): AMS KS2698,
Austinmer, 6 January 1959, N. Mitchell; AMS KS4812, Avalon Beach, 14
March 1976, Porritt; AMS KS4837, Avoca, 11 January 1975; AMS KS4197,
Balgowlah, 7 January 1960, Burch; AMS KS4946, Balmoral, 18 April 1974,
Robinson & Hoey; AMS KS4902, Bankstown, 14 April 1965; AMS KS5137,
Baulkham Hills, 18 January 1973, M. Gray; AMS KS4080, Bayview, 2 April
1974, S. Thompson; AMS KS2956, Beecroft, 25 April 1979, D. Dean; AMS
KS3999, Belrose, 19 January 1967, Anderson; AMS KS1152, “The Mists”,
Berambing, December 1976, K.J. Street; AMS KS8376, Berowra, December
1981, J. Hingley; AMS KS1852, Bilgola Plateau, 26 May 1977, H. Newton;
AMS KS1148, Bilpin, 17 January 1930, E.M. Hunt; AMS KS4961, Burraneer
Bay, 15 December 1972; AMS KS1154, Camden, 21 February 1974, G.
Seymour; AMS KS4914, Campsie, 12 February 1975, R.N. Peck; AMS
KS4915, Caringbah, 4 April 1972, M. Sinclair; AMS KS4773, Cardiff, 21
January 1979; AMS KS4194, Castlecrag, May 1963,1. Webb; AMS KS4484,
Charlestown, 1 February 1980, G. Anderson; AMS KS4372, Chatswood, 16
April 1972, W. Feonard; AMS KS3990, Cheltenham, 2 April 1974, S.K.
Pearson; AMS KS14383, Cherrybrook, 1 April 1984, M. Backing; AMS
KS2957, Chester Hill, 27 May 1979, J. Pauli; AMS KS4395, Clarevale Beach,

16 January 1976, K.H. Sinfield; AMS KS4378, Clifton Gardens, 20 April
1972, Dr C.A. Monticone; AMS KS5013, Cobbitty, 11 March 1959, T. Down;
AMS KS4066, Collaroy, near Dee Why Fagoon, 14 December 1958, A.
Kabanoff; AMS KS5072, Cromer, 6 April 1980, M. Duncan; AMS KS3036,
Denistone, 26 April 1979, M. Richison; AMS KS4001, Dundas, 1 March

1976, B. & A.C. Oldsen; AMS KS7470, Dural, March 1981; AMS KS2432,
East Maitland, H. Miller; AMS KS4815, Eastwood, 30 March 1976, D.
Griffith; AMS KS7477, Elizabeth Bay, 5 May 1981, D.T. Cave; AMS KS1153,
Engadine, 28 November 1972, S. Couglin; AMS KS1132, Epping, January

1977, W.J. Bunton; AMS KS1143, Faulconbridge, 8 January 1973, Stewart;
AMS KS5694, Fox Valley Reserve, nearWahroonga, 22 June 1980, B. Henke;
AMS KS4071, Frenchs Forest, 5 June 1972, N. Hailstone; AMS KS 13409,
Galston, 2 February 1981, G. Jackson; AMS KS 10976, Galston Gorge, 4
May 1983, G. Jackson; AMS KS1156, Gladesville, 20 February 1970,
McElwaine; AMS KS13752, Gordon, 22 December 1983, M. Gray; AMS
KS4771, Gosford, 21 January 1979, R.E. Mascord; AMS KS4005, Grays
Point, 5 February 1977, P. Poppett; AMS KS4382, Greenwich/Willoughby;
AMS KS1160, Hartley Vale, 8 December 1952, E.S. Miller; AMS KS4297,
Hazelbrook, 8 January 1980, S. Clark; AMS KS5071, Heathcote, 20 April
1980, R. Witchard; AMS KS2323, Helensburgh, 6 January 1979, N. West;
AMS KS1157, Hornsby, 3 March 1976, Reed; AMS KS4949, Hurstville, 5
June 1972, A. Osten; AMS KS1161, Kembla Heights, 11 November 1963,
G. Pearce; AMS KS8678, East Beach, Kiama area, 20 January 1982, D.
Bennedett; AMS KS4377, Killara, February 1972, P. Mayman; AMS KS3200,
Kurrajong Heights, 30 November 1969, G. Hunt; AMS KS4944, Fakemba,
7 January 1948; AMS KS1159, Fane Cove, 22 April 1930, ? Graham; AMS
KS1146, Fawson, March 1956, R. Schleicher; AMS KS8920, Findfield, 14
April 1982, B. Henstridge; AMS KS4078, McMahons Point, May 1972, N.
Mayfield; AMS KS6067, Manly, 3 November 1980; AMS KS 10783,
Matcham, near Gosford, 27 March 1983, van Derbruggen; AMS KS1162,
Minnamurra Falls, 12 May 1964, R. Mascord; AMS KS4022, Mona Vale, 26
February 1973, T. Govranik; AMS KS4948, Mortdale, 13 February 1975;
AMS KS5314, Mosman, 19 June 1980, S. Pfeiffer; AMS KS4947, Mount
Colah, 25 January 1979, M. Gray; AMS KS1155, Mount Irvine, 14November
1944, E.F. Troughton; AMS KS5016, Mount St. Thomas, near Wollongong,

17 June 1956, F. Evans; AMS KS2146, MountWilson, 15 November 1978-11
December 1978, C. Horseman; AMS KS4009, Neutral Bay, AMS KS3189,
Newport, September 1959, A.J. Wright; AMS KS13560, Northbridge, 30
December 1983, J. Mayer; AMS KS4950, Parramatta, 25 March 1959, W.G.
Ashford; AMS KS4951, Peakhurst, 24 April 1945, H. Inder; AMS KS2712,

Pennant Hills, 21 March 1979, Thompson; AMS KS4045, Penrith, 25 August
1979, A. Johnson; AMS KS4399, Potts Point, 5 July 1956, P. Berry Smith;
AMS KS4015, Pymble, 14 March 1949; AMS KS7466, Queenscliff, April
1981; AMS KS4027, Rhodes, 12 May 1930, T. White; AMS KS4921,
Riverwood, 12 April 1966, R.E. Mascord; AMS KS2995, Roseville, 3 June
1979, ? Hidder; AMS KS4869, Rydalmere, 13 March 1974, R. Hood; AMS
KS4070, Ryde, January 1971; AMS KS4007, St. Ives, 12 May 1971; AMS
KS4083, Scotland Island, 1 April 1974; AMS KS3457, Springwood, 1978,

A. McFean; AMS KS4922, Stanmore, 1949, January; AMS KS4971, Telopea,
25 April 1971, K.P. Reid; AMS KS2713, The Oaks, Apr 1979, P. McIntosh;
AMS KS4203, The Spit, 4 July 1948, Page; AMS KS4013, Thomleigh; AMS
KS4373, Turramurra, 1 April 1970; AMS KS8664, Wahroonga, 31 January
1982, A.S. Martin; AMS KS4008, Waitara, 11 March 1949; AMS KS 10782,
Warrawee, 12 April 1983, Ralleston; AMS KS5146, Waverley, 15 May 1973,
F. Gibson; AMS KS4367, Waverton, 4 March 1971, Dr C. Browne; AMS
KS5131, West Pennant Hills, 19 May 1973, R. McDonald; AMS KS6148,
West Ryde, 1 January 1980, W. Bennett; AMS KS4000, Whale Beach, 21
April 1974; AMS KS4025, Willoughby, 30 July 1973, A.C. Jones; AMS
KS3185, Winston Hills, 21 July 1973, M. Gray; AMS KS3404,AMS KS5720,
Woonona, near Bulli, December 1960, S.H. Roberts; AMS KS4662,
Wyoming, near Gosford, 14 March 1980; AMS KS2670, New Fambton, near
Newcastle, 1979, F. Hallinan; AMS KS1150, Newcastle, 13 March 1963,

B. C. Dyson; AMS KS4125, Wyong, 18 March 9178; AMS KS2923,
Benandarah State Forest, 22 March 1979-25 April 1979, C. Horseman; AMS
KS5205, Foxground, near Gerringong, 10 April 1980, G. Wishart; AMS
KS5170, Ryde, 33°49'S 151°06F, 20 May 1980, A. Brown. New South
Wales (females): AMS KS4076, Artarmon, Apr 1927, J. Stewart; AMS
KS4930, Ashfield, 20 December 1930, E. Hudson; AMS KS 1826, Austinmer,
23 November 1959, F.E. Havand; AMS KS1377, Avalon, 15 May 1978, F.
Reneman; AMS KS5017, Avoca, 20 September 1968, N. Tweedale; AMS
KS4825, Balgowlah, 2 May 1970, G. Smith; AMS KS4987, Bankstown, 19
April 1949, E.A. Cox; AMS KS5140, Baulkham Hills, 18 January 1973, M.
Gray; AMS KS2716, Bayview, 20 April 1979, B. Makins; AMS KS4358,
Beecroft, February 1928, G.A. Heumann; AMS KS4903, Belmore, 25 April
1924, M. Codd; AMS KS4803, Belrose, 19 January 1967, Anderson; AMS
KS1827, Bilpin, 12 April 1972, R. McDonald; AMS KS7564, Birmingham
Gardens, near Newcastle, 6 December 1978, R. Mascord; AMS KS1674,
Blackheath, 24 April 1948, R. McKay; AMS KS4887, Bowral; AMS KS8733,
Bundanoon, February 1982, B. Erdman; AMS KS4913, Camden, 21 February
1974, G. Seymour; AMS KS4408, Berowra, 15 September 1973, J. Disney;
AMS KS3524, Carlingford, 26 September 1979, J. Armitage; AMS KS3175,
Castle Hill, Apr 1979, F. Pearce; AMS KS4206, Castlecrag, 17 July 1973, J.
Bunce; AMS KS4242, Chatswood, 14 November 1928, R. Barnes; AMS
KS4325, Cheltenham, Apr 1972, D. Fevy; AMS KS4211, Clarevale Beach,
F. & M. Mortimer; AMS KS4030, Clifton Gardens, 21 April 1961, Bradley;
AMS KS3188, Collaroy, February 1928, E. Osborne; AMS KS4400,
Collaroy Plateau, July 1966, B. Tye; AMS KS4892, Colo Vale, July 1958,
D.H. Wallace; AMS KS4905, Coogee, November 1926, Stokes; AMS
KS4233, Cremorne, 21 June 1927, F.E. Grutzmacher; AMS KS4962,

Figure 7. Collection records for Atrax robustus (squares) and A.
yorkmainorum (circles).

300

Records of the Australian Museum (2010) Vol. 62

Cronulla, 13 January 1971; AMS KS4828, Dee Why, May 1932, Rowe;
AMS KS4804, Dundas, January 1975, M. Denham; AMS KS4865, Dural,
18 January 1971, M. Thomley; AMS KS3319, Eastwood, 8 July 1979, J.
Cullen; AMS KS4829, Elanora, 1 March 1967, S. Marshall; AMS KS4989,
Elizabeth Bay, 9 January 1958, D.T. Cave; AMS KS1830, Emu Plains, 30
July 1957, G.D. Sutherland; AMS KS4770, Enfield, 5 June 1905; AMS
KS4546, Epping, 17 February 1980, J. Brand; AMS KS4990, Five Dock,
May 1928, R. Williams; AMS KS4361, Forestville, 19August 1971; AMS
KS8868, Freeman’s Reach, near Windsor, 26 January 1982, M. Gray;
AMS KS8377, French’s Forest, 24 October 1981, R. Scott; AMS KS4661,
Galston, 3 February 1980; AMS KS4963, Garrie Beach, 7 October 1965,
R. Mascord; AMS KS4839, Gosford, 25 September 1964, E. Worrell; AMS
KS4207, Greenwich, 5 May 1970, K. Burns; AMS KS4966, Guildford
West, 27 August 1957, S.R. Kirkwood; AMS KS1220, Hazelbrook, 23
November 1976, G. Weatherspoon; AMS KS1675, Hazelbrook, February
1976; AMS KS4985, Heathcote, 26 June 1975, R. Witchard; AMS KS4967,
Helensburgh, 11 June 1951, L. Reed; AMS KS4225, Hornsby, 7 May 1940;
AMS KS1208, Jamieson Valley, 22 June 1953, A.W. Gayley; AMS KS4065,
Killara, 8 March 1928; AMS KS5078, Kingsgrove, 1952; AMS KS4993,
Kingswood, May 1962; AMS KS4410, Kuring-ai Chase National Park, 23
January 1980, M. Gray; AMS KS1310, Kurrajong, February 1978; AMS
KS4945, Lakemba, 17 March 1930; AMS KS4853, Lane Cove, 15 March
1973; AMS KS4968, Leichhardt, 2 September 1913, D. Hunter; AMS
KS1834, Linden, 3 September 1950, K. Baames; AMS KS5031, Lindfield,
29 September 1962, M. & V. Gregg; AMS KS4234, McMahons Point, Apr
1972, J. Suich; AMS KS6025, Manly, Manly Hospital, 3 November 1980;
AMS KS4907, Matraville South, 20 October 1952, A. Brown; AMS
KS4238, Meadowbank, March 1963, R.H. Hall; AMS KS1829, Megalong
Valley, R. lbow; AMS KS4856, Middle Harbour, 6 September 1933,
Kaufman; AMS KS4896, Mittagong, 30 December 1975, M. & V. Gregg;
AMS KS4360, Mona Vale, 9 November 1975, J. Walker; AMS KS1125,
Morpeth; AMS KS4204, Mortdale, 31 January 1974, C. Stewart; AMS
KS4403, Mosman, 3 March 1932, Patton; AMS KS3693, Mount Colah, 29
July 1979, J. Fairlie; AMS KS4189, Mount Fairy, 25 November 1979; AMS
KS1835, Mount Irvine, 14 November 1944, E. Troughton; AMS KS2960,
Mount Keira, 12 May 1979, Walsh; AMS KS4832, Mount Kuring-ai, 5
November 1972; AMS KS4808, Naremburn, 26.ii. 1961, M. Fairbum; AMS
KS1126, Narara, 20 March 1930, B.R. Everingham; AMS KS4405,
Narrabeen, 29 September 1957, L.B. Trapp; AMS KS4046, Neutral Bay, 3
November 1979; AMS KS797, Newport, 7 December 1977; AMS KS4064,
Normanhurst, 6 April 1969, ? Dymock; AMS KS4085, Northbridge, 26
July 1975, D. Dickerson; AMS KS3401, Northmead, March 1973, J.
Deviana; AMS KS4195, North Narrabeen, 11 June 1952, L.J. Lizara; AMS
KS4232, North Sydney, May 1925, Dr S.J.W. Moreau; AMS KS4908,
Oatley, 15 February 1963; AMS KS4858, Palm Beach, 1 February 1951,
H.R. Swaine; AMS KS4909, Parramatta, 21 March 1910; AMS KS1131,
Peakhurst, 15 November 1969, D. Fantom; AMS KS1837, Pearl Beach,
24.iii.1966, J. Cooks; AMS KS1930, Pennant Hills, 6 October 1978, S.
el-Issa; AMS KS4045, Penrith, 25 July 1979, A. Johnson; AMS KS4969,
Penshurst, R. Pollett; AMS KS1844, Picton, November 1946, J.K.
McFarlane; AMS KS4072, Pittwater, 1927, A. Musgrave; AMS KS4824,
Point Clare, T.A. Trewheeler; AMS KS4910, Potts Point, 5 October 1969,
C. Catterall; AMS KS4931, Punchbowl, 17 July 1947, F:J. Donald; AMS
KS4809, Pymble, November 1971, A. Ritchie; AMS KS1850, Richmond,
12 October 1977, Ambulance Station; AMS KS3168, Roseville, 18 January
1948, H. Chadwick; AMS KS4964, Royal National Park, October 1966, R.
Mascord; AMS KS4911, Rushcutters Bay, 24 May 1971, S.J. Penlington;
AMS KS4810, Ryde, 28 October 1969, J. Garden; AMS KS4084, St. Ives,
March 1967; AMS KS4912, St. Peters, 2 February 1959, D.J. Gilks; AMS
KS5240, Scotland Island, 29 February 1980, J. Lowry; AMS KS5023,
Somersby, J Brown; AMS KS1839, Springwood, 4 August 1971; AMS
KS1135, Sydney, 17 November 1969; AMS KS2715, Terry Hills, 30 April
1979, R. Gleeson; AMS KS4891, The Oaks, 25 February 1956; AMS
KS4899, Thirroul, September 1933, D. King; AMS KS796, Thomleigh, 8
January 1978; AMS KS4844, Tuggerah, September 1927, H.W. Legge;
AMS KS4376, Turramurra, 1928, H.S. Wales; AMS KS13791, Wahroonga,
2 February 1984, A.S. Martin; AMS KS4845, Wamberal, 4 December 1972,
J.N. Kgour; AMS KS4243, Warrawee, 24 December 1972, ? Rickets; AMS
KS4063, West Pennant Hills, 15 May 1973, S. Zucher; AMS KS6069, West
Ryde, 10 November 1980; AMS KS4846, West Wallsend, 30 May 1956, J.
Donne; AMS KS4847, Weston, 4 November 1969,1. Waugh; AMS KS4357,
Willoughby, 17 August 1969; AMS KS5151, Winston Hills, 9 September
1972, J. Deviana; AMS KS1848, Wirrimbirra Sanctuary, 9 July 1969, M.N.
Fackender; AMS KS4862, Woolwich, May 1927, H. Switzer; AMS KS4848,
Woy Woy, 13 July 1933, B. Hynson; AMS KS8295, Yagoona, February
1981; AMS KS1274, Yerranderie, 20 February 1978, H. Kenny.

Diagnosis. CL 9.22-12.30 (male). Differs from other species
by having a relatively spinose palpal tibia (4-11) with dorsal
spines present. Differs from A. sutherlandi by having a more
elongate embolus (BulbW/EmbL 0.30) (Fig. 6B), and from
A. yorkmainorum by larger body size (CL 9.22-12.30) and
shallower offset of embolus base from tegulum (Fig. 6B).

Male (AMS KS4016, Gordon, New South Wales, 32°38’S
148°39'E, F. Hatton, 10 December 1949). — Size. Carapace
length 10.20, width 8.70. Abdomen length 9.52, width
7.38. — Colour. Basic colour pattern. — Carapace. Longer
than wide. Height 2.86. Frontal width 5.14. Fovea narrow,
procurved. Mid-dorsal cephalic setae reach fovea. Anterior
strial setae numerous. Anterolateral carapace angle with
small, weak bristles. — Eyes. Central eye region weakly
raised. Eye group width 1.85. Median ocular quadrangle
length 0.70; anterior width 0.76, posterior width 1.13.
Diameters: AME 0.28, ALE 0.58, PLE 0.38, PME 0.26.
— Chelicerae. Cheliceral groove narrow, weakly divergent.
Groove length 2.92, middle width 0.41. Cheliceral teeth: 18
central, occupying proximal half of groove; 12 prolateral;
13 retrolateral. — Labium. Much wider than long, apically
indented. Length 1.08, width 1.68. Labiosternal sigilla
broad, entire. Cuspules c. 300, number high, in crescentic
transverse band. — Sternum. Ovoid, long. Length 5.45,
width 3.88. Posterior sigilla ovoid, broad. Strong bristles
on posterior margin. — Palp. Bulb about as long as tibia.
Tegular region rather small, slightly wider than long. Base
of embolus weakly offset from tegulum. Embolus shaft
long, gently curved, strongly tapered. Distal embolus
weakly twisted, ejaculatory groove wide. Bulb length
4.62, width 1.12. Embolus length 3.60, midwidth 0.25.
Length of femur 4.35, patella 1.96, tibia 4.71. Width of
tibia 1.80. Spination: femur 7, patella 5, tibia 10 (dorsal
spines present). Some distal femoral bristles and spines
sinuous. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.43

4.21

6.36

6.69

3.87

29.56

2

7.75

3.86

6.36

6.28

3.89

28.14

3

7.14

3.18

5.15

6.03

3.94

25.44

4

8.32

3.81

6.80

7.41

4.40

30.74

Tibia I width 1.50. Femora I and II with dorsal spines. Leg

I unmodified, tibial and metatarsal spines numerous. Tibia

II with large, conical, apophysis set just proximal of centre.
A few ventral tibial spines placed proximally, absent in
distal half (except apical ventral spines). Distal tibia II
ventrally concave. Metatarsus II sinuous, curved proximally,
ventral spines placed upon and distal to a small, subcentral
apophysis. Anterior coxal hairs unmodified. Scopulae
legs I-IV: tarsus, weak; distal metatarsus, absent to weak.
Tarsal claw teeth legs I, II: superior 13, 11; inferior 2, 2.
Trichobothria legs I, II: tarsus 20, 19; metatarsus 15, 12;
tibia p9 r9, p8 r8. — Leg spination. Leg I; femur 9 (d6 p3),
patella 13 (p2), tibia 51 (p8), metatarsus 29, tarsus 20. Leg II:
femur 12 (d6 p6), patella 6 (p2-3); tibia 18 (p4), metatarsus
22(pl), tarsus 23. Leg III: femur d7 (plus strong bristles),
patella 5 (p3 r2), tibia 15 (p2 r3), metatarsus 25, tarsus 25.
Leg IV: femur 0 (long bristles present), patella rl, tibia 18 (pi
r4), metatarsus 20, tarsus 21. — Abdomen. Posterior lateral
spinnerets, lengths: total 4.68; basal segment 1.52; middle
1.08; apical 2.08. Apical segment width 0.60.

Gray: Australian funnel-web spiders

301

Table 2. Male morphological data— Atrax robustus (n =

24).

character

range

mean

character

range

mean

character

ratio

SD

CL*

9.22-12.30

10.48

MtlS

24-32 (p0-l)

27

CW/CL

0.89

0.033

CW

8.27-10.59

9.28

TalS

12-27

19

CH/CW

0.35

0.025

CH

2.86-4.06

3.28

Ti2S

13-27 (p2-5)

18

CFW/CL

0.55

0.038

CFW

4.49-7.38

5.78

STC2teeth

10-15

13

CHGW/L

0.14

0.015

ChGL

2.84-3.60

3.09

BulbL

4.08-9.06

5.79

SW/SL

0.74

0.043

ChGW

0.41-0.55

0.44

EmbL

3.28-5.26

4.25

LL/LW

0.75

0.052

ChGCT

10-29

20

BulbW

1.12-1.48

1.28

PLSAPW/L

0.25

0.032

LL

1.08-1.62

1.41

EmbmidW

0.20-0.30

0.27

BulbW/EmbL

0.30

0.055

LW

1.60-2.16

1.87

PalpTibL

4.65-6.61

5.53

EmbmidW/L

0.06

0.012

CUSPS*

261-494

340

PalpTibW

1.59-2.08

1.83

PalpTibW/L

0.33

0.035

SL

5.30-6.56

5.81

PalpTibS*

4-11

8

BulbL/Tib L

0.97

0.068

SW

3.88-5.83

4.31

PalpPatS*

1-8

5

PLSAPW

0.46-0.64

0.53

PalpFem*

5-9

6

PLSAPL

1.64-2.40

2.03

FelS

2-10

6

PalS

6-13 (p2-4)

10

Female (AMS KS13470, Gordon, N.S.W., 32°38’S 148°39’E,
1 November 1983, C. Horseman). — Size. Carapace length
12.63, width 10.41. Abdomen length 14.82, width 12.07.
— Colour. Basic colour pattern. — Carapace. Much longer
than wide. Height 2.72, frontal width 8.28. Cephalic length
8.50. Fovea narrow, procurved, anterior margin slightly
indented. Mid-dorsal cephalic setae numerous, reach back
to fovea. Anterior strial setae numerous. Strial and marginal
setae numerous, rather short. Anterolateral carapace angle
with several short, weak bristles. — Eyes. Central eye region
slightly raised. Eye group width 2.60. Diameters: AME 0.26,
ALE 0.60, PLE 0.44, PME 0.36 Interdistances: AME-AME
0.39, AME-ALE 0.27, ALE-PLE 0.27, ALE-PME 0.14,
PME-PME 1.05. Median ocular quadrangle length 0.90,
anterior width 0.94, posterior width 1.73. — Chelicerae.
Groove narrow, margins diverging distally; groove length
3.70, middle width 0.59. Cheliceral teeth: 29 central,
occupying basal half of groove; 15 prolateral; 14 retrolateral.
— Labium. Wider than long, anterior margin weakly
indented. Length 2.18, width 2.64. Cuspules occupying
central two thirds of labium. Labiosternal sigilla entire. —
Sternum. Ovoid. Length 6.89, width 5.17. Posterior sigilla
ovoid: length 1.10, width 0.60. — Palp. Spination: patella
pi (bristle-like), tibia 5, tarsus 12. Trichobothria: tibia pll
rlO, tarsus 22. Tarsal claw with 10 teeth. — Legs. 4123. Legs
I, IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

9.50

5.32

7.32

6.40

3.44

31.98

2

8.45

4.88

6.15

5.79

3.28

28.55

3

7.46

3.94

4.80

5.66

3.69

25.55

4

9.00

4.88

6.71

7.18

4.22

31.99

Tibia I width 2.30. Metatarsus I proximal width 1.44. Coxae
I, II with short, anteromedial bristle-like setae. Tarsal claw
teeth legs I, II: superior 11,11; inferior III, IV. Trichobothria
legs I, II: tarsus 24, 24; metatarsus 21, 19; tibia plO r9, p9
rlO. — Leg spination. Leg I: femur pi, patella pi, tibia 3 (p2),
metatarsus 11, tarsus 15. Leg II: femur p2, patella p2, tibia
5 (p2), metatarsus 14, tarsus 14. Leg III: femur 0, patella

3 (p2 rl), tibia 8 (p2 r2), metatarsus 23 (p9 r2), tarsus 17.
Leg IV; femur 0, patella rl, tibia 6 (pi r2), metatarsus 22
(p6 r2), tarsus 25. — Abdomen. Posterior lateral spinnerets-
lengths: total 6.20; basal segment 2.40, middle 1.32, apical
2.48. Apical segment width 0.77. — Genitalia. Spermathecae
long, slender; gently curved toward midline. Basal separation
twice the width of a spermatheca. Length 1.50, width 0.41.

Distribution. Sydney, Central Coast and Illawarra regions
and west to the Blue Mountains (Fig. 7).

Comments. Specimens from the Hunter River/Newcastle
region are typically larger than other individuals (male CL
c. 12 mm) and the palpal tibia and embolus may be more
elongate.

Mating position in A. robustus. Mating was observed for
A. robustus (Gray, 1986), a species with well-developed
apophyses on tibia and metatarsus II. The position adopted
during its mating is almost identical to that recorded for H.
formidabilis (Fig. IF: Walker, in Mascord, 1980; Walker,
1982), another species with large apophyses, the A. robustus
male, uses his second legs to clasp the basal parts of the
female femora II, locking them in the cavity between
the tibial and metatarsal apophyses. At the same time he
braces his first legs against the chelicerae and palps of the
rearing female, pushing her up and backwards. The main
difference in the H. formidabilis mating was the distal
crossing of the bracing first legs (Fig. IF)—this was not
seen in the A. robustus matings. Hickman (1964) described
mating in H. infensa, a species lacking leg II apophyses.
The accompanying plate shows the spiders in an almost
identical position to that observed for A. robustus (although
Hickman’s notes are somewhat contradictory).

Snazel & Allison (1989) noted the marked similarities
between mating in H. formidabilis and Macrothele, another
genus in which leg II modifications are present or absent.
Coyle (1986) observed similar mating behaviour in Euagrus
sp., a diplurid genus with tibia II apophyses.

302

Records of the Australian Museum (2010) Vol. 62

Atrax yorkmainorum n.sp.

Figures 7-10; Tables 3, 34

Etymology. The species is named for Dr Barbara York Main
and Professor Bert Main, in recognition of their remarkable
achievements in Arachnology and Ecology.

Types. Holotype male: AMS KS1042, Batlow, New South Wales, 35°30'S
148°08'E, H.V. Smith, 16 June 1951. Paratypes (AMS). New South Wales.
Males: AMS KS4104, Bago Forest, nr. Batlow, 35°41'S 148°09’E, 5
November 1979, J. Kovacs; AMS KS4607, Batlow-Tumut area, 35°25'S
148°10'E, 5 March 1980, N. Robinson; AMS KS4718, Tumbarumba,
35°45'S 148°00'E, 25 Mar AMS KS1043, Batlow, 35°30'S 148°08’E. 1980;
AMS KS12361, AMS KS1050, Wondalga, 35°23'S 148°07’E, 14 October
1948, C.H. Jagoe; Females : AMS KS16451, Batlow, 35°30’S 148°08'E,
16 June 1951, H.Y. Smith; AMS KS5054, Bago Forest, 35°41'S 148°09’E,
8 January 1979, J. Kovacs; AMS KS1047, Tumut, 35°18'S 148°13'E, 4
May 1977, B. Buckley. Australian Capital Territory. Males : Uriarra Forest,
Brindabella Range, 35°18'S 148°56F, 9 July 1983, M.R. Gray; AMS
KS13396, Tidbinbilla Nature Reserve, 35°28’S 148°54'E, 26 October 1983,
W. Osborne. Females : AMS KS3542, Tidbinbilla Nature Reserve, 35°28'S
148°54'E, November 1978, P. Ormay; AMS KS928, Gibraltar Falls, 35°29'S
148°56'E, 24 October 1971, T. McGregor.

Other material examined. New South Wales (males). AMS KS13534,
Bago Forest, near Batlow, 5 November 1979, J. Kovacs; AMS KS1041,
Batlow, December 1948, C.H. Jagoe; AMS KS1042, Dora Dora National
Park, Jingellic near Albury, 16 January 1979, A.B. Rose (AMS). New
South Wales (females): AMS KS1041, Batlow, December 1948, C.B.
Jagoe; AMS KS4608, Batlow-Tumut area, 5 March 1980, N. Robinson;
AMS KS7566, Cooma, 19 January 1980, R. Mascord; AMS KS13545,
Gundagai, 25 October 1983. Australian Capital Territory (males). AMS
KS 13397, Tidbinbilla Nature Reserve, 26 October 1983, W. Osborne; AMS
KS13592, Canberra area, 35°17'S 149°13F, 13 Dec 1983, T. Mitchell.
Australian Capital Territory (females): AMS KS654, Honeysuckle
Creek, 24 February 1977, M. Noble; AMS KS12032, Gudgenby National
Park; AMS KS929, near Piccadilly Circus, 25 January 1972, K. Watson.

Figure 8. Atrax yorkmainorum , male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
( D ) cephalothorax and chelicerae, dorsal; (E) cheliceral groove
teeth; ( F) palp, prolateral; (G) leg II, prolateral. Scale lines 1 mm.

Diagnosis. CL 7.39-9.22 (male). Usually smaller then A.
robustus and differs from it by palp lacking dorsal tibial
spines, embolus base more strongly offset from tegulum,
and apical part of embolus set at angle to strongly curved
embolic shaft (Fig. 10B,C). Differs from A. sutherlandi by
its more elongate, slender and curved embolus.

Male (holotype). — Size. Carapace length 8.42, width
7.48. Abdomen length 8.50, width 6.90. — Colour. Basic
colour pattern. — Carapace. Height 2.69. Frontal width
4.57. Fovea narrow, procurved. Mid-dorsal cephalic setae
few, reach fovea. Anterior strial setae present. Anterolateral
carapace angle with small, weak bristles. — Eyes. Central
eye region weakly raised. Eye group width 1.66. Median
ocular quadrangle length 0.57, anterior width 0.69, posterior
width 1.05. Diameters: AME 0.20, ALE 0.36, PLE 0.35,
PME 0.19. — Chelicerae. Cheliceral groove narrow, margins
subparallel, diverging slightly distally. Groove length 2.48,
middle width 0.26. Cheliceral teeth: 16 central, confined
to basal half of groove; 11 prolateral; 12 retrolateral. —
Labium. Much wider than long, apically weakly indented.
Length 1.49, width 1.08. Labiosternal sigilla broad, entire.
Cuspules c. 193, number moderate, in crescentic transverse
band; frequency moderate. — Sternum. Ovoid. Length 4.43,
width 3.62. Posterior sternal sigilla ovoid, broad. Strong
bristles on posterior margin. — Palp. Tegular area small,
slightly wider than long. Embolus long, slender; shaft tapered
and strongly curved. Distal embolus very slender, apical
region weakly twisted and set at angle to the embolic axis;
ejaculatory groove narrow. Embolus base strongly offset
from tegulum. Bulb length 4.16, width 1.10. Embolus length
3.23, midwidth 0.18. Length of femur 3.48, patella 1.57,
tibia 4.61. Width of tibia 1.71. Spination: femur 3, patella
0, tibia 3 (all bristle-like). Distal femoral bristles and spines
sinuous. — Legs. 4123.

Figure 9. Atrax yorkmainorum, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cheliceral
groove teeth; (D) spermathecae; (E) spinnerets; (F) cephalothorax
and chelicerae, dorsal; (G) leg III, prolateral, patella, tibia,
metatarsus. Scale lines 1 mm.

Gray: Australian funnel-web spiders

303

Figure 10. Atrax yorkmainorum, male (. A,D-K , AMS KS13592; B,C, AMS KS1034): (A) palp, prolateral; (B) bulb; (C) embolus, distal;
(. D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; (H,I) distal patella and tibia II— H,
ventral, /, prolateral; ( J,K) metatarsus II— J, ventral, K, prolateral. Scale lines: 0.3 mm, except A,F,K 0.2 mm and B 1.0 mm.

304

Records of the Australian Museum (2010) Vol. 62

Table 3. Male morphological data— Atrax yorkmainorum (n = 10).

character

range

mean

character

range

mean

character

ratio

SD

CL

7.83-9.22

8.47

MtlS

12-26 (p0-l)

19

CW/CL

0.92

0.027

CW

7.14-8.60

7.76

TalS

7-13

10

CH/CW

0.35

0.021

CH

2.53-3.03

2.72

Ti2S

13-20 (pl-3)

16

CFW/CL

0.56

0.042

CFW

4.14-5.20

4.69

STC2teeth

12-15

13

CHGW/L

0.11

0.016

ChGL

2.33-2.70

2.59

BulbL

3.93-4.78

4.36

SW/SL

0.77

0.032

ChGW

0.22-0.35

0.28

EmbL

2.92-3.71

3.37

LL/LW

0.72

0.023

ChGCT

9-18

14

BulbW

1.08-1.22

1.12

PLSAPW/L

0.28

0.023

LL

1.03-1.30

1.16

EmbmidW

0.18-0.25

0.20

BulbW/EmbL

0.34

0.019

LW

1.41-1.77

1.58

PalpTibL

4.32-5.10

4.72

EmbmidW/L

0.06

0.008

CUSP

167-228

199

PalpTibW

1.56-1.87

1.69

PalpTibW/L

0.36

0.022

SL

4.19-5.06

4.56

PalpTibS

1-4

3

BulbL/TibL

0.92

0.047

SW

3.28-3.83

3.52

PalpPatS

0-2

<1

PLSAPW

0.36-0.51

0.42

PalpFemS

3-4

3

PLSAPL

1.15-1.84

1.51

FelS

5-13

8

PalS

5-12 (p2-4)

8

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.40

3.40

4.98

4.73

2.69

22.20

2

5.98

3.26

4.69

4.61

2.61

21.15

3

5.30

2.79

4.01

4.59

3.06

19.75

4

6.19

2.92

5.00

5.44

3.40

22.95

Tibia I width 1.35. Femora I and II with dorsal spines.
Leg I unmodified, tibial spines numerous. Tibia II with
large, conical apophysis placed centrally. Metatarsus II
sinuous, curved proximally with a weak, spined apophysis
centrally; ventral spines lacking proximally. Anterior coxal
hairs unmodified. Tarsal scopulae legs I-IV: weak. Distal
metatarsal scopulae legs I-IV: absent to weak. Tarsal claw
teeth leg I, II: superior 14,13; inferior 0,2. Trichobothria legs
I, II: tarsus 15,15; metatarsus 14,14; tibia p8 r8, p8 r8. — Leg
spination. Leg I: femur 7 (d3 p4) (some bristle-like), patella
5 (p2), tibia 36 (p2), metatarsus 19, tarsus 10. Leg II: femur
10 (d4 p6) (plus several bristle-like), patella 5 (p2), tibia 16
(p2), metatarsus 16 (pi), tarsus 10. Leg III: femur 0 (long
bristles), patella 5 (p4 rl), tibia 14 (p2 r3), metatarsus 23,
tarsus 12. Leg IV: femur 0 (long bristles), patella rl, tibia 13
(pi r3), metatarsus 20, tarsus 17. Abdomen. Posterior lateral
spinnerets, lengths: total 4.41; basal segment 1.63; middle
1.18; apical 1.60. Apical segment width 0.41.

Female (paratype AMS KS16451). — Size. Carapace length
8.57, width 7.14. Abdomen length 10.27, width 7.89. —
Colour. Basic colour pattern. — Carapace. Much longer than
wide. Height 3.16; frontal width 5.64. Cephalic length 6.05.
Fovea narrow, procurved. Mid-dorsal cephalic setae few,
alternating large and small, reach back to fovea. Anterior
strial setae numerous. Anterolateral carapace angle with
weak bristles. — Eyes. Eye region slightly raised centrally.
Eye group width 1.65. Diameters: AME 0.20, ALE 0.45, PLE
0.35, PME 0.26. Interdistances: AME-AME 0.23, AME-
ALE 0.15, ALE-PLE 0.14, PLE-PME 0.07, PME-PME
0.59. Median ocular quadrangle length 0.59, anterior width
0.63, posterior width 1.05. — Chelicerae. Cheliceral groove
narrow, margins subparallel, weakly divergent distally.
Groove length 2.63, middle width 0.35. Cheliceral teeth: 16
central in basal half of groove; 11 prolateral, 12 retrolateral.
— Labium. Wider than long, anterior margin indented.
Length 1.36, width 1.78. Cuspules spread in shallow v-shape
across anterior two-thirds of labium. Labiosternal sigilla

entire or narrowed centrally. — Sternum. Ovoid, moderately
wide. Length 4.59, width 3.84. Posterior sigilla moderately
short, ovoid: length 0.75, width 0.37. — Palp. Spination: tibia
4, tarsus 14. Trichobothria: tibia p6 r7, tarsus 12. Tarsal claw
with 7 teeth. — Legs. 4123. Legs I, IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.98

3.57

4.56

3.71

2.24

20.06

2

5.30

3.37

3.88

3.47

2.24

18.26

3

4.56

2.82

3.15

3.30

2.38

16.21

4

5.59

3.18

4.28

4.38

2.75

20.18

Tibial width 1.51. Metatarsus I proximal width 1.02. Coxal
setae unmodified. Tarsal claw teeth legs I, II: superior 9, 9;
inferior 2,2. Trichobothria legs I, II: tarsus 13,15; metatarsus
12, 11; tibia p9 r8, p8 r7. — Leg spination. Leg I: femur 0,
patella 0, tibia 2, metatarsus 11, tarsus 10. Leg II: femur
0, patella 0, tibia 2 (pi), metatarsus 12, tarsus 8. Leg III:
femur 0, patella pd3, tibia 6 (p2 rl), metatarsus 20 (p6 rd2).
Leg IV: femur 0, patella 0, tibia 4, metatarsus 14, tarsus
12. — Abdomen. Posterior lateral spinnerets, lengths: total
4.20; basal segment 1.73, middle 1.08, apical 1.39. Apical
segment width 0.45. — Genitalia. Spermathecae elongate,
strongly curved, approximated basally and apically but
well separated in middle. Apical region slightly wider than
remainder of spermatheca. Length 1.84, width 0.35.

Distribution. Southeastern New South Wales and Australian
Capital Territory (Fig. 7).

Atrax sutherlandi n.sp.

Figures 11-14; Tables 4, 34

Etymology. The species is named for the late Professor
Struan Sutherland, whose venom research resulted in the
production of a successful funnel-web spider antivenom.

Types. Holotype male: AMS KS1076, Bermagui, New South Wales, 36°25'S
150°04'E, K Walker. Paratypes. New South Wales (AMS). Males : AMS
KS7464, Bega, 36°40’S 149°54’E, K. Walker; AMS KS 1595, Nadgee Nature
Reserve; AMS KS1074, Bermagui, 36°25'S 150°04E, K. Walker; AMS
KS 11748, Bondi State Forest, nr. Bombala, 37°06'S 149° 15E, 14 November
1980, G. Gowing et al.; AMS KS1075, Bermagui, 36°25’S 150°04E, K.
Walker. Females: AMS KS5127, Bermagui, 36°25'S 150°04’E, 18 May
1973, M. Gray; AMS KS4886, Bockelo Creek, 18 May 1972, M. Gray;

Gray: Australian funnel-web spiders

305

Figure 11. Collection records for Atrax sutherlandi.

AMS KS11848, Bondi State Forest, nr. Bombala, 37°06'S 149°15’E, 14
November 1980, G. Gowing; AMS KS3618, Merimbula, 36°53’S 149°54'E,
6 September 1976, H. Harvey. Victoria. Males. K11078 (MV), Noorinbee,
37°31'S 149° 10E, Vic., 12 November 1969, A. Neboiss; K11088 (MV),
Cann River, 37°34’S 149°09'E, 5 March 1964. Females (AMS): AMS
KS3397, Mallacoota Inlet, 37°33’S 149°46E, 9 October 1971, S. Sutherland;
AMS KS1092, Genoa, 37°28’S 149°35’E, 1 November 1964.

Other material examined. New South Wales (males): AMS KS8467,
Bombala, 1981; AMS KS5118, Bega, 17 May 1973, Ambulance Station;
Merimbula, March 1965 (MV); AMS KS9561, Monga; AMS KS5120,
Mount Doctor George, near Bega, 17 May 1974; AMS KS6072, Mumbulla
State Forest, near Bega, November 1980, R. Wells; AMS KS1085,
Bemboka, 10 May 1970, M. Clune; AMS KS 14232, Bermagui, 8 April
1984; AMS KS11528, Bondi State Forest, near Bombala, 14 November
1980, G. Gowing et al.; AMS KS1084, Brogo, 4 December 1962, K.
Walker; AMS KS5497, Kioloa State Forest, 4 October 1979-30 October
1979, C. Horseman; AMS KS5121,Beckelo Creek, 18May 1973,M. Gray;

Figure 12. Atrax sutherlandi, male: (A) cephalothorax and
chelicerae, lateral; (B) leg II, prolateral; (C) cheliceral groove teeth;
(D) cephalothorax and chelicerae, dorsal; (E) sternum, labium and
maxilla; (F) spinnerets; (G) palp, prolateral. Scale lines 1 mm.

AMS KS5119, Bega, March 1974, Ambulance Station; AMS KS5126,
Bega, Autumn 1973; AMS KS1086, Bemboka, July 1977, G. Wishart;
AMS KS1074, Bermagui, K. Walker; AMS KS3619, Bombala, 17 January
1930, A.J. Barrett; AMS KS11445, Bondi State Forest, near Bombala, 31
March 1981, G. Gowing; Brown Mountain, 30 March 1967, R.J.B. (ANIC);
AMS KS5042, Candelo, 17 May 1973, M. Gray; AMS KS7299, Cooma, 6
April 1981, A.N. Wallace; AMS KS1102, Merimbula, 5 June 1964; AMS
KS1595, Nadgee Nature Reserve; AMS KS1088, Narooma, 29 January
1959, R Martin; AMS KS3624, Narooma, 15 June 1971, J. Cobcroft;
AMS KS1083, Nerrigundah, 1 October 1977, C. Cowall. Victoria (males):
AMS KS1055, Murrindal, 2 km N. of, 3 June 1973, L. Windsor; Buchan,
9 January 1964, Dingey & Penshurst (MV); AMS KS1087, Cobargo, June
1957, C.G. Bradford; Cobargo, 5 November 1964 (MV); Mallacoota,
26 September 1963 (MV); AMS KS1057, Gelantipy, 6 May 1947; AMS
KS1091, Mallacoota, May 1957; Mallacoota, January 1963, C. McArthur;
AMS KS3397, Mallacoota Inlet, October 1971, S. Sutherland; AMS
KS1055, Murrindal, 3 June 1973, L. Windsor; AMS KS1095, Noorinbee,
17 June 1957, N.A. Wakefield; AMS KS1096, Noorinbee, 25 September
1963, D.S. Broome; AMS KS1108, Wingan, 27 February 1946.

Diagnosis. CL 7.22-10.20 (male). Differs from other Atrax
spp. by having a relatively shorter and wider embolus, and
a shorter palpal tibia (PalpTibW/ L 0.43) (Fig. 14A-C;
Table 4).

Male (holotype). — Size. Carapace length 7.56, width 6.59.
Abdomen length 6.93, width 5.76. — Colour. Basic colour
pattern. — Carapace. Longer than wide. Height 2.55. Frontal
width 4.12. Fovea narrow, procurved. Mid-dorsal cephalic
setae reach fovea, anterior strial setae present. Anterolateral
carapace angle with small, very weak bristles. — Eyes.
Central eye region weakly raised. Eye group width 1.58.
Median ocular quadrangle length 0.63, anterior width 0.71,
posterior width 1.03. Diameters: AME 0.19, ALE 0.44, PLE
0.32, PME 0.25. — Chelicerae. Cheliceral groove narrow,
margins subparallel except distally where they diverge.
Groove length 2.27, middle width 0.24. Cheliceral teeth:
23 central, confined to basal half of groove; 13 prolateral;

Figure 13. Atrax sutherlandi, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) leg III,
prolateral, patella, tibia, metatarsus; ( D ) cephalothorax and
chelicerae, dorsal; (E) spinnerets; (F) spermathecae; (G) cheliceral
groove teeth. Scale lines 1 mm.

306

Records of the Australian Museum (2010) Vol. 62

Table 4. Male morphological data— Atrax sutherlandi (n = 17).

character

range

mean

character

range

mean

character

ratio

SD

CL

7.22-9.64

8.23

MtlS

18-35

23

CW/CL

0.87

0.039

CW

6.49-8.45

7.21

TalS

9-24

13

CH/CW

0.36

0.025

CH

2.13-3.36

2.62

Ti2S

10-23 (1-4)

15

CFW/CL

0.62

0.039

CFW

3.81-5.57

4.45

STC2teeth

12-14

13

CHGW/L

0.12

0.018

ChGL

2.15-3.16

2.41

BulbL

2.79-4.40

3.26

SW/SL

0.70

0.042

ChGW

0.19-0.34

0.28

EmbL

2.02-3.01

2.34

LL/LW

0.70

0.056

ChGCT

10-32

21

BulbW

0.90-1.21

1.02

PLSAPW/L

0.25

0.023

LL

0.92-1.39

1.07

EmbmidW

0.15-0.22

0.19

BulbW/EmbL*

0.44

0.043

LW

1.36-1.77

1.53

PalpTibL

3.24-4.56

3.65

EmbmidW/L*

0.08

0.012

CUSP

142-295

209

PalpTibW

1.34-1.85

1.56

PalpTibW/L*

0.43

0.035

SL

4.25-5.47

4.83

PalpTibS

1-3

3

BulbL/TibL

0.89

0.077

SW

2.82-3.85

3.41

PalpPatS

0-1

<1

PLSAPW

0.38-0.66

0.44

PalpFemS

0-4

2

PLSAPL 1.26-2.07 1.53

FelS 2-10 6

PalS 5-16 (p0-3) 10

13 retrolateral. — Labium. Much wider than long, apically
weakly indented. Length 0.99, width 1.49. Labiosternal
sigilla broad, entire. Cuspules c. 218, number moderate,
grouped in broad, central band. — Sternum. Ovoid, long.
Length 4.73, width 3.28. Posterior sigilla ovoid. Bristles on
posterior margin strong. — Palp. Tegular area wider than
long. Embolus of moderate length, shaft gently curved and
strongly tapered. Distal embolus slender, weakly twisted,
ejaculatory groove wide. Embolus base moderately to
strongly offset from tegulum. Bulb length 2.82, width 0.96.
Embolus length 2.10, midwidth 0.19. Length of femur 3.32,
patella 1.47, tibia 3.26. Width of tibia 1.48. Spination: femur
3, patella 0, tibia 2. Distal femur with sinuous spines and
strong bristles. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.63

3.16

4.23

4.08

2.41

19.51

2

5.10

2.98

4.10

4.00

2.43

18.60

3

4.75

2.61

3.41

3.92

2.81

17.50

4

5.28

2.79

4.61

4.80

3.04

20.52

Tibia I width 1.26. Femora I and II with dorsal spines. Leg

I unmodified, patella and tibial ventral spines numerous.
Tibia II with large, conical apophysis placed just proximal
of centre. Metatarsus II sinuous, strongly curved proximally,
with a small, spined, central apophysis. Anterior coxal hairs
unmodified. Tarsal scopulae legs I-IV: weak to moderate.
Distal metatarsal scopulae legs I-IV: absent to weak.
Tarsal claw teeth legs I, II: superior 13, 13; inferior 3, 2.
Trichobothria legs I, II: tarsus 14, 13, metatarsus 14, 14;
tibia p9 r9, p9 r8. —Leg spination. Leg I: femur d6, patella

II (p2), tibia 39 (p2), metatarsus 24, tarsus 12. Leg II: femur
d7, patella 5 (p2), tibia 16 (p3), metatarsus 16 (pi), tarsus
13. Leg III: femur 0 (long bristles), patella 6 (p5 rl), tibia
11 (p4 r3), metatarsus 15, tarsus 11. Leg IV: femur 0 (long
bristles), patella rl, tibia 8 (pi r2), metatarsus 21, tarsus
17. — Abdomen. Posterior lateral spinnerets, lengths: total
3.82, basal segment 1.36; middle 0.98; apical 1.48. Apical
segment width 0.41.

Female (paratype AMS KS3397). — Size. Carapace length
10.20, width 8.16. Abdomen length 10.88, width 9.18. —
Colour. Basic colour pattern. — Carapace. Much longer than

wide. Height 3.74; frontal width 6.39. Cephalic length 7.11.
Fovea narrow, procurved. Mid-dorsal cephalic setal row with
alternating large and small bristles which extend back to the
fovea. Anterior strial setae numerous. Anterolateral carapace
angle with weak bristles. — Eyes. Eye region weakly raised.
Eye group width 1.99. Diameters: AME 0.21, ALE 0.41, PLE
0.45, PME 0.27. Interdistances: AME-AME 0.27, AME-
ALE 0.20, ALE-PLE 0.20, PLE-P.M.E 0.12, PME-PME
0.81. Median ocular quadrangle length 0.78, anterior width
0.78, posterior width 1.34. — Chelicerae. Cheliceral groove
narrow, margins subparallel, divergent distally. Groove
length 3.26, middle width 0.47. Cheliceral teeth: 25 central,
occupying basal half of groove; 14 prolateral; 14 retrolateral.
— Labium. Much wider than long, anterior margin indented.
Length 1.60, width 2.28. Cuspules occupy anterior two-thirds
of labium. Labiosternal sigilla entire. — Sternum. Long,
ovoid. Length 6.12, width 4.01. Posterior sigilla short, ovoid:
length 0.48, width 0.27. — Palp. Spination: tibia 5, tarsus
11. Trichobothria: tibia p8 r8, tarsus 18. Tarsal claws with
8 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.48

4.15

5.64

4.84

2.72

24.83

2

6.58

3.88

4.73

4.32

2.65

22.11

3

5.81

3.16

3.50

4.35

2.92

19.74

4

7.11

3.84

5.03

5.54

4.01

25.53

Tibia I width 1.77. Metatarsus I proximal width 1.13.
Tarsal claw teeth legs I, II; superior 10, 11; inferior 3, 2.
Trichobothria legs I, II: tarsus 20, 16; metatarsus 20, 13;
tibia plO r9, p9 r9. — Leg spination. Leg I: femur 0, patella
0, tibia 2, metatarsus 12, tarsus 15. Leg II: femur pi, patella
0, tibia 4 (p2), metatarsus 11, tarsus 12. Leg III: femur 0,
patella pd3 rl, tibia 7 (p2 r2), metatarsus 17 (p6 rd2), tarsus
17. Leg IV: femur 0, patella 0, tibia 7 (r2), metatarsus 21
(p4 rd2), tarsus 18 — Abdomen. Posterior lateral spinnerets,
lengths: total 5.55; basal segment 2.04, middle 1.16, apical
2.35. Apical segment width 0.54. — Genitalia. Spermathecae
moderately elongate, gently curved, basally approximated.
Length 1.66, width 0.52.

Distribution. Southeastern New South Wales and north¬
eastern Victoria (Fig. 11).

Gray: Australian funnel-web spiders

307

Figure 14. Atrax sutherlandi, male ( A,D-K , AMS KS1102; B,C,
AMS KS1075): (A) palp, prolateral; (B) bulb; (Q embolus, distal;
(. D,E ) distal patella and tibia I— D, ventral, E, prolateral; (F,G)
metatarsus I— F, ventral, G, prolateral; (//,/) distal patella and
tibia II— H, ventral, I, prolateral; (J,K) metatarsus II— J, ventral, K,
prolateral. Scale lines: 0.3 mm, except B 1.0 mm and D,F,H 0.2 mm.

308

Records of the Australian Museum (2010) Vol. 62

Illawarra n.gen.

Type species Illawarra wisharti new species.

Diagnosis. Differs from other atracine species as follows—in
males, by having broad midventral spine row on male tarsi
I-IV (Fig. 3B); in females, by having a robust, aspinose leg
I with metatarsus partially fused to tarsus and tarsal claws
enlarged.

Description. With the characters of the Atracinae. Colour
light to dark brown with distinct abdominal chevrons.
Carapace longer than wide; relatively low with caput weakly
raised (CH/CW 0.37) and narrow frontally (CFW/CL 0.54).
Cheliceral paturon less robust than in other genera. (Figs.
16A,B; 17B,C). Cheliceral groove narrow, lateral margins
parallel; central tooth row short; basal; prolateral tooth row
may be incomplete; retrolateral row complete (Fig. 16C).
Labium short (LL/LW 0.70) (Fig. 16D), cuspule numbers
low to moderate (85-154). Apical segment of posterior
lateral spinnerets relatively short. Leg I more robust than
other legs, especially in females; female leg I without spines,
tarsal claws enlarged and metatarsus thickened proximally
(wedge-shaped) and partially fused to the tarsus (Fig. 17H,I).
Male tarsi highly spinose (43-93 spines), with two lateral
spine rows and a full midventral spine row (Fig. 3B). Female
tarsi with few spines. Male femora I, II with slender dorsal
spines (Fig. 16F,H); legs I, II without apophyses; tibia I, II
and metatarsus I, II spinose, spines slender and scattered
(Fig. 18D-J). Male palpal patella a little wider than femur.
Palpal bulb with a broad tegular area; groove between
tegulum and subtegulum sometimes wide, exposing middle
haematodocha.; embolus short, wide. (Figs. 161, 18B).
Female spermathecae two very short, distally broad sacs
(Fig. 17E).

Included species. Illawarra wisharti n.sp.

Distribution. The Illawarra region of southern New South
Wales.

Illawarra wisharti n.sp.

Figures 3B, 15-18; Tables 5, 34

Etymology. The species is named for Graeme Wishart,
Australian Museum Associate, idiopid spider researcher,
and collector of many mygalomorph spiders in the Illawarra
region.

Types. Holotype male: AMS KS5357, Nowra, New South Wales, 34°52'S
150°36'E, 17 March 1980. Paratypes (AMS). New South Wales. Males :
AMS KS15748, Willowvale near Gerringong, 34°45’S 150°47'E, 25 Mar
1985, G. Wishart; AMS KS920, Avondale, 34°30’S 150°45'E, March 1956
R.K. Hayes; AMS KS2721, Figtree, 34°25'S 150°50E, 16 March 1979,
J. Lewis; AMS KS919, AMS KS5355, Nowra, 34°52'S 150°36’E, 7 April
1980; AMS KS8921, Swan Lake, nr. Sussex Inlet, 35°10'S 150°34'E, 10
April 1982, A. McLaughlin; AMS KS15746, Willowvale, nr Gerringong,
34°52'S 150°36'E, 29 April 1977, G. Wishart; AMS KS16450, Unanderra,
34°27'S 150°50'E, 8 February 1979, J. Wafle; AMS KS5356, Bangalee Scout
Camp, Nowra area, 14 March 1980; Females : AMS KS30273, Willowvale,
4km W. of Gerringong, 34°35'S 149°37'E, November 1979, G. Wishart;
AMS KS2667, AMS KS4778, Nowra South, 34°54'S 150°35'E, March
1959; AMS KS 13781, Farmborough Heights, near Unanderra, 34°27'S
150°48'E, J. Wafle, 8 February 1979.

Figure 15. Collection records for Illawarra wisharti.

Other material examined. New South Wales (males): AMS KS921,
Willowvale, 4 km W. of Gerringong, 11 May 1977, G. Wishart; AMS
KS922, Willowvale, 4 km W. of Gerringong, 20 May 1977, G. Wishart; AMS
KS1448, Willowvale, 4 km W. of Gerringong, 5 April 1978, G. Wishart;
AMS KS918, Ulladulla, Ulladulla Ambulance Station; AMS KS10796,
Unanderra, January 1983, M. Senior.

Diagnosis. CL 6.77-10.13 (male). Male tarsi I-IV strongly
spinose (tarsus I with 43-93 spines), with a middle ventral
spine row as well as lateral ventral rows (Fig. 3B). Female
leg I robust, aspinose; metatarsus I thickened proximally and
partially fused with tarsus I, tarsal claws enlarged (Fig. 17H,I).

Figure 16. Illawarra wisharti, male: (A) cephalothorax and
chelicerae, dorsal; ( B ) cephalothorax and chelicerae, lateral; (C)
cheliceral groove teeth; ( D ) sternum, labium and maxilla; (E)
spinnerets; ( F ) leg I, prolateral; (G) palp, prolateral; (H) leg II,
prolateral; ( I) palpal organ. Scale lines 1 mm.

Gray: Australian funnel-web spiders

309

Table 5. Male morphological data— Illawarra wisharti (n = 12).

character

range

mean

character

range

mean

character

ratio

SD

CL

6.77-10.13

8.00

MtlS

28-76 (p0-l)

48

CW/CL

0.90

0.031

CW

6.36-9.36

7.18

TalS*

43-93

70

CH/CW*

0.37

0.02

CH

2.50-2.83

2.66

Ti2S

10-17 (p0-l)

13

CFW/CL*

0.54

0.021

CFW

3.94-5.17

4.32

STC2teeth*

6-10

8

CHGW/L*

0.11

0.016

ChGL

2.01-2.42

2.19

BulbL

1.73-2.11

1.89

SW/SL

0.82

0.021

ChGW

0.20-0.32

0.23

EmbL

1.02-1.22

1.12

LL/LW*

0.70

0.038

ChGCT

7-17

11

BulbW

0.76-0.98

0.84

PLSAPW/L

0.45

0.034

LL

0.86-1.18

0.98

EmbmidW

0.10-0.19

0.14

BulbW/EmbL*

0.76

0.047

LW

1.31-1.62

1.39

PalpTibL

2.36-2.92

2.65

EmbmidW/L

0.13

0.024

CUSP*

85-154

113

PalpTibW

1.10-1.46

1.24

PalpTibW/L

0.47

0.022

SL

3.60-4.79

4.28

PalpTibS

0-3

1

BulbL/TibL

0.72

0.027

SW

2.95-3.91

3.51

PalpPatS

0-3

1

PLSAPW

0.30-0.42

0.35

PalpFemS

0-1

1

PLSAPL

0.68-0.93

0.79

FelS*

3-16

9

PalS

0-4 (p0-l)

2

Male (holotype). — Size. Carapace length 7.54, width 6.75.
Abdomen length 7.88, width 5.90. — Colour. Basic colour
pattern, often brown overall, with distinct, narrow abdominal
chevrons. — Carapace. Longer than wide, weakly raised.
Height 2.62. Frontal width 4.06. Fovea procurved. Mid¬
dorsal cephalic setae few, almost reach fovea. Anterior
strial setae absent basally. Anterolateral angle of carapace
with weak hairs only. — -Eyes. Central eye region sessile
to slightly raised. Eye group width 1.62. Median ocular
quadrangle length 0.78, anterior width 0.75, posterior width
1.05. Diameters: AME 0.30, ALE 0.48, PLE 0.27, PME 0.24.
— Chelicerae. Anterodorsal paturon bristles strong, sparse.
Cheliceral groove very narrow, margins parallel. Groove
length 2.15, middle width 0.26. Cheliceral teeth: 10 central,
confined to base of groove; 7 prolateral; 13 retrolateral. —
Labium. Much wider than long, broadly indented apically.
Length 0.90, width 1.37. Labiostemal sigilla broad, complete.

Figure 17. Illawarra wisharti, female: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C)
cephalothorax and chelicerae, dorsal; ( D ) cheliceral groove teeth;
(. E ) spermathecae; (F) spinnerets; (G) leg III, prolateral, patella,
tibia, metatarsus. (H,I) Leg I, metatarsus and tarsus— H, prolateral;
I, dorsal. Scale lines 1 mm.

Cuspules c. 122, number low to moderate. — Sternum. Ovoid,
moderately broad. Length 4.12, width 3.32. Posterior sigilla
of moderate size, ovoid. — Palp. Tegular area almost as long
as wide, pear shaped. Middle haematodocha sometimes
exposed, in broad tegular-subtegilar groove. Embolus short,
moderately broad and gently curved, embolus weakly to
moderately twisted distally. Embolus weakly offset from
tegulum. Bulb length 1.80, width 0.80. Embolus length 1.08,
midwidth 0.12. Length of femur 2.96, patella 1.24, tibia 2.36.
Width of tibia 1.13. Spination: spines bristle-like; femur 2
sinuous, patella 1 sinuous, tibia 0. Sinuous bristles on femur
and patella, strong bristles on tibia. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.64

3.06

5.08

5.12

2.54

22.44

2

5.72

2.79

4.10

4.59

2.71

19.91

3

5.33

2.54

3.44

4.55

2.95

18.81

4

6.31

2.87

4.55

5.95

3.53

23.21

Tibia I width 1.44. Femora I—III with dorsal spines, bristle
like spines on femur IV. Metatarsus I slightly thickened
proximally, dorsal sigilla broad. Metatarsus II slightly bent
proximally, tibial and metatarsal spines scattered. Anterior
coxal hairs normal. Tarsal spines very numerous with a mid-
ventral spine row as well as lateral ventral spine rows. Tarsal
scopulae very weak to absent, distal metatarsal scopulae
absent. Tarsal claw teeth legs I, II: superior 8, 10; inferior 0,
2. Trichobothria legs I, II: tarsus 10, 8; metatarsus 9,10; tibia
p5 r5, p6 r5. — Leg spination. Leg I: femur 9 (d7 p2), patella
1, tibia23 (pi) metatarsus 56 (pi), tarsus 93. LegII: femur 7
(d6), patella p2, tibia 16 (pi), metatarsus 48, tarsus 79, Leg
III: femur 5, patella 9 (p6 r3), tibia 25 (p7 r6), metatarsus
38, tarsus 80. Leg IV: femur 0 (strong bristles), patella 4 (p2
r2), tibia 19 (p4 r6), metatarsus 42, tarsus 67. — Abdomen.
Numerous weak dorsal bristles, hair cover sparse. Posterior
lateral spinnerets with apical segment rather short. Lengths:
total 2.68; basal segment 1.12, middle 0.66, apical 0.90.
Apical segment width 0.35.

Female (paratype AMS KS2667). — Size. Carapace length
8.12, width 6.72. Abdomen length 9.79, width 7.48. —
Colour. Commonly brown in colour overall, abdominal
chevrons distinct. Otherwise colour pattern basic. —
Carapace. Longer than wide, moderately-weakly raised and

310

Records of the Australian Museum (2010) Vol. 62

Figure 18. Illawarra wisharti, male (. A,D-J , AMS KS15748; B,C, AMS KS5356): (A) palp, prolateral; ( B ) bulb; (C) embolus, distal; (D,E)
distal patella and tibia I— D, ventral, E, prolateral; (E) metatarsus I, ventral; ( G,H) distal patella and tibia II—G, ventral, 77, prolateral;
(7,7) metatarsus II—7, ventral, J, prolateral. Scale lines: 0.3 mm, except A 0.2 mm, and B 1.0 mm.

frontally narrow. Height 2.87, frontal width 4.72. Cephalic
length 6.19. Fovea strongly procurved, anterior margin
indented. Mid-dorsal cephalic setae long, in single row, may
or may not extend back to fovea. Anterior strial setae absent
basally. Anterolateral carapace angle with a few very weak

bristles. — Eyes. Central eye region slightly raised. Eye group
width 1.56. Diameters: AME 0.21, ALE 0.43, PLE 0.24,
PME 0.18 Interdistances: AME-AME 0.24, AME-ALE
0.18, ALE-PLE 0.22, PLE-PME 0.12, PME-PME 0.66.
Median ocular quadrangle length 0.52, anterior width 0.60,

Gray: Australian funnel-web spiders

311

posterior width 1.05. — Chelicerae. Cheliceral groove very
narrow, margins parallel. Groove length 2.14, middle width
0.32. Cheliceral teeth: 5 central, confined to basal third of
groove; 6 prolateral, a gap separating the distal tooth from
the rest; 13 retrolateral. — Labium. Wider than long, anterior
margin widely indented. Cuspules distributed in shallow
V-shaped band on anterior half of labium. Labiosternal
sigilla narrow. — Sternum. Broad. Length 4.29, width 3.70.
Posterior sigilla ovoid: length 0.72, width 0.38. — Palp.
Spination: tarsus 4. Trichobothria: tibia p5 r5, tarsus 5. Tarsal
claw long, slender with two basal teeth. — Legs. 1423. Leg
I robust, spines absent.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.66

3.28

4.43

3.44

2.00

18.81

2

4.43

2.62

2.79

2.61

1.74

14.19

3

4.13

2.17

2.11

2.61

1.89

12.91

4

5.91

2.79

3.32

3.77

2.38

18.17

Tibia I width 1.62. Metatarsus I wedge shaped, wide
proximally with a large dorsal sigilla, fused distally with
tarsus; proximal width 1.15. Tarsus I claws enlarged, strong,
hook-like. Coxal hairs unmodified, long. Tarsi I-IV with few
ventral spines, all lateral. Tarsal claw teeth few, basal—legs
I, II: superior claws 3, 2; inf erior 0, 0. Trichobothria legs I,
II: tarsus 8, 10: metatarsus 10, 8; tibia p6 r5, p5 r6. Short
setae interspersed with trichobothria. — Leg spination.
Leg I: spines absent. Leg II: femur 0, patella 0, tibia 0,
metatarsus 5, tarsus 4. Leg III: femur 0, patella p3, tibia 5
(p3), metatarsus 9 (p3), tarsus 8. Leg IV: femur 0, patella 0,
tibia 2, metatarsus 9 (p3), tarsus 8. — Abdomen. Posterior
lateral spinnerets with short apical segment. Lengths: total
2.38; basal segment 1.14, middle 0.52, apical 0.44. Apical
segment width 0.35. — Genitalia. Spermathecae very short,
only slightly longer than wide. Length 0.78, width 0.58.
Apical two-thirds bulbous.

Distribution. Illawarra region of New South Wales (Fig. 15).

Comments. Burrows are found in the ground litter layer or
under rocks. The entrance is at ground level and lacks silk
trip-lines. The burrow silk lining is very weak to absent.
The spiders are often noted to have a characteristic “ant¬
like” smell.

Hadronyche L. Koch

Hadronyche L. Koch, 1873: 463. Type species by monotypy
Hadronyche cerberea L. Koch, 1873. Ausserer, 1875:
143. Simon, 1892: 186; 1903: 969. Hogg, 1901: 274.
Rainbow, 1911: 121; 1913: 4; 1914: 252. Roewer, 1942:
208. Bonnet, 1957: 2070. Main, 1976:74; 1985:42. Gray,
1978:125. Gray, 1981: 265; 1988: 114. Raven, 1980:225
(synonymized with Atrax); 2000: 225. Goloboff, 1993: 6.
Hedin & Bond, 2006: 467. Platnick, 2010.

Styphlopis- Rainbow, 1913: 5. Type species by monotypy
Styphlopis insularis Rainbow, 1913. Roewer, 1942: 208.
Bonnet, 1958: 4193. Gray, 1978: 125 (synonymized
with Atrax ). Raven, 1980: 255. Main, 1985: 40. First
synonymized by Gray, 1988: 114.

Pseudatrax. -Rainbow, 1914: 260. Type species by
monotypy Pseudatrax moreaui Rainbow, 1914. Bonnet,
1958: 3807. Gray, 1978: 125 (synonymized with Atrax).
Raven, 1980: 255. Main, 1985: 40. First synonymized
by Gray, 1988: 114.

Anepsiada.-Rainbow & Pulleine, 1918: 167. Type species
by monotypy Anepsiada ventricosa Rainbow & Pulleine,
1918. Roewer, 1942: 208. Bonnet, 1955: 323. Gray,
1978: 125 (synonymized with Atrax). Gray, 1981: Raven,
1980: 255. Main, 1985: 40. First synonymized by Gray,
1988: 114.

Diagnosis. Differs from Atrax in tibia II being either
unmodified or having a blunt, rounded apophysis or
apophyseal swelling Differs from both Atrax and Illawarra
in having caput moderately to strongly raised and cheliceral
paturon more robust. Differs from Illawarra by male tarsi
having two instead of three ventral spine rows.

Description. With characters of the Atracinae. Carapace
broad, often not much longer than wide, or as wide;
caput moderately to strongly raised (CH/CW 0.40-0.53)
and wide frontally (CFW/CL 0.61-0.83). (Figs. 20B,E;
23B,G). Cheliceral paturon typically more robust than in
other Atracinae, thicker both dorsoventrally and laterally.
Cheliceral groove a wide to narrow V-shape with central
cheliceral teeth distributed along its full length, in a
staggered row (males—Fig. 20G) or one to several irregular
rows (females—Fig. 2ID); or, less commonly, central
teeth confined to a short, basal row (Fig. 97E). Labium
subquadrate, often almost as long as wide (LL/LW 0.86-
1.20) (Fig. 20C), sometimes shorter (LL/LW 0.64-0.79) (Fig.
18E). Sternal sigilla ovoid to narrowly elongate. Terminal
segment of posterior lateral spinnerets short to moderately
long digitiform. Male tibia II either unmodified with ventral
spines grouped or scattered (Fig. 43H,I); or with a rounded
spined ventral apophysis or low apophyseal swelling (Figs.
32H,I; 36H,I). Metatarsus II either sinuous (proximoventrally
concave) with a small, spined mid-ventral apophysis (Fig.
29K), or weakly sinuous to unmodified (Fig. 70K). Tibia
and metatarsus I ventrally spinose (tibial spines may extend
retrolaterally); metatarsus sometimes proximally thickened.
Male palpal patella large, often wider than the femur, or about
as wide. Palpal organ morphology variable but, compared
to Atrax spp., the tegular area of the bulb is often larger,
the embolus broader and shorter with the distal ejaculatory
groove narrow and slanting above the flange-like lower
margin (Figs. 25B,C; 32B,C). Spermathecal shape variable,
basic pattern a pair of relatively short, broad sacs.

Included species. 31 species They are placed here in four
species groups: the lamingtonensis group; the adelaidensis
group; the infensa group; and the cerberea group.

Distribution. Eastern Australia from northeast Queensland
to Tasmania and the Gulf Ranges region of South Australia.

Comments on genera and species groups. Both Atrax and
Illawarra are easily characterized genera with few species.
By contrast, Hadronyche is a diverse and speciose genus,
mainly characterized by the caput being relatively higher
and broader than in the other two genera. The separation
of Atrax and Hadronyche was supported by results from
electrophoretic studies (Gray, 1988), and more recently,
Raven (2000) has suggested that greater palpal patella/femur
width may be an apomorphy of Hadronyche. Another source
of differentiation between Hadronyche and both Atrax and
Illawarra comes from mass profile analyses of funnel-web

312

Records of the Australian Museum (2010) Vol. 62

spider venom peptides by Wilson (2001). This study found
that Atrax robustus and Illawarra wisharti had similar venom
profiles. However, the Hadronyche species sampled, viz.
Hadronyche versuta, H. cerberea and H. formidabilis, had
profiles that were similar to each other but quite different
from those observed for Atrax and Illawarra spp.

Of the four putative Hadronyche species groups
recognized here, only two seem likely to be natural groups—
the morphologically distinctive lamingtonensis group and the
isolated door building adelaidensis group. Both groups lack
leg II modifications and retain the short labium present in
Atrax and Illawarra. The short, basal central cheliceral tooth
row seen in these genera is also present in the lamingtonensis
group. However, in the adelaidensis group this is replaced by
a long central tooth row, a character shared with species in
the infensa and cerberea groups. The infensa and cerberea
groups are united by the longer labium (with a few exceptions
in the latter group). The infensa group is characterized by the
absence of dorsal femoral spines—these spines are present
in most, but variable in a few, cerberea group species. The
cerberea group is currently a diverse “dump” taxon for
species of uncertain affinities, and includes all Hadronyche
species with leg II modifications.

cerberea species group

Description. (Figs. 30-32, 41-43). Small to large sized
Atracinae. Male femora I, II usually with dorsal spines,
sometimes spines absent. Labium relatively long, (LL/LW
0.86-0.99), sometimes shorter (LL/LW 0.64-0.79) (see
comments below). Male tibia II shape and spination variable:
unmodified cylindrical to weakly sinuous, with a few ventral
spines clustered proximally or scattered; or sinuous with a
spinose, rounded apophysis or apophyseal swelling placed
mid-ventrally to proximally. Metatarsus II either cylindrical
to weakly sinuous and without an apophysis, or sinuous
(often ventrally concave proximally) with a mid-ventral
apophysis/apophyseal swelling. Male palp with embolus
often moderately short and broad, weakly curved and twisted.
Ratio of bulb length to palpal tibia length 0.64-0.77. Central
cheliceral tooth row long. Apical segment of posterior lateral
spinnerets short to long (PLSAPW/L 0.29-0.51).

Included species: H. versuta (Rainbow), H. formidabilis
(Rainbow), H. venenata (Hickman), H. cerberea L. Koch,
H. modesta (Simon), H. meridiana Hogg, H. pulvinator
(Hickman), H. marracoonda n.sp., H. jensenae n.sp., H.
tambo n.sp., H. monaro n.sp., H. emmalizae n.sp., H. alpina
n.sp., H. nimoola n.sp., H. mascordi n.sp.

Distribution. Cerberea group species are found mainly from
south of the Hunter River region in mid-eastern New South
Wales, into Victoria and Tasmania. Only H. formidabilis
occurs north of the Hunter River and into southeastern
Queensland.

Comments. The cerberea group is a diverse and problematic
“convenience” grouping of 15 species. These spiders
usually have dorsal spines on male femora I or II, but these
spines may be absent in some individuals of H. alpina, H.
versuta, H. tambo and H. nimoola. Eight of the 14 species
for which males are known have variably developed
apophyseal structures on tibia II and, usually, metatarsus

II. The remaining species, H. nimoola, H. mascordi, H.
tambo , and H. marracoonda , lack such structures, except
H. monaro which has a weak apophysis on metatarsus II.
Most cerberea group species have a relatively long labium
(a character also present in infensa group species). However,
a putatively related group of four species from Victoria and
Tasmania— H. modesta, H. meridiana, H. jensenae and H.
pulvinator —have a shorter labium (LL/LW 0.69-0.79). Gray
(1987,1988) placed them as a separate species group. A short
labium state also exists in H. mascordi (LL/LW males 0.64),
but this is probably associated with the partial fusion of the
labium with the sternum in this species. Excluding these 5
species, the remainder have a male LL/LW almost identical
to that of the infensa group.

Hadronyche cerberea L. Koch

Figures 19-22; Tables 6, 34

Hadronyche cerberea L. Koch, 1873: 463. Karsch, 1878:

798. Main, 1985: 43. Gray, 1988: 114.

Types. Syntypes: females from Sydney region, New South Wales, lost (see
below). Neotype, here designated as follows: one male, registered AMS
KS6873 in the research collection of the Australian Museum, Sydney; with
label-data: Blackwall, nearWoy Woy, New South Wales, 33°30'S 151°20'E,
R. McDonald, 10 June 1973, from paperbark tree.

Other material examined. New South Wales (males): AMS KS6874,
Antonio, 21 September 1973, R. McDonald; AMS KS4597, Avalon, 15
January 1980; AMS KS8729, Awabakal Nature Reserve, near Newcastle,
7 August 1981, G. Anderson; AMS KS4104, Bago Forest way, nearBatlow,
5 November 1979, AMS KS. Kovacs; AMS KS6871, Bilgola, 18 February
1974, B. Mercer; AMS KS6872, Bilgola Heights, 5 February 1978; AMS
KS2261, Blackheath, November 1978; AMS KS4782, Empire Bay, near
Gosford, 18 March 1978, L. Abra; AMS KS8568, Gosford, 12 April 1979;
AMS KS 13367, Gosford area; AMS KS1347, Kuringai Chase National
Park, 1978, M. Arena; AMS KS 14058, Mudgee, 13 February 1984, W.
Bennett; AMS KS 13346, Mullion Creek, near Orange, September 1983,
D. Ogilvy; AMS KS 16587, Olinda, near Rylstone; AMS KS5850, Orange;
AMS KS5900, Springwood, 21 February 1930; AMS KS4485, Swansea,
4 February 1980, G. Anderson; AMS KS4783, Terrigal, 18 March 1978,
L. Abra; Terrigal, 10 January 1959, T.E. Dence; AMS KS8923, Toukley,
4 April 1982, R. Hay; AMS KS5901, Valley Heights, 21 October 1921,
R.S. Thornthwaite; AMS KS 10784, Walang, near Bathurst, 23 March
1983, A. Boesen; AMS KS5902, Wentworth Falls, 27 April 1943. New

Figure 19. Collection records for Hadronyche cerberea.

Gray: Australian funnel-web spiders

313

South Wales (females): AMS KS6874, Antonio/Rydal, 21 September
1973, R. McDonald; AMS KS3438, Baulkham Hills, 27 March 1979, K.
Moore; AMS KS6870, Bobbin Head, 18 March 1972, A.B. Rose; AMS
KS7296, Bombala, 16 April 1981; AMS KS2331, Burrawang, May 1977, R.
Merrick; Collaroy Plateau, February 1975, A. Goodridge; AMS KS13978,
Dooralong, 24 February 1984, L. Abra; AMS KS3461, Gosford, May 1979,
D. Jones; AMS KS4606, Highfields, 7 February 1980, G. Anderson; AMS
KS5362, Monga, 1980, D. Rowell; Mount Fairy, 25 November 1979; AMS
KS5903, Newcastle area, 1973; AMS KS5842, Pearl Beach, 27 July 1980,
V. Serventy; AMS KS5976, AMS KS6217, Wamberal, 6 March 1980, M.
Gray & C. Horseman.

Comments on syntype loss and erection of neotype.

with the inadequacy of the original description based on
female characters and loss of the type material, the identity
of Hadronyche cerberea L. Koch, the type species of the
genus, has long been conjectural. The syntype female
specimens, labelled “Sydney”, were formerly housed in
the Staatlische Museum, Stuttgart, West Germany. They
were lost during World War II (pers. comm., Dr M. Janus,
formerly Hauptconservater, Staatlische Museum, Stuttgart).
Determination of the identity of H. cerberea was facilitated
by the presence of only two common atracine species in the
Sydney region. These are Atrax robustus, the Sydney funnel-
web spider, and a then unidentified tree dwelling species
of Hadronyche, common in forest habitats of southeastern
New South Wales, including the Sydney region (where
wood cutting and clearing activities could have facilitated
encounters with this spider). The description of the female

Figure 20. Hadronyche cerberea, male: (A) spinnerets; ( B )
cephalothorax and chelicerae, lateral; (C) sternum, labium and
maxilla; (D) palp, prolateral; (E) cephalothorax and chelicerae,
dorsal; ( F ) leg II, prolateral; (G) cheliceral groove teeth; (H)
sternum, anterolateral bristles. Scale lines 1 mm.

spider given by Koch (1873) indicated that he was dealing
with a species of Hadronyche. Consequently, the decision
was made to identify Koch’s H. cerberea with the tree
dwelling Sydney Hadronyche species (Gray, 1981, 1988).

Designation of a neotype will stabilise the taxonomic
status of Hadronyche cerberea L. Koch as the type species
of the genus. The presence of a raised caput, a long labium
and a bluntly rounded tibia II apophysis refers the species
unequivocally to the genus Hadronyche as here defined.
A diagnosis and description of the species is given below.
The original type locality was given as “Sydney” but this
locality could refer to anywhere from the Hawkesbury River/
Central Coast region in the north to the Illawarra region in the
south, where these spiders are common. The neotype male
specimen is held in the research collections of the Australian
Museum. Sydney.

Diagnosis. CL 7.18-8.77 (male). Tree-dwelling spiders in
sclerophyll forests. Differs from most species by presence (in
both sexes) of thorn-like setae on coxae I, II and a bristle-like
tuft of setae at anterolateral sternal angles (Figs. 20H, 21H);
from H. formidabilis by smaller size of tibia II apophysis;
and from H. emmalizae by embolus almost straight (Fig.
22B), not strongly curved.

Neotype male Size. Carapace length 8.18, width 7.83.
Abdomen length 7.87, width 6.97. — Colour. Basic colour

Figure 21. Hadronyche cerberea, female: (A) spermathecae; ( B ) ceph¬
alothorax and chelicerae, lateral; (C) cephalothorax and chelicerae,
dorsal; (D) chehceral groove teeth; (E) sternum, labium and maxilla;
(F) leg III, prolateral, patella, tibia, metatarsus; (G) spinnerets; (H)
coxa I, anteromedial thom-like setae. Scale lines 1 mm.

314

Records of the Australian Museum (2010) Vol. 62

Figure 22. Hadronyche cerberea , male (A,D,F-H,J,K, AMS KS4783; B, C,E,I, AMS KS16587): (A) palp, prolateral; (B) bulb; (Q embolus,
distal; ( D,E ) distal tibia I, retrolateral— D, small spines (Terrigal); E, large spines (Rylstone); (EG) metatarsus I— F, ventral, G, prolateral;
(FI,I) distal patella and tibia II, ventral— H, Terrigal; /, Rylstone; ( J) tibia II, prolateral; (K) metatarsus II, prolateral. Scale lines: 0.3 mm,
except B 1.0 mm and E,1 0.2 mm.

Gray: Australian funnel-web spiders

315

Table 6. Male morphological data— Hadronyche cerberea (n = 12).

character

range

mean

character

range

mean

character

ratio

SD

CL

7.18-8.77

8.17

MtlS

26-46

34

CW/CL

0.95

0.034

CW

6.84-8.39

7.77

TalS

8-17

10

CH/CW

0.46

0.018

CH

3.12-3.94

3.54

Ti2S

15-39 (p0-l)

26

CFW/CL

0.71

0.02

CFW

5.25-6.31

5.81

STC2teeth

10-14

12

CHGW/L

0.21

0.022

ChGL

2.00-2.70

2.33

BulbL

2.66-3.07

2.82

SW/SL

0.75

0.031

ChGW

0.37-0.58

0.43

EmbL

1.64-2.11

1.86

LL/LW

0.97

0.08

ChGCT

15-26

20

BulbW

1.01-1.24

1.09

PLSAPW/L

0.42

0.047

LL

1.28-1.89

1.51

EmbmidW

0.18-0.24

0.20

BulbW/EmbL

0.60

0.056

LW

1.36-1.74

1.57

PalpTibL

4.02^.82

4.37

EmbmidW/L

0.10

0.014

CUSP

194-293

252

PalpTibW

1.48-1.93

1.78

PalpTibW/L

0.41

0.027

SL

4.92-5.81

5.26

PalpTibS

0-4

2

BulbL/TibL

0.64

0.025

SW

3.63-4.35

3.95

PalpPatS

0

—

PLSAPW

0.34-0.68

0.50

PalpFemS

0-3

1

PLSAPL

1.00-1.42

1.18

FelS

2-12

7

PalS

3-14 (pi—5)

7

pattern. Abdomen colour variable, light maroon brown
to dark brown. — Carapace. Slightly longer than wide,
moderately raised. Height 3.65. Frontal width 5.99. Fovea
procurved. Mid-dorsal cephalic setae reach fovea. Anterior
strial setae present. Anterolateral angle of carapace with
strong bristles. — Eyes. On slight tubercle. Eye group width
2.05. Median ocular quadrangle length 0.77, anterior width
1.10, posterior width 1.35. Eye diameters: AME 0.32, ALE
0.47, PLE 0.32, PME 0.27. — Chelicerae. Groove broad,
margins diverging distally, length 2.36, middle width 0.56.
Cheliceral teeth: 17 central, running full length of groove, 10
prolateral; 9 retrolateral. — Labium. Almost as long as wide;
shape as in female. Length 1.64, width 1.66. Labiosternal
sigilla entire. Cuspules c. 223, number moderate to high.
— Sternum. Ovoid to broadly ovoid. Length 5.37, width
4.06. Posterior sigilla large, broadly elongate. Bristle-like
hairs grouped in anterolateral angles. — Palp. Tegular area
slightly longer than wide. Embolus shaft weakly curved,
of medium length and taper. Distal embolus weakly to
moderately twisted. Embolus weakly offset from tegulum.
Bulb length 2.86, width 1.09. Embolus length 1.88, midwidth
0.19. Length of femur 4.12, patella 1.72, tibia 4.30. Width
of tibia 1.79. Spination: femur 3, tibia 2. Sinuous bristles on
distal femur. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.93

3.24

6.03

5.66

2.95

24.81

2

6.72

3.44

5.74

5.74

2.99

24.63

3

7.28

3.08

5.00

5.41

3.03

23.80

4

7.46

3.40

6.40

6.48

3.36

27.10

Tibia I width 1.23. All femora with dorsal spines. Metatarsus
I proximally thickened, larger ventral spines grouped
proximally. Ventral tibia II with centrally placed, low, spinose
apophyseal swelling; distoventral tibia concave and without
spines (except apically). Metatarsus II moderately to weakly
sinuous with small central, spined apophysis. Coxae I and II
with basally thickened and thorn-like setae on anteromedial
surface. Tarsal and distal metatarsal scopulae weak on legs
I and II, well developed on legs III and IV. Tarsal claw teeth
legs I, II: superior 12, 11; inferior 4. Trichobothria legs I,
II: tarsus 14, 14; metatarsus 12,13; tibia p6 r6, p5 r7. — Leg
spination. Leg I: femur 12(d9p3), patella 6(p5 vl), tibia
41(p2), metatarsus 29, tarsus 8. Leg II: femur 6(d5pl), patella

p3, tibia 19(pl), metatarsus 20, tarsus 11. Leg III: femur d8,
patella 5(p4 rl), tibia 9(p2 r3), metatarsus 26(pd3 rd3), tarsus
16. Leg IV: femur d6, patella 2(pl rl), tibia 6(r2), metatarsus
19, tarsus 13. — Abdomen. Posterior lateral spinnerets with
moderately short apical segment. Lengths: total 3.14; basal
segment 1.44, middle 0.70, apical 1.00. Apical width 0.46.

Female (AMS KS6869, Canalack Gully, Hornsby Heights,
33°39'E 151°05'S, New South Wales, 19 Lebruary 1973, M.
Gray and M. Robinson, from tree web). — Size. Carapace
length 9.95, width 9.93. Abdomen length 14.62, width
12.24. — Colour. Basic colour pattern except that abdomen
is often weakly pigmented dorsally (light maroon brown
colour) and dark pigment is weak or absent ventrally (light
maroon colour). — Carapace. Broad, about as long as wide,
cephalic area strongly raised. Height 4.52; frontal width
8.67. Cephalic length 6.66. Lovea strongly procurved,
anterior margin slightly indented. Mid-dorsal cephalic setae
numerous, reach back to fovea. Anterior strial setae present.
Anterolateral carapace angle with numerous bristles. — Eyes.
Eye region raised centrally. Eye group width 2.76. Diameters:
AME 0.34, ALE 0.54, PLE 0.48, PME 0.29. Interdistances.
AME-AME 0.41, AME-ALE 0.43, ALE-PLE 0.37, PLE-
PME 0.19, PME-PME 1.87. Median ocular quadrangle
length 0.85, anterior width 1.16, posterior width 1.83. —
Chelicerae. Groove wide, margins diverge distally; groove
length 3.26, middle width 1.02. Cheliceral teeth: 45 central
in 2-3 irregular rows occupying full length of groove; 11
prolateral; 12 retrolateral. — Labium. Long; as long as wide
or longer, sides sloping to a narrower rounded apex, anterior
margin not indented apically. Length 2.38, width 2.31.
Cuspules grouped on anterior two-thirds. Labiosternal sigilla
broad, entire. — Sternum. Ovoid. Length 6.90, width 5.10.
Bristle-like hairs grouped in anterolateral angles. Posterior
sigilla elongate: length 1.43, width 0.54. —Palp. Spination:
tibia 3, tarsus 6. Trichobothria: tibia p8 r8, tarsus 13. Tarsal
claw with 7 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.45

4.01

6.00

5.29

2.99

25.74

2

6.83

3.77

5.30

4.98

2.97

23.85

3

6.04

3.26

4.18

4.69

2.99

21.16

4

7.72

3.98

5.81

5.81

3.13

26.45

316

Records of the Australian Museum (2010) Vol. 62

Tibia / width 1.73. Metatarsus I proximal width 1.14. Coxae
I, II with basally thickened thorn-like setae anteromedially.
Tarsal claw teeth legs I, II: superior 10, 10; inferior 4, 4.
Trichobothria legs I, II: tarsus 17, 15; metatarsus 18, 16;
tibia p8 r7, p8 r7. —Leg spination. Leg I: femur 0, patella
0, tibia 3, metatarsus 13, tarsus 8. Leg II: femur 0, patella
0, tibia 3 (pi), metatarsus 13, tarsus 9. Leg III: femur rd2,
patella p5 rl, tibia 7(p3 r2), metatarsus 15 (p3 r2), tarsus 8.
Leg IV: femur 0, patella 0, tibia 5 (r3), metatarsus 11, tarsus
10. — Abdomen. Posterior lateral spinnerets with short apical
segment. Lengths: total 4.05; basal segment 1.60, middle
1.02, apical 1.43. Apical segment width 0.65. — Genitalia.
Spermathecae short, digitiform, apical third slightly narrower
than basal region; basal separation equivalent to spermatheca
width. Spermatheca length 1.09, width 0.63.

Distribution. Southeastern coast and highlands in New
South Wales as far north as the Hunter River, Central Coast
region (Fig. 19).

Comments. Thorn-like setae (short, strongly thickened
basally) on coxae I, II (Fig. 21H) and bristle-like tuft of
anterolateral sternal hairs (Fig. 20H) are well developed in
both sexes. Thorn-like setae are also present in males of H.
formidabilis and, more weakly, in some H. tambo and H.
emmalizae males.

Male tibia I retroventral spines are sometimes reduced in
size (Fig. 22D). The extent of such variation in this widely
distributed species needs further examination.

Figure 23. Hadronyche versuta, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) leg II,
prolateral (NB, dorsal femoral spines present or absent); ( D )
cheliceral groove teeth; (E) spinnerets; (F) palp, prolateral; (G)
cephalothorax and chelicerae, dorsal. Scale lines 1 mm.

Biology. This is a tree-dwelling species that is widely
distributed in open forest habitats. It is often associated with
rough-barked trees such as Melaleuca, Banksia, Casuarina
and some eucalypts (Fig. ID). Burrows are associated with
tree-trunk fractures, rot-holes and borer holes, with the
entrance sheet and trip-lines often disguised by bark/leaf
detritus and frass.

Hadronyche versuta (Rainbow)

Figures 1, 23-26; Tables 7, 34

Atrax versuta Rainbow, 1914: 253.

Aname bicolor.- Rainbow, 1914: 233.

Pseudatrax moreaui- Rainbow, 1914: 261.

Atrax bicolor.- Hickman, 1964: 107 (transferred from Aname
after Rainbow & Pulleine, 1918: 139). Main, 1985: 40.
First synonymized by Gray, 1988, 114.

Atrax moreaui. -Main, 1985: 40 (transferred from Pseud¬
atrax). First synonymized by Gray, 1988, 114.

Hadronyche versuta.-Gmy, 1988: 114 (transferred from
Atrax).

Types. Two subadult syntypes: AMS KS969 (old catalogue number
K12907). Jenolan, New South Wales, J. Wiburd, 1901.

Other material examined. New South Wales (males): AMS KS844,
Antonio Creek, 28 April 1973, R. McDonald; AMS KS1008, Antonio,
26 December 1974, R. McDonald; AMS KS999, Blackheath, 11

Figure 24. Hadronyche versuta, female: (A) cheliceral groove
teeth; ( B ) cephalothorax and chelicerae, lateral; (C) sternum,
labium and maxilla; ( D ) cephalothorax and chelicerae, dorsal; (E)
leg III, prolateral, patella, tibia, metatarsus; ( F) spinnerets; (G)
spermathecae. Scale lines 1 mm.

Gray: Australian funnel-web spiders

317

Table 7. Male morphological data —Hadronyche versuta (n = 15).

character

range

mean

character

range

mean

character

ratio

SD

CL

7.84-10.39

9.16

MtlS

19-39

29

CW/CL

0.96

0.032

CW

8.13-9.84

8.83

TalS

8-26

14

CH/CW

0.44

0.027

CH

3.49-4.42

3.90

Ti2S*

37-75(p0-l)

48

CFW/CL

0.70

0.039

CFW

5.95-7.17

6.43

STC2teeth

11-15

13

CHGW/L

0.22

0.019

ChGL

2.20-2.87

2.54

BulbL

2.88-3.44

3.25

SW/SL

0.72

0.025

ChGW

0.37-0.67

0.55

EmbL

1.84-2.40

2.21

LL/LW

0.98

0.055

ChGCT

16-32

22

BulbW

1.15-1.34

1.26

PLSAPW/L*

0.49

0.071

LL

1.56-1.88

1.70

EmbmidW

0.24-0.41

0.30

BulbW/EmbL

0.57

0.031

LW

1.60-1.84

1.74

PalpTibL

3.79-4.90

4.51

EmbmidW/L

0.14

0.016

CUSP

227-413

292

PalpTibW

1.82-2.24

1.98

PalpTibW/L

0.44

0.023

SL

5.37-6.39

5.90

PalpTibS*

4-13

8

BulbL/TibL

0.72

0.029

SW

3.90-4.76

4.28

PalpPatS

0-4

1

PLSAPW

0.39-0.68

0.54

PalpFemS

2-7

4

PLSAPL

0.82-1.49

1.11

FelS*

0-3

1

PalS

4-14 (pi—3)

9

January 1930, Dr V.K. Spence; AMS KS4431, Bomaderry, 19 January
1980, G. Wishart; AMS KS13790, Bungonia Heights, near Goulburn,
21 January 1984; AMS KS1179, Callala Beach via Nowra, near Jervis
Bay, January 1959, J. Coppin; AMS KS3607, Capertee, 25 July 1973,
R. McDonald; AMS KS1182, Erowal Bay, Georges Basin, 3 December
1950, E.C. Hammond; AMS KS4620, Gerroa, 24 December 1979, D.
Tidmarsh; AMS KS2433, Goulburn, 24 January 1979, T. Smith; AMS
KS8535, Goulburn, December 1981; AMS KS2962, Gulgong, May
1979, Gillman; AMS KS5305, Hargraves, near Mudgee, 4 June 1980,
C. Knott; AMS KS997, Hazelbrook, February 1976; AMS KS1177,
Honeymoon Bay, near Jervis Bay, 25 December 1954, R Harvey;
AMS KS846, AMS KS857-8, Jenolan, February 1932, J.C. Wiburd;
AMS KS1178, Jervis Bay, October 1958, K. Sanders; AMS KS860,
Kanangra-Boyd National Park, 21 May 1971, M. Gray; AMS KS1003,
Mount Wiburd, Kanangra-Boyd National Park, 9 January 1973, G.S.
Hunt; AMS KS1200, Katoomba, 24 January 1950, C. Batty; AMS
KS851, Echo Point, Katoomba, 14 February 1953, A. Henry; AMS
KS843, Leura, December 1972, Maguire; AMS KS856, Lithgow area,
1961, Lithgow Pharmacy; AMS KS1007, Mittagong, N.S.W., R.N.
Lochhead; AMS KS996, Mount Irvine, 1959, P.G. Yalder; AMS KS2493,
Mount Wilson, 12 January 1979-7 February 1979, C. Horseman; AMS
KS1174, Nowra, February 1975; AMS KS3357-8, Nowra, 12 January
1979, Sister Green; AMS KS7690, Olinda, 28 May 1981, J. Kirk; AMS
KS1176, Shoalhaven Heads, 7 April 1969, B. Virtue; AMS KS4123,
Springwood, 1 February 1979, Mascord; AMS KS6840, Springwood, 1
February 1979, Mascord; AMS KS 1001, Wentworth falls, January 1951,
H. Holland; AMS KS1002, Wentworth falls, January 1951, H. Holland;
AMS KS1114, Rydal, November 1973, R. McDonald; AMS KS1118,
Mittagong, 12 February 1977, B. Telfer; AMS KS1014, Kandos, June
1968, G. Daniels; AMS KS1016, Mudgee, 29 April 1974, Ambulance
Stn.; AMS KS1855, Peel, near Bathurst, 20 May 1974, P. Wolfe; AMS
KS1010, Rylstone, 24 km N. on Bylong Road; AMS KS1013, Sodwalls,
29 January 1973, R. McDonald; AMS KS4411, Turondale, Apr 1979.
New South Wales (females): AMS KS1671, Bilpin; AMS KS4776,
Blackheath, 14 March 1959, Heywood; AMS KS4664, Bundanoon,
February 1980; AMS KS1179, Callala Beach, January 1959, J. Coppin;
AMS KS8296, Culburra, near Nowra, September 1966, R. Esgate;
AMS KS6281, Currarong, 6.x. 1980; AMS KS1207, Faulconbridge, 7
December 1976, C. Watson; AMS KS793, Gerroa, 28 March 1977, G.
Wishart; AMS KS 12625, Goulburn, 16 March 1983; AMS KS6732,
Goulburn, January 1981, Ambulance Stn.; AMS KS1115, Hazelbrook,
8 October 1973, K. Lay; AMS KS9964, Jenolan, N.S.W., December
1979; AMS KS1862, Kanangra-Boyd National Park, 26 November
1974, M. Gregg; AMS KS3385, Kangaroo Valley, 1 July 1979, N.L.
Boomer; AMS KS1859, Katoomba, N.S.W., February 1927, F. Walford;
AMS KS1860, Kurrajong Heights, 4 March 1973, R. McDonald; AMS
KS4127, Lake Conjola, 29 April 1979, R. Mascord; AMS KS5926,
Marulan, September 1980; AMS KS1117, Medlow Bath, 12 October
1945; AMS KS6216, Moss Vale, 6 October 1980; AMS KS10677,
Nowra, 20 January 1983, W. Lamond; AMS KS5332, Shoalhaven Heads,
11 June 1980, H. Pepper; AMS KS2262, Sussex Inlet, 29,xi.l978, S.
Prince; AMS KS1672, Wentworth Falls, 31 March 1936, K.K. Graham;
AMS KS3814, Mittagong, 21 October 1979, B. Day; AMS KS1012.

Diagnosis. CL 7.84-10.39 (male). Differs from most species
by male tibia II having spinose rounded apophysis with
ventral spines always extending onto distal tibia (Fig. 25H,I);
from H. emmalizae by having embolus almost straight; from

H. formidabilis by having a less prominent tibia II apophysis
and shorter spinnerets (Fig. 25C,E; Table 7).

Male (AMS KS4477), Lithgow, New South Wales, 33°28’S
150°09'E, J.W. Rayner, 18 January 1980). — Size. Carapace
length 9.51, width 9.14. Abdomen length 10.58, width
6.97. — Colour. Basic colour pattern. Dorsum of abdomen
usually with definite paler patch anteriorly flanked by small
sigilla. — Carapace. Slightly longer than wide, moderately
raised. Height 3.80. Frontal width 6.81. Fovea procurved.
Mid-dorsal cephalic setae reach fovea. Anterior strial setae
present. Anterolateral angle of carapace with strong bristles.
— Eyes. Sessile. Eye group width 2.39. Median ocular
quadrangle length 0.86, anterior width 1.07, posterior width

I. 58. Diameters: AME 0.33, ALE 0.54, PLE 0.47, PME 0.29.
— Chelicerae. Cheliceral groove of moderate length and
width, margins diverging distally. Groove length 2.72, middle
width 0.58. Cheliceral teeth: 21 central, running full length of
groove; 11 prolateral; 9 retrolateral. — Labium. About as long
as wide. Length 1.80, width 1.78. Labiosternal sigilla entire.
Cuspules c. 288, number moderate to high. — Sternum.
Ovoid. Length 5.90, width 4.27. Posterior sigilla large, broad,
ovoid. — Palp. Tegular area wider than long. Embolus shaft
broad, weakly tapered and curved. Distal embolus weakly to
moderately twisted. Embolus slightly offset from tegulum.
Bulb length 3.32, width 1.29. Embolus length 2.32, midwidth
0.32. Length of femur 4.24, patella 2.12, tibia 4.87. Width of
tibia 1.99. Spination: femur 2, patella 1, tibia 6. Distal femur
with sinuous bristles. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.75

3.67

6.28

6.22

3.75

27.67

2

7.41

3.66

5.85

6.20

3.69

26.81

3

6.56

3.36

5.33

5.67

3.73

24.65

4

8.28

3.36

6.88

6.97

4.10

29.59

Tibia I width 1.51. Femora I and II with few, weak dorsal
spines, occasionally none. Metatarsus I proximally weakly
thickened, larger ventral spines grouped proximally. Tibia
II apophysis a low, blunt swelling, centrally placed; ventral

318

Records of the Australian Museum (2010) Vol. 62

Figure 25. Hadronyche versuta, male (. A,D-K , AMS KS851; B,C,
AMS KS3358): (A) palp, prolateral; ( B ) bulb; (C) embolus, distal;
(D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G)
metatarsus I— F, ventral, G, prolateral; (//,/) distal patella and tibia
II—77, ventral, 7, prolateral; (J,K) metatarsus II— J, ventral, K,
prolateral. Scale lines: 0.3 mm, except A,E 1.0 mm and 7 0.2 mm.

spines numerous, larger spines grouped upon apophysis,
shorter spines in weakly excavated distoventral region.
Metatarsus II strongly sinuous with a prominent spined
apophysis. Coxal hairs normal. Scopulae weak or lacking

on first and second tarsi and distal metatarsi; well developed
on tarsi and distal metatarsi III and IV. Tarsal claw teeth legs
I, II: superior 13, 13; inferior 3, 3. Trichobothria legs I, II;
tarsus 14, 11; metatarsus 18, 18; tibia p7 r6, p8 r8. —Leg

Gray: Australian funnel-web spiders

319

Figure 26. Collection records for Hadronyche versuta (squares)
and H. emmalizae (circles).

spination. Leg I: femur dl, patella 14 (p3 vl 1), tibia 56 (pi),
metatarsus 25, tarsus 12. Leg II: femur dl, patella 7 (p3
r4), tibia 45 (pi), metatarsus 23, tarsus 18. Leg III: femur
0, patella 9 (p8 vl), tibia 13 (p4 r2), metatarsus 30, tarsus
17. Leg IV: femur 0, patella 0, tibia 7 (rl), metatarsus 21,
tarsus 21. — Abdomen. Numerous strong bristles dorsally.
Posterior lateral spinnerets with rather short apical segment.
Lengths: total 4.29; basal segment 1.76, middle 1.04, apical
1.49. Apical segment width 0.65.

Female (AMS KS1233), Mt. Wiburd, Kanangra-Boyd
National Park, New South Wales, 33°49’S 150°01'E, G.S.
Hunt, 10 January 1973). — Size. Carapace length 11.18,
width 11.20. Abdomen length 16.12, width 12.72. — Colour.
Basic colour pattern. Abdomen usually dark maroon brown,
sometimes paler. — Carapace. About as wide as long,
strongly raised, broad frontally. Height 5.54; frontal width
10.29. Cephalic length 9.86. Fovea strongly procurved,
anterior margin slightly indented. Mid-dorsal cephalic setae
numerous, reach back to fovea. Anterior strial setae present.
Antero-lateral carapace angle with numerous bristles. —
Eyes. Ocular area sessile or slightly raised centrally. Eye
group width 3.60. Diameters: AME 0.37, ALE 0.72, PLE
0.60, PME 0.38. Interdistances: AME-AME 0.66, AME-
ALE 0.50, ALE-PLE 0.46, PLE-PME 0.25, PME-PME
1.61. Median ocular quadrangle length 0.97, anterior width
1.42, posterior width 2.36. — Chelicerae. Cheliceral groove
margins diverge distally, groove wide. Groove length 3.60,
middle width 1.06. Cheliceral teeth: 66 central, in several
irregular rows, occupying full length of groove; 11 prolateral;
10 retrolateral. — Labium. Long; about as long as wide,
rounded, anterior margin not indented. Length 2.90, width
2.88. Cuspules grouped on anterior half. Labiosternal sigilla
entire. — Sternum. Long. Length 8.46, width 5.74. Posterior
sigilla elongate, length 1.64, width 0.76. — Palp. Spination:
tibia 7, tarsus 11. Trichobothria: tibia p8 r8, tarsus 13. Tarsal
claw with 4 teeth. — Legs. 1423 or 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.69

5.17

8.35

5.78

3.36

31.35

2

7.81

4.76

6.19

5.45

3.28

27.49

3

6.72

3.53

4.33

4.84

3.16

22.58

4

8.94

4.92

6.12

5.07

3.31

28.36

Tibia V width 2.00. Metatarsus I proximal width 1.31. Coxa

I, II with weakly basally thickened or unmodified setae
frontally. Tarsal claw teeth legs I, II: superior 9, 9; inferior

4, 4. Trichobothria legs 1, II: tarsus 18, 18; metatarsus 18,
17; tibia plO r9, p8 r8. — Leg spination. Leg I: femur 0,
patella 0, tibia 7, metatarsus 11, tarsus 8. Leg II: femur 0,
patella p2, tibia 5, metatarsus 16, tarsus 13. Leg III: femur
0, patella p8, tibia 10 (p5 r2), metatarsus 20 (p7 r4), tarsus
12. Leg IV: femur 0, patella 0, tibia 1, metatarsus 12, tarsus
19. — Abdomen. Posterior lateral spinnerets stout with short,
thick apical segment. Lengths: total 5.11; basal segment
2.34, middle 1.23, apical 1.54. Apical segment width
0.94. — Genitalia. Spermathecae large, digitiform, slight
constriction separating apical third from basal two thirds.
Length 1.66, width 0.67.

Distribution. Blue Mountains to Illawarra region of New
South Wales (Fig. 26). Specimens probably attributable to
this species have been collected from as far south as the Eden
region near the NSW border.

Notes. Retreat burrows are often built within rotting logs and
stumps with entrance silk sometimes disguised by rotting
wood particles and prey remains.

Hadronyche emmalizae n.sp.

Figures 26-29; Tables 8, 34

Etymology. The species is named for my daughter, Emma
Elizabeth Jensen Gray.

Types. Holotype male: AMS KS13464, Geehi Rest Area, Swampy Plains
River, Kosciuszko National Park, New South Wales, 36°22'S 148°10'E,
A.B. Rose, 30 November 1983. Paratypes (AMS). New South Wales.
Males : AMS KS4114, Bago State Forest, 35°41'S 148°09'E, 20 November
1979, L. Hedt; AMS KS6648, Geehi, 36°23'S 148°10’E, 27 January 1981,

J. Robson; KS 113352, Khancoban, 36°13'S 148°08’E, 4 December 1980,

5. Bates. Females : AMS KS 10675, Paddy’s River Dam, Bago State Forest,
35°41'S 148°09E, L. Hedt, 5 November 1983; AMS KS4096, Bago, 35°37'S
147°59'E, 1 January 1979, L. Hedt; AMS KS4256-7, Bago State Forest,
35°41'S 148°09'E, 11 December 1979, F. Hedt.

Diagnosis. CL 8.06-9.78 (male). Differs from other species
in having both a rounded apophysis on tibia II and a strongly
curved embolus (Fig. 29B).

Male (holotype). — Size. Carapace length 9.78, width 8.91.
Abdomen length 7.68, width 7.14. — Colour. Basic colour
pattern. — Carapace. A little longer than wide, cephalic area
moderately raised. Height 4.22; frontal width 6.87. Fovea
weakly procurved, anterior margin not indented. Mid-dorsal
cephalic setae reach fovea. Anterior strial setae numerous.
Anterolateral carapace angle with strong bristles. — Eyes.
Eye region raised. Eye group width 2.20. Median ocular
quadrangle length 0.82, anterior width 1.04, posterior width
1.54. Diameters: AME 0.31, ALE 0.48, PLE 0.36, PME 0.22.
— Chelicerae. Cheliceral groove margins diverge distally.
Groove length 2.87, middle width 0.68. Cheliceral teeth:

320

Records of the Australian Museum (2010) Vol. 62

21 central, occupying full length of groove; 11 prolateral;
10 retrolateral. — Labium. About as long as wide, apically
weakly indented. Length 1.73, width 1.70. Labiosternal
sigilla entire, broad. Cuspules c. 229, number moderate to
high, placed in anterior third to half of labium. — Sternum.
Ovoid. Length 5.58, width 4.25. Posterior sternal sigilla
narrow, elongate. — Palp. Tegular area wider than long.
Embolus offset from tegulum; embolic shaft moderately wide
and strongly curved with distal section recurved at angle to
shaft. Bulb length 3.57, width 1.43. Embolus length 2.41,
midwidth 0.27. Length of femur 5.17, patella 1.80, tibia 5.03.
Width of tibia 2.26. Spination: femur 1, patella 0, tibia 7 (3
dorsal). Distal femur with sinuous bristles. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.16

3.81

6.80

6.56

3.88

29.21

2

7.86

3.64

6.66

5.88

3.88

27.74

3

6.63

3.06

4.96

5.68

3.91

24.24

4

8.43

3.33

6.60

6.83

4.93

30.12

Tibia I width 2.35. Femora 1 and II with dorsal spines.
Metatarsus I thickened proximally, ventral spines numerous.
Metatarsus II strongly sinuous with ventral spined apophysis;
ventral spines absent proximally. Ventral tibia II with
a rounded, spined apophysis centrally; spines lacking
proximally and distally (apart trom distal apical spines).
Coxae frontal thorn-like setae weak or absent. Tarsal,
distal metatarsal scopulae: leg I, II weak to absent; leg 3, 4

Figure 27. Hadronyche emmalizae, male: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) palp,
prolateral; (D) cephalothorax and chelicerae, dorsal; (£) spinnerets;
( F ) cheliceral groove teeth; (G) leg II, prolateral. Scale lines 1 mm.

moderate-strong. Tarsal claw teeth legs I, II: superior 13,11;
inferior 0,0. Trichobothria legs I, II: tarsus 13,13; metatarsus
17, 12; tibia p7 r6, p7 r7. — Leg spination. Leg I: femur 6
(d4p2), patella v2, tibia 43, metatarsus 24, tarsus 21. Leg II:
femur 6 (d5pl), patella p2, tibia 31, metatarsus 32, tarsus 26.
Leg III: femur d5 (bristle-like), patella p5, tibia 9 (p2 r2),
metatarsus 30, tarsus 34. Leg IV: femur 0 (strong bristles),
patella 0, tibia 3, metatarsus 27, tarsus 39. — Abdomen.
Posterior lateral spinnerets with moderately long apical
segment. Lengths: total 3.31; basal segment 1.29, middle
0.61, apical 1.41. Apical segment width 0.49.

Female (paratype AMS KS10675) — Size. Carapace length
11.05, width 10.34. Abdomen length 12.92, width 9.35. —
Colour. Basic colour pattern. — Carapace. Slightly longer
than wide, cephalic area broad, strongly raised. Height 5.10;
frontal width 9.69. Cephalic length 7.96. Fovea procurved.
Mid-dorsal cephalic setae numerous, reach back to fovea.
Anterior strial setae present. Strial and marginal carapace
hairs numerous. Anterolateral carapace angle with numerous,
slender bristles. — Eyes. Central eye region slightly raised.
Eye group width 2.72. Diameters. AME 0.29, ALE 0.65,
PLE 0.36, PME 0.28. Interdistances: AME-AME 0.44,
AME-ALE 0.37, ALE-PLE 0.29, PLE-PME 0.20, PME-
PME 1.31. Median ocular quadrangle length 0.85, anterior
width 1.09, posterior width 1.84. — Chelicerae. Cheliceral
groove wide, margins diverging distally. Groove length 3.24,

Figure 28. Hadronyche emmalizae, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
(D) cheliceral groove teeth; (E) cephalothorax and chelicerae,
dorsal; ( F ) spermathecae; (G) leg III, prolateral, patella, tibia,
metatarsus. Scale lines 1 mm.

Gray: Australian funnel-web spiders

321

Figure 29. Hadronyche emmalizae, male (A,D-K, AMS KS113352, Khancoban, NSW; B,C, AMS KS4114): (A) palp, prolateral; (5)
bulb; (C) embolus, distal; ( D,E) distal patella and tibia I— D, ventral, E, prolateral; (F, G) metatarsus I— F, ventral, G, prolateral; (H,I)
distal patella and tibia II— H, ventral, I, prolateral; (J, K) metatarsus II— J, ventral, K, prolateral. Scale lines: 0.3 mm, except A,B 1.0 mm.

322

Records of the Australian Museum (2010) Vol. 62

Table 8. Male morphological data— Hadronyche emmalizae (n = 3).

character

range

mean

character

range

mean

character

ratio

SD

CL

8.06-9.78

8.89

MtlS

24-35 (p0-l)

28

CW/CL

0.93

0.021

CW

7.68-8.91

8.25

TalS

11-21

14

CH/CW

0.46

0.04

CH

3.40-4.22

3.81

Ti2S

25-35

31

CFW/CL

0.72

0.049

CFW

6.12-6.87

6.41

STC2teeth

11-14

13

CHGW/L

0.24

0.006

ChGL

2.52-2.87

2.71

BulbL

3.31-3.67

3.51

SW/SL

0.79

0.025

ChGW

0.60-0.68

0.64

EmbL

2.19-2.72

2.44

LL/LW

0.99

0.064

ChGCT

17-22

20

BulbW

1.23-1.46

1.37

PLSAPW/L

0.39

0.04

LL

1.65-1.73

1.70

EmbmidW

0.22-0.27

0.25

BulbW/EmbL

0.57

0.11

LW

1.67-1.80

1.72

PalpTibL

4.62-5.03

4.78

EmbmidW/L

0.10

0.021

CUSP

222-260

237

PalpTibW

2.09-2.26

2.18

PalpTibW/L

0.46

0.017

SL

5.22-5.58

5.41

PalpTibS

7

7

BulbL/TibL

0.74

0.038

SW

4.15-4.39

4.26

PalpPatS

0-2

1

PLSAPW

0.41-0.59

0.49

PalpFemS

3-5

4

PLSAPL

1.09-1.41

1.28

FelS

2-6

4

PalS

2-6 (pi-2)

3

middle width 0.99. Cheliceral teeth: 47 central, occupying
full length of groove in 2-3 irregular rows; 12 prolateral; 11
retrolateral. — Labium. Slightly wider than long, rectangular,
anterior margin weakly indented. Length 2.24, width 2.52.
Cuspules occupying anterior half of labium. Labiostemal
sigilla entire, slightly narrowed centrally. — Sternum.
Ovoid. Length 7.28, width 5.95. Anterolateral angles with
basally thickened, bristle-like setae. Posterior sigilla narrow,
elongate, length 1.84, width 0.54. — Palp. Spination: patella
1 prolateral bristle, tibia 15, tarsus p6, 3 ventral bristles.
Trichobothria: tibia p8 r7, tarsus 11. Tarsal claws with 7
teeth. — Legs. 1423.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.43

4.52

6.49

5.47

3.47

28.38

2

7.65

4.25

5.44

4.96

3.40

25.70

3

6.22

3.12

4.15

4.27

3.26

21.02

4

8.13

3.60

6.36

5.71

3.88

27.68

Tibia I width 1.98. Metatarsus I proximal width 1.29. Coxae
I, II with several thorn-like setae fronto-medially. Tarsal claw
teeth legs I, II: superior 10, 10; inferior 2, 2. Trichobothria
legs I, II: tarsus 16,16; metatarsus 19,18; tibia plO r9, p9 r9.
—Leg spination. Leg I: femur 0, patella 0, tibia 2, metatarsus
13, tarsus 11. Leg II: femur 0, patella 0, tibia 3, metatarsus
12, tarsus 12. Leg III: femur pi, patella pd8, tibia 11 (p7 r2),
metatarsus 20 (p4 r4), tarsus 14. Leg IV: femur 0, patella 0,
tibia 3 (pi), metatarsus 23, tarsus 25. — Abdomen. Frontal
abdomen with a group of strong, spine-like bristles. Posterior
lateral spinnerets with short apical segment. Lengths: total
4.85; basal segment 2.24, middle 0.99, apical 1.70. Apical
segment width 0.89. — Genitalia. Spermathecae short. Basal
two thirds strongly dilated, wider than digitiform apical third.
Well separated basally (by the width of a spermatheca);
apices rounded, diverging slightly from each other. Length
1.33, width 0.80.

Distribution. South from Bago State Forest to the
Khancoban-Geehi region, western side of Snowy Mountains,
New South Wales (Fig. 26).

Hadronyche formidabilis (Rainbow)

Figures IF, 30-33; Tables 9, 34

Atrax formidabilis Rainbow, 1914: 255. 1955: 772. Main,
1967: 40. Walker, 1982: 4. Main, 1985: 40.

Hadronyche formidabilis. -Gray, 1988: 114 (transferred
from Atrax).

Types. Holotype male: AMS KS1038 (old catalogue number K35282).
Richmond River, New South Wales

Other material examined. New South Wales (males): QMB S13892,
Dorrigo, 10 February 1984, R. Gordon; AMS KS950, Gibraltar Range,
near Glen Innes, 12 May 1966, Schultz (matured as male 27 December
1967); AMS KS935, Karuah State Forest, near Dungog, 13 January 1976;
AMS KS4514, Lakes Way, near Forster, 15 January 1980, G. Sanders;
AMS KS951, South Grafton, July 1965, C. Snook; AMS KS8791, Taree,
29 December 1981; AMS KS934, Taree, 9 January 1976; AMS KS952,
Tea Gardens, 3 January 1969, van Dreuten; AMS KS13390, Ulong, 60
km W. of Coff’s Harbour, 23 November 1983, C. Martin; AMS KS3245,
Wollomombi via Armidale, 2 May 1973, M. Wyndham; AMS KS946,
Wongwibinda, 1951, D. Wright; AMS KS949, Armidale Caravan Park;
QMB S383, Armidale, Apr 1979; AMS KS947, Barrington Tops, 12
November 1961, B. Salkind; AMS KS1667, Comboyne, 16 January 1946,
H.J. Davidson; AMS KS4775, Dalby, 17 February 1938, N. Geary. New
South Wales (females): AMS KS13391, Ulong, 23 November 1983, C.
Martin; AMS KS 13392, Ulong, 23 November 1983, C. Martin; AMS
KS1313, Upper Allyn River area, 10 February 1978, A. D’Ombrain; AMS
KS3547, Wauchope, 10 October 1934; AMS KS7293, Wilson Creek,
31 March 1981, P. Giraud; AMS KS3234, Wollomombi via Armidale, 2
May 1973, M. Wyndham; AMS KS946, Wongwibinda, 1951, D. Wright;
AMS KS1864, Bellingen, AMS KS1058, Cascade, 6 February 1976, R.
Holmes; AMS KS 14225, Dorrigo, 2 April 1984; AMS KS4663, Dungog
area, February 1980; AMS KS5175, Grevillia Saw Mill, May 1980; AMS
KS1367, Kempsey, September 1977, B. Mercer; AMS KS2652, Kempsey,
February 1979, Daniels; AMS KS13628, Lismore, 30 April 1982; AMS
KS3544, Murwillumbah, 1 April 1971, J.O’Reilly; AMS KS5108, Newee
Creek, 8 April 1980, T. Foley; AMS KS8363, Niangla, 2 October 1981, M.
Keat. Queensland (males): QMB S186, Danabah, 1-7 March 1976, V.E.
Davies & R. Raven; QMB S185, O’Reilly’s, Lamington Plateau, 1 January
1973, R. Raven; QMB S184, Binna Burra, 12 March 1954, W. McIntyre;
QMB S187, Binna Burra, 18 December 1976, T. Gynther.

Diagnosis. CL 10.03-12.30 (male). Large, tree dwelling
Atracinae. Differs from most species by tibia II having a
large, rounded spinose apophysis with spines extending
onto distoventral tibia (Figs. 30C; 32H,I); Differs from

Gray: Australian funnel-web spiders

323

H. cerberea, H. versuta and H. emmalizae by the more
prominent tibia II apophysis (Fig. 30C) and longer
spinnerets, PLSAPW/L 0.29 (Fig. 30D; Table 9). Females
differ from other atracine species by presence of dense cover
of long, fine hairs laterally and ventrally on tibiae & metatarsi

I, II (Fig. 3 IB).

Redescription of male holotype. Size. Carapace length
11.39, width 10.27. Abdomen length 10.20, width 7.48. —
Colour. Basic colour pattern. — Carapace. Longer than wide,
cephalic area moderately raised. Height 4.76; frontal width
7.48. Fovea weakly procurved, anterior margin indented
centrally. Mid-dorsal cephalic setae extend almost to fovea.
Anterior strial setae absent basally. Anterolateral carapace
angle with strong bristles. — Eyes. Central eye region raised.
Eye group width 2.65. Median ocular quadrangle length
0.90, anterior width 1.17, posterior width 1.80. Diameters:
AME 0.37; ALE 0.56; PLE 0.39; PME 0.29. —Chelicerae.
Cheliceral groove rather narrow, margins weakly divergent
distally. Groove length 3.75, middle width 0.61. Cheliceral
teeth: 22 central, occupying full length of groove; 12
prolateral; 11 retrolateral. — Labium. Slightly wider than
long, apically weakly indented. Length 1.96, width 2.19.
Labiosternal sigilla entire, broad. Cuspules c. 338 cuspules,
number high. — Sternum. Long, ovoid. Length 6.87, width
4.56. Several bristles grouped at anterolateral angles.
Posterior sternal sigilla broad, elongate. — Palp. Tegular
area wider than long. Embolus of moderate length, weakly

Figure 30. Hadronyche formidabilis, male: (A) cephalothorax
and chelicerae, lateral; ( B ) cheliceral groove teeth; (C) leg III,
prolateral; (D) spinnerets; (E) sternum, labium and maxilla; (F)
palp, prolateral; (G) cephalothorax and chelicerae, dorsal. Scale
lines 1 mm.

offset from tegulum. Shaft moderately wide, weakly curved,
moderately twisted distally. Bulb length 3.60, width 1.36.
Embolus length 2.51, midwidth 0.29. Length of femur 5.24,
patella 2.38, tibia 5.40. Width of tibia 2.31. Spination: femur
3; patella 2; tibia 12, (3 dorsal). Distal femur with sinuous
bristles. — Legs. 1423. Legs I and IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

10.54

4.65

9.01

8.55

4.76

37.51

2

9.04

4.69

8.02

7.93

4.62

34.30

3

8.36

3.84

6.72

7.13

4.54

30.59

4

10.47

4.37

9.04

8.87

4.72

37.47

Tibia I width 1.72. Femora I and II with dorsal spines.
Metatarsus I with numerous ventral spines grouped on
thickened proximal region. Tibia I with many ventral spines,
distal spines short, prolateral spines present. Tibia II with
a large, rounded, strongly spined ventral apophysis placed
slightly proximal of centre. Ventral spines proximal and
distal to apophysis few and small; prolateral spines present.
Metatarsus II strongly sinuous with a prominent spined
ventral apophysis; ventral spines mainly on and distal to
apophysis, few proximally. Thom-like setae present on coxae
I, II, frontal. Tarsal and distal metatarsal scopulae: legs I, II
very weak to absent; legs 3,4 weak to moderate. Tarsal claw
teeth legs I, II: superior 13,12; inferior 3,3. Trichobothrialegs
I, II: tarsus 16,14; metatarsus 16,14; tibia p8r8, p8r8. — Leg
spination. Leg I: femur 11 (d3 pd8), patella 11 (p5), tibia 131

Figure 31. Hadronyche formidabilis, female: (A) cephalothorax
and chelicerae, dorsal; ( B ) leg I, hair cover; (C) leg III, prolateral,
patella, tibia, metatarsus; (D) spinnerets; (E) cheliceral groove teeth;
(F) cephalothorax and chelicerae, lateral; (G) sternum, labium and
maxilla; (H) spermathecae. Scale lines 1 mm.

324

Records of the Australian Museum (2010) Vol. 62

Figure 32. Hadronyche formidabilis, male ( A,D-K , AMS KS935;
B,C, AMS KS947): (A) Palp, prolateral; (5) bulb; (C) embolus,
distal; ( D,E ) distal patella and tibia I— D, ventral, E, prolateral;
( F,G ) metatarsus I— F, ventral, G, prolateral; (El,I) distal patella and
tibia II— H, ventral, I, prolateral; (J,K) metatarsus II— J, ventral, K,
prolateral. Scale lines: 0.3 mm, except B 1.0 mm.

(p7), metatarsus 45, tarsus 44 (2 mid ventral). Leg II: femur
14 (d7pd7), patella 5(p4), tibia 65 (p2), metatarsus 44, tarsus
35 (3 mid ventral). Leg III: femur pdlO, patella 13 (pl2 rl),
tibia 24 (p6 r4), metatarsus 32 (p3 rd2), tarsus 24. Leg IV:

femur 0 (strong bristles), patella rl, tibia 11 (r3), metatarsus
26, tarsus 30. — Abdomen. Posterior lateral spinnerets with a
rather long apical segment. Lengths: total 5.32; basal segment
2.09; middle 1.36; apical 1.87. Apical segment width 0.58.

Gray: Australian funnel-web spiders

325

Table 9. Male morphological data— Hadronyche formidabilis (n = 10).

character

range

mean

character

range

mean

character

ratio

SD

CL*

10.03-12.03

11.06

MtlS

36-69 (p0-l)

49

CW/CL

0.91

0.02

CW

9.39-11.29

10.06

TalS

18-71 (0-5 midv) 39

CH/CW

0.45

0.026

CH

4.10-5.08

4.50

Ti2S*

38-88 (pO-2)

63

CFW/CL

0.67

0.031

CFW

6.36-8.64

7.41

STC2teeth

10-13

11

CHGW/L*

0.15

0.021

ChGL

3.23-3.94

3.55

BulbL

3.40-4.01

3.66

SW/SL

0.73

0.029

ChGW

0.45-0.61

0.53

EmbL

2.04-2.70

2.48

LL/LW

0.92

0.021

ChGCT

15-34

22

BulbW

1.30-1.56

1.39

PLSAPW/L*

0.29

0.024

LL

1.74-2.04

1.87

EmbmidW

0.26-0.33

0.30

BulbW/EmbL

0.56

0.043

LW

1.94-2.26

2.04

PalpTibL

4.88-6.12

5.44

EmbmidW/L

0.12

0.009

CUSP*

331-417

371

PalpTibW

2.13-2.58

2.30

PalpTibW/L

0.43

0.04

SL

6.27-7.28

6.72

PalpTibS

6-16

9

BulbL/TibL

0.67

0.044

SW

4.51-5.44

4.93

PalpPatS

1-3

2

PLSAPW

0.50-0.67

0.57

PalpFemS

2-5

3

PLSAPL

1.67-2.29

1.99

FelS

2-17

9

PalS*

5-25 (p2-8)

11

Female (AMS KS4663), Dungog area, New South Wales,
32°23'S 151°45'E, February 1980). —Size. Carapace length
14.84, width 12.22. Abdomen length 14.28, width 10.54. —
Colour. Basic colour pattern. Abdomen dark or light maroon
brown. — Carapace. Clearly longer than wide, strongly
raised. Height 6.46; frontal width 10.66. Cephalic length
11.36. Fovea procurved. Mid-dorsal cephalic setae weak
but numerous anteriorly, few posteriorly, do not reach fovea.
Anterior strial setae absent basally. Anterolateral carapace
angle with many bristles. — Eyes. Eye region sessile, very
slightly raised centrally. Eye group width 3.53. Diameters:
AME 0.38, ALE 0.78, PLE 0.70, PME 0.40. Interdistances:
AME-AME 0.58, AME-ALE 0.44, ALE-PLE 0.38, PLE-
PME 0.12, PME-PME 1.54. Median ocular quadrangle
length 1.08, anterior width 1.11, posterior width 2.38. —

Figure 33. Collection records for Hadronycheformidabilis (squares)
and H. alpina. (circles).

Chelicerae. Cheliceral groove narrow, margins diverging
distally. Groove length 4.72, middle width 0.82. Cheliceral
teeth: 44 central, in irregular double row, occupying full
length of groove; 13 prolateral; 13 retrolateral. — Labium.
Almost as long as wide, rounded, anterior margin broadly
indented. Length 3.08, width 2.95. Cuspules occupying
anterior three quarters of labium. Labiostemal sigilla entire.
— Sternum. Long. Length 9.10, width 6.60. Posterior sigilla
elongate: length 1.85, width 0.80. — Palp. Spination: tibia 4,
tarsus 7. Trichobothria: tibia plO r9, tarsus 22. Tarsal claw
with 6 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

10.40

5.60

8.36

7.01

4.10

35.47

2

9.43

5.49

7.13

6.48

3.98

32.51

3

8.12

4.51

5.41

5.99

3.98

29.01

4

10.66

5.62

8.20

7.63

4.59

36.70

Tibiae and metatarsi I, II with dense cover of long, fine hairs
ventrally and laterally. Tibiae I, II with 0-1 spines. Tibia I
width 2.62. Metatarsus I proximal width 1.64. Coxae I, II
with a few thorn-like setae frontally. Tarsal claw teeth legs I,
II: superior 7, 6; inferior 3, 3. Trichobothria legs I, II: tarsus
19,15, metatarsus 26,23; tibia p9 r9, p8 r8. — Leg spination.
Leg I: femur 0, patella 0, tibia 1, metatarsus 6, tarsus 7. Leg
II: femur 0, patella 0, tibia 1, metatarsus 8, tarsus 9. Leg III:
femur 0, patella pdl3 rl, tibia 11 (p4 r3), metatarsus 21 (p5
rd3), tarsus 11. Leg IV: femur 0, patella 0, tibia 4, metatarsus
7, tarsus 22. — Abdomen. Posterior lateral spinnerets stout,
apical segment thick and moderately long. Lengths: total
6.73; basal segment 2.54; middle 1.44; apical 2.75. Apical
segment width 0.94. — Genitalia. Spermathecae broad,
converge slightly anteriorly. Apical third rounded, clearly
set off from basal two thirds by a constriction. Spermatheca
length 1.49, width 0.65.

Distribution. Northeastern New South Wales, from the
Hunter River, to southeastern Queensland (Fig. 33). A
single record exists from southeastern New South Wales
(Robertson, southern highlands) but remains unconfirmed.

Comments. This is the largest atracine spider known. Both
males and especially females have noticeably narrower
carapaces than comparable species. They are a tree dwelling

326

Records of the Australian Museum (2010) Vol. 62

species associated with tall open forest and rainforest
habitats. Their retreats are associated with trunk/branch
fracture holes, rotten heartwood pipes and epiphytic growths.
During timber milling, these spiders are regularly found in
the rotting wood and frass environment of heartwood pipes
in large trees such as Tallowwood ( Eucalyptus microcorys).
They probably feed on wood eating beetles and associated
fauna inhabiting the tree pipe habitat.

Hadronyche alpina n.sp.

KS876, Mt. Kosciuszko, Kosciuszko National Park, 36°27'S 148°15'E, 6
January 1929, H.O. Fletcher; AMS KS10602, Thredbo, 36°26'S 148°27’E,
13 January 1983, G. Jackson. Australian Capital Territory. Male: AMS
KS870, Mt. Ginini, Brindabella Ranges, 35°31'S 148°46'E, 5 December
1966, R. Wood.

Other material examined. New South Wales (males): AMS KS878,
Australian Alps, 1950, S.G. Alley; AMS KS876, Mount Kosciuszko,
Kosciuszko National Park, 6 January 1929, H.O. Fletcher; AMS KS873,
Mount Kosciuszko, Kosciuszko National Park, 31 January 1974; AMS
KS874, Mount Kosciuszko near summit, Kosciuszko National Park, 9
February 1952, C.E. Chadwick.

Figures 33-36; Tables 10, 34

Etymology. The specific epithet refers to the species
presence in the Snowy Mountains alpine region of New
South Wales.

Types. Holotype male: AMS KS872, Mt. Kosciuszko, Kosciuszko National
Park, New South Wales, 36°27’S 148° 15E, 13 January 1968, J. Child.
Paratypes (all AM). New South Wales. Males: AMS KS 13808, Kosciuszko
National Park, 5 December 1983, J. Gold; AMS KS8515, South Ramshead,
Kosciuszko National Park, 36°31’S 148°14'E, 16 December 1981, W.S.
Osborne; AMS KS875, West slope of Mt. Kosciuszko, Kosciuszko
National Park, 36°27'S 148°15’E, 17 December 1971, R.F. Jensz; AMS
KS871, Seaman’s Hut, Mt. Kosciuszko, Kosciuszko National Park, 36°27'S
148°15'E, 28 November 1966, H. Cogger; AMS KS23643, Charlotte
Pass area, Kosciuszko National Park, 36°26'S 148° 19E, J. Molan; AMS
KS877, Mt. Kosciuszko, 7 January 1929, H.O. Fletcher & A. Musgrave.
Females: AMS KS8730, Daner’s Gap, Kosciuszko National Park, 36°21'S
148°28'E,W.S. Osborne, 12 January 1982; AMS KS8514, South Ramshead,
Kosciuszko National Park, 36°31'S 148°14'E, 16 December 1981; AMS

Figure 34. Hadronyche alpina , male: (A) cephalothorax and
chelicerae, lateral; ( B ) sternum, labium and maxilla; (C) cheliceral
groove teeth; (D) cephalothorax and chelicerae, dorsal; (E) palp,
prolateral; ( F ) spinnerets; (G) leg II, prolateral (NB, dorsal femoral
spines present or absent). Scale lines 1 mm.

Diagnosis. CL 8.16-10.06 (male). Apophyseal swelling
present on tibia II (Fig. 36H,I). Differs from H. versuta
by shorter embolus and H. emmalizae by shorter, straight
embolus(BulbW/EmbL 0.76) (Fig. 36B);. from H. cerbereci
by absence of coxal thorns; from H. venenata by presence of
metatarsus II apophysis (Fig. 36K); and from H. meridiana
and H. modesta by longer labium (Fig. 34B).

Male (holotype). — Size. Carapace length 9.31, width 9.28.
Abdomen length 9.02, width 6.72. — Colour. Basic colour
pattern. — Carapace. Almost as wide as long, moderately
raised. Height 3.81. Frontal width 5.95. Fovea weakly
procurved-straight. Mid-dorsal cephalic setae almost reach
fovea. Anterior strial area with none or few setae. Strong
bristles on anterolateral carapace angle. — Eyes. Sessile
to slightly raised. Eye group width 2.28. Median ocular

teeth; ( B ) cephalothorax and chelicerae, lateral; (C) sternum,
labium and maxilla; ( D ) cephalothorax and chelicerae, dorsal; (E)
spermathecae; ( F ) spinnerets; (G) leg III, prolateral, patella, tibia,
metatarsus. Scale lines 1 mm.

Gray: Australian funnel-web spiders

327

Figure 36. Hadronyche alpina, male ( A,D-K , AMS KS23643; B, C,
AMS KS877): (A) palp, prolateral; ( B ) bulb; (C) embolus, distal;
(. D,E ) distal patella and tibia I— D, ventral, E, prolateral; (F,G)
metatarsus I— F, ventral, G, prolateral; ( H,1 ) distal patella and
tibia II— H, ventral, /, prolateral; (J,K) metatarsus II— J, ventral,
K, prolateral. Scale lines: 0.3 mm, except A 0.5 mm, D 1.0 mm
and F,H, 0.2 mm.

quadrangle length 1.10, anterior width 1.04, posterior width
1.44. Diameters: AME 0.30, ALE 0.47, PLE 0.41, PME
0.21. — Chelicerae. Cheliceral groove rather narrow, margins
subparallel to weakly divergent. Groove length 3.18, middle

width 0.58. Cheliceral teeth: 29 central, running full length
of groove, many often proximally placed; 10 prolateral; 11
retrolateral. — Labium. Slightly wider than long. Length
1.60, width 1.74. Labiostemal sigilla, broad entire. Cuspules

328

Records of the Australian Museum (2010) Vol. 62

Table 10. Male morphological data— Hadronyche alpina (n = 10).

character

range

mean

character

range

mean

character

ratio

SD

CL

8.16-10.06

9.21

MtlS

25-45

35

CW/CL

0.96

0.038

CW

7.63-9.32

8.81

TalS

8-15

12

CH/CW

0.41

0.021

CH

3.31-4.08

3.65

Ti2S

21-48

32

CFW/CL

0.66

0.045

CFW

5.44-6.36

6.04

STC2teeth

13-16

14

CHGW/L

0.18

0.029

ChGL

2.53-3.18

2.84

BulbL

2.61-3.20

2.87

SW/SL

0.77

0.037

ChGW

0.40-0.63

0.51

EmbL

1.59-1.96

1.75

LL/LW

0.89

0.038

ChGCT

4-29

15

BulbW

1.17-1.67

1.31

PLSAPW/L*

0.33

0.042

LL

1.07-1.64

1.49

EmbmidW

0.21-0.28

0.23

BulbW/EmbL*

: 0.76

0.059

LW

1.25-1.84

1.67

PalpTibL

3.94-4.76

4.42

EmbmidW/L

0.14

0.011

CUSP*

110-140

128

PalpTibW

1.87-2.28

2.10

PalpTibW/L

0.48

0.033

SL

4.66-6.05

5.55

PalpTibS

0-7

3

BulbL/TibL

0.66

0.041

SW

3.67-4.72

4.29

PalpPatS

0-1

1

PLSAPW

0.41-0.56

0.48

PalpFemS

3-5

4

PLSAPL

1.29-1.78

1.46

FelS

0-8

3

PalS

3-10 (pi-4)

6

c. 130, number low to moderate. — Sternum. Ovoid. Length
5.58, width 4.32. Posterior sigilla large, elongate. — Palp.
Tegular area slightly wider than long. Embolus short, weakly
curved, with short, weakly twisted distal part; basal embolus
slightly or not offset from tegulum. Bulb length 2.83, width
1.34. Embolus length 1.65, midwidth 0.27. Length of femur
4.31, patella 1.97, tibia 4.43. Width of tibia 2.12. Spination:
femur 3; tibia 2. Femur with distal sinuous bristles. — Legs.
4213. Legs I, 2 subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.36

4.06

6.39

6.31

4.17

29.29

2

8.16

4.10

6.89

6.52

4.10

29.77

3

6.93

3.47

5.46

5.82

4.02

25.70

4

8.36

3.85

7.16

6.97

4.67

31.01

Tibia I width 1.62. Femora I and II usually with, sometimes
without, dorsal spines. Femora III and IV with strong, long
bristles. Metatarsus I slightly thickened proximally, larger
ventral spines grouped proximally. Tibia II with small
apophyseal swelling, centrally placed with spines grouped
upon it. Tibia II weakly concave distoventrally. Metatarsus
II sinuous with a prominent ventral, spined apophysis.
Coxal hairs unmodified. Scopulae well developed on tarsi
of all legs and distal metatarsi of legs II-IV. Tarsal claw
teeth legs I, II: superior 4; inferior 2. Trichobothria legs I,
II: tarsus 12,13; metatarsus 17,16; tibia p9 r8, p7 r6. — Leg
spination. Leg I: femur d3 (plus bristles), patella 10 (p4
v6), tibia 72 (p4), metatarsus 39, tarsus 12. Leg II: femur
d3 (plus bristles), patella 3 (p2 vl), tibia 32, metatarsus 21,
tarsus 12. Leg III: femur 0, patella 8 (p7 vl), tibia 10 (p4 r2),
metatarsus 22, tarsus 10. Leg IV: femur 0, patella 0, tibia
2, metatarsus 17, tarsus 18. — Abdomen. Posterior lateral
spinnerets with moderately long apical segment. Lengths:
total 3.36; basal segment 1.20, middle 0.64, apical 1.52;
apical width 0.49.

Female (paratype AMS KS8730). — Size. Carapace length
10.02, width 9.35. Abdomen length 15.64, width 11.22. —
Colour. Basic colour pattern. — Carapace. Slightly longer
than wide, strongly raised. Height 5.07, frontal width 8.23.
Cephalic length 7.14. Fovea procurved. Mid-dorsal cephalic

setae almost reach or reach fovea. Anterior strial setae few to
absent basally. Anterolateral carapace angle with a few weak
bristles and hairs. — Eyes. Central eye region weakly raised.
Eye group width 2.72. Diameters: AME 0.25, ALE 0.52, PLE
0.44, PME 0.20. Interdistances: AME-AME 0.44, AME-
ALE 0.31, ALE-PLE 0.42, PLE-PME 0.22, PME-PME
1.16. Median ocular quadrangle length 0.98, anterior width
1.03, posterior width 1.69. — Chelicerae. Cheliceral groove
margins diverge distally. Groove length 3.59, middle width
0.82. Cheliceral teeth: 28 central, in single row, occupying
full length of groove; 11 prolateral; 9 retrolateral. — Labium.
Slightly wider than long, sides sloping medially to convex
anterior margin. Length 2.04, width 2.41. Cuspules on
anterior half of labium. Labiosternal sigilla entire, narrow.
— Sternum. Ovoid. Length 6.90, width 5.24. Posterior sigilla
elongate: length 1.36, width 0.58. — Palp. Spination: tibia
2, tarsus 7. Trichobothria: tibia p6 r7, tarsus 8. Tarsal claw
with 6 teeth. — Legs. 1423. Legs I, IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.15

4.32

5.81

4.79

3.20

26.27

2

7.00

3.94

5.24

4.28

2.86

23.32

3

6.12

3.37

3.77

3.92

2.99

20.17

4

7.75

4.22

5.62

5.09

3.33

26.01

TibialsNi&lh. 1.84. Metatarsus I proximal width 1.26. Coxal
with anterior setae thickened basally. Tarsal claw teeth legs I,
II: superior 11,11; inferior 3,3. Trichobothria legs I, II: tarsus
11,12; metatarsus 14,14; tibia p9 r7, p7 r8. — Leg spination.
Leg I: femur 0, patella 0, tibia 1, metatarsus 8, tarsus 6. Leg
II: femur 0, patella 0, tibia 1, metatarsus 9, tarsus 8. Leg
III: femur 0, patella pd8, tibia 15 (p8 r4), metatarsus 20 (p9
rd3), tarsus 16. Leg IV: femur 0, patella 0, tibia 2, metatarsus
22, tarsus 33. — Abdomen. Posterior lateral spinnerets with
short apical segment. Lengths: total 3.95; basal segment
1.63, middle 0.82, apical 1.50. Apical segment width 0.67.
— Genitalia. Spermathecae about twice as long as wide,
digitiform, apical half slightly enlarged. Length 0.95, width
0.45. Spermathecae diverge slightly anteriorly.

Distribution. Kosciuszko National Park, New South Wales
and Brindabella Range, Australian Capital Territory (Fig. 33).

Gray: Australian funnel-web spiders

329

Hadronyche venenata (Hickman)

Figures 37-40; Tables 11, 34

Atrax venenatus Hickman, 1927: 63. Hickman, 1967: 20.
Main, 1985: 42.

Hadronyche venenata- Gray, 1988: 114 (Transferred from
Atrax).

Types. Holotype male: QVM 957-13-16 Type 28. Newtown Creek, Hobart,
Tasmania, V.V. Hickman, 22 December 1925. Allotype female: QVM
1957-13-17 Type 29. Data as for holotype.

Other material examined. Tasmania (males): AMS KS6071, Lambert
Park, near Hobart, 3 June 1980, M. Gray; Bicheno, 22 January 1972, B.
Stephenson (QVM); Cape Lodi, near Bicheno, 5 March 1978, A. McBain
(QVM); Cole’s Bay, 27 April 1972, N.S. Freeman (QVM); Cole’s Bay, 13
March 1978, Monaghan (QVM); Cole’s Bay, 24 February 1971, M. O’Toole
(QVM); Mathina, 16 February 1972 (QVM): Mathina, 10 February 1971,

J. Turner (QVM); AMS KS972, Mount Ben Lomond, 11 April 1926, A.L.
Meston; J1916 (TM), GeilstonBay, 16 May 1984, J. McDavitt; J1360 (TM),
Rosny, Hobart, 17 April 1978, Mr Ward; E938 (TM), Blackman’s Bay, 5
May 1974, E. Cunliffe; Hobart, June 1971, K. Hamilton (QVM); J1917
(TM), Winnebah, May 1984, F. Wagner; J 739 (TM), Tinderbox, 8 March
1971, D. Milledge; J1113 (TM), West Hobart, 26 March 1976, R. van de
Uusse; J1025 (TM), Taroona, Hobart, 9 March 1975, Ms Stottard. Tasmania
(females): Blackman’s Bay, Apr 1983 (TM); AMS KS975, Cascades, 3
January 1967; Coles Bay, February 1970, N. Lawson (TM); Cromwell, 20
April 1971, N. Toombs (TM); Golden Ridges, 13 February 1946, W. Bart
(QVM); Hobart, June 1971, K. Hampton (QVM); Liffy, 28 October 1972,

K. Watson (QVM); Mathina, 10 February 1971, J. Turner (QVM); Mount
Victoria, 14 January 1972, N.B. Brown (QVM); Mount Young, 18 January
1971, ? Simpson (QVM); Poatina, 28 September 1963, F. Fishwick (QVM);
Ringarooma, 7 June 1963, J. Kidd (QVM); Rosetier, 7 June 1971 (TM);
AMS KS974, Sandy Bay, 12 December 1966, J. Cossum; St. Marys, 16 July
1971, K. Cook (QVM); AMS KS6214, Taroona, near Hobart, 8 September
1980, R. Parrott; AMS KS973, Trevallyn, 1 February 1928.

Figure 38. Hadronyche venenata, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
( D ) cephalothorax and chelicerae, dorsal; (E) leg II, prolateral;
( F ) palp, prolateral; (G) cheliceral groove teeth. Scale lines 1 mm.

Figure 37. Collection records for Hadronyche venenata

Diagnosis. CL 6.88-9.59 (male). Differs from other
Hadronyche spp. by having a weak spined apophyseal
swelling on tibia II but metatarsus II not modified(Figs.
38E, 40H-K). Male caput relatively weakly raised (Fig.
38B; CH/CW 0.40).

Redescription of male holotype. Size. Carapace length 6.88,
width 6.19. Abdomen length 5.30, width 4.42. — Colour.
Basic colour pattern. — Carapace. Longer than wide,
cephalic area moderately raised, rather narrow frontally.
Height 2.46. Frontal width 3.69. Fovea straight, narrow.
Mid-dorsal cephalic setae reach fovea. Anterior strial setae

Figure 39. Hadronyche venenata, female: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) leg
III, prolateral, patella, tibia, metatarsus; ( D ) cephalothorax and
chelicerae, dorsal; ( E ) cheliceral groove teeth; (F) spermathecae;
(G) spinnerets. Scale lines 1 mm.

330

Records of the Australian Museum (2010) Vol. 62

Figure 40. Hadronyche venenata, male {A,D-K, AMS KS6071; B,C, AMS KS972): (A) palp, prolateral; (B) bulb; (C) embolus, distal;
(. D,E ) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; ( H,I) distal patella and tibia II— H,
ventral, /, prolateral; ( J,K) metatarsus II— J, ventral, K, prolateral. Scale lines: 0.3 mm, except A 0.2 mm and B 1.0 mm.

present. Anterolateral carapace angle with several weak
bristles. — Eyes. Central eye region raised. Anterior median
eyes small. Eye group width 1.60. Median ocular quadrangle
length 0.58, anterior width 0.70, posterior width 1.09.
Diameters: AME 0.17, ALE 0.38, PLE 0.26, PME 0.20. —

Chelicerae. Groove margins subparallel, weakly divergent
distally; length 2.02, middle width 0.26. Cheliceral teeth:
16 central, occupying full length of groove; 9 prolateral;
7 retrolateral. — Labium. Slightly wider than long, weakly
indented apically. Length 1.14, width 1.24. Labiosternal

Gray: Australian funnel-web spiders

331

Table 11. Male morphological data— Hadronyche venenata (n = 12).

character

range

mean

character

range

mean

character

ratio

SD

CL

6.88-9.59

8.52

MtlS

24-53 (p0-l)

37

CW/CL

0.95

0.029

CW

6.19-8.82

8.11

TalS

9-24

14

CH/CW*

0.40

0.016

CH

2.46-3.62

3.27

Ti2S

16-39

26

CFW/CL

0.65

0.047

CFW

3.69-6.45

5.49

STC2teeth

12-16

13

CHGW/L*

0.13

0.014

ChGL

2.02-2.83

2.53

BulbL

1.58-2.83

2.44

SW/SL

0.71

0.056

ChGW

0.26-0.39

0.32

EmbL

1.33-1.84

1.57

LL/LW

0.93

0.029

ChGCT

12-24

17

BulbW

0.94-1.44

1.09

PLSAPW/L*

0.36

0.057

LL

1.14-1.60

1.44

EmbmidW

0.16-0.22

0.18

BulbW/EmbL

0.67

0.037

LW

1.24-1.72

1.56

PalpTibL

3.03-1.18

3.77

EmbmidW/L

0.12

0.008

CUSP

213-325

269

PalpTibW

1.41-1.96

1.77

PalpTibW/L

0.47

0.042

SL

3.94-5.92

5.28

PalpTibS

4-8

6

BulbL/TibL

0.65

0.032

SW

2.84-4.20

3.53

PalpPatS

1-3

1

PLSAPW

0.32-0.64

0.46

PalpFemS*

5-11

7

PLSAPL

0.84-1.53

1.43

FelS*

0-3

1

PalS

4-14 (pi—3)

9

sigilla narrowed toward midline. Cuspules c. 220, number
moderate to high. — Sternum. Ovoid, moderately long.
Length 3.94, width 2.84. Posterior sigilla ovoid, broad. —
Palp. Tegular area wider than long. Embolus rather short,
base hardly offset from tegulum. Embolus shaft weakly
curved to straight, only slightly tapered distally. Distal part
of embolus weakly twisted and short. Bulb length 2.07,
width 0.94. Embolus length 1.37, midwidth 0.16. Length
of femur 3.00, patella 1.34, tibia 3.03. Width of tibia 1.41.
Spination: femur 4, patella 1, tibia 7 (3 dorsal). Some distal
femoral spines and bristles sinuous. — Legs. 4123. Legs I,
IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.99

2.87

4.94

4.55

2.96

21.31

2

5.90

2.83

4.72

4.26

2.80

20.51

3

5.03

2.43

3.85

3.53

2.79

17.62

4

5.99

2.68

4.84

4.90

2.94

21.35

Tibia I width 1.17. Femora I and II with dorsal spines. Tibia

I ventral spines numerous. Metatarsus I slightly thickened
proximally, larger ventral spines grouped proximally. Tibia

II with weak apophyseal swelling in proximal half, ventral
spines grouped upon it; distal ventral tibia II weakly concave,
without spines (except apically). Metatarsus II without
apophysis, straight to weakly sinuous (midventral region
somet im es slightly thickened) with strong midventral to
distal spines. Coxal setae unmodified. Scopulae legs I-IV:
tarsus, weak to moderate; metatarsus, absent to weak.
Tarsal claw teeth legs I, II: superior 13, 13; inferior 2, 1.
Trichobothria legs I, II: tarsus 10, 10, metatarsus 10, 11;
tibia p6 r6, p6 r6. — Leg spination. Leg I: femur 4(d3 pi),
patella 6(p2 v4), tibia 46, metatarsus 28, tarsus 12. Leg II:
femur d3, patella pi, tibia 26, metatarsus 16, tarsus 9. Leg
III: femur d3, patella p3, tibia 11 (p2 r2 d2), metatarsus 15,
tarsus 10. Leg IV: femur d4, patella 0, tibia 9 (r3), metatarsus
16, tarsus 12 — Abdomen. Posterior lateral spinnerets: apical
segment moderately long. Lengths: total 2.36; basal segment
0.96, middle 0.56; apical 0.84. Apical segment width 0.32.

Redescription of female allotype. Size. Carapace length
7.38, width 6.81. Abdomen length 10.61, width 8.70.
— Colour. Basic colour pattern. 3-4 pairs abdominal
chevrons, middle pairs distinct and almost meeting mid-

dorsally. — Carapace. Slightly longer than wide, strongly
raised. Height 3.03. Frontal width 5.13. Cephalic length
5.34. Fovea slightly procurved-straight, anterior margin
slightly indented. Mid-dorsal cephalic setae numerous and
run back to fovea. Anterior strial setae present. Anterolateral
carapace angle with several hairs and weak bristles. —
Eyes. Central eye region slightly raised. Eye group width
1.91. Diameters: AME 0.18, ALE 0.45, PLE 0.37, PME
0.24. Interdistances: AME-AME 0.30, AME-ALE 0.18,
ALE-PLE 0.27, PLE-PME 0.14, PME-PME 0.79. Median
ocular quadrangle length 0.60, anterior width 0.70, posterior
width 1.25. — Chelicerae. Groove margins diverge weakly
distally; length 2.20, middle width 0.40. Cheliceral teeth:
18 central, in single row occupying full length of groove;
11 prolateral; 11 retrolateral. — Labium. Slightly wider
than long, rounded, apically indented. Length 1.47, width
1.65. Cuspules occupying anterior two-thirds. Labiosternal
sigilla constricted at midline. — Sternum. Ovoid, moderately
long. Length 4.80, width 3.54. Posterior sigilla ovoid:
length 0.78, width 0.42. — Palp. Spination: tibia 2, tarsus
7. Trichobothria: tibia p5 r4, tarsus 9. Tarsal claw with 3
teeth. — Legs. 1423.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.58

3.12

4.31

3.44

2.13

18.58

2

5.08

2.92

3.49

3.02

2.05

16.56

3

4.26

2.46

2.71

2.67

2.05

14.15

4

5.25

2.87

3.90

3.36

2.42

17.80

Tibia / width 1.40. Metatarsus I proximal width 0.97. Coxal
setae unmodified. Tibia I, II with few or no spines. Tarsal
claw teeth legs I, II: superior 10,9; inferior 3,3. Trichobothria
legs I, 2: tarsus 10, 9; metatarsus 11, 10; tibia p7 r7, p6 r5.
—Leg spination. Leg I: femur 0, patella 0, tibia 0, metatarsus
11, tarsus 8. Leg II: femur 0, patella 0, tibia 0, metatarsus
10, tarsus 8. Leg III: femur 0, patella pd2, tibia 6 (p3 lr),
metatarsus 13 (p4 rl), tarsus 12. Leg IV: femur 0, patella 0,
tibia 2 (rl) metatarsus 9, tarsus 13. — Abdomen. Posterior
lateral spinnerets with short apical segment. Lengths: total
3.02; basal segment 1.30, middle 0.72, apical 1.00. Apical
segment width 0.52. — Genitalia. Spermathecae digitiform,
about three times longer than wide. Length 0.82, width
0.29. Apical third set off from basal two thirds by a slight
constriction.

332

Records of the Australian Museum (2010) Vol. 62

Distribution. Eastern Tasmania (Fig. 37).

Comments. Hickman (1927,1967) described tibia II of this
species as lacking “spurs or apophyses”, but with the “basal
half ... slightly expanded and ... heavily spined”. This low
spinose swelling on tibia II is interpreted here as a weak
apophyseal swelling.

Hadronyche marracoonda n.sp.

Figures 41-44; Tables 12, 34

Etymology. The specific epithet is an Aboriginal word
meaning toward the west, a reference to the western slopes
distribution of this species in New South Wales.

Types. Holotype male: AMS KS892, Wagga Wagga Teachers College,
Wagga Wagga, New South Wales, 35°06'S 147°22'E, 20 April 1971.
Paratypes (all AM). New South Wales. Males: AMS KS890, “The Decca”,
Bigga area, 34°04'S 149°09'E, 12 May 1972, M. Chudleigh; AMS KS13353,
3km E. of Binda, 34°04'S 149°09'E, 28 March 1983, M. Gray & C.
Horseman; AMS KS5223, Crookwell, 34°27’S 149°28'E, 19 May 1980,
L.E. Willis; AMS KS879, Oberon, 33°41’S 149°52E, May 1974, J. Bearup;
AMS KS2719, Tumbarumba, 35°45’S 148°00'E, 10 April 1979, J. Frost;
AMS KS43617, CarcoarDam, 55km SW of Bathurst, 33°37’S 149°14'E, 17
May 1995, J. Chaffey; AMS KS6876, Caloola via Newbridge, 12 February
1981, Callan. Females: AMS KS 10831, Bigga, 34°04’S 149°09’E, 60 km
N.W. of Crookwell, 23 March 1983; AMS KS5740, Adelong area, 35°17'S
148°03'E, July 1980, M. Pearce; AMS KS8796, Gocup, 35°13’S 148°12E,
9 March 1982, B. Laird; AMS KS1371, Oberon, 33°41'S 149°52'E, 16
May 1978, D.H. Clowes, AMS KS10681, Orange, 33°16'S 149°06'E, 17

Figure 41. Hadronyche marracoonda, male: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cheliceral
groove teeth; ( D ) palp, prolateral; (E) cephalothorax and chelicerae,
dorsal; ( F ) spinnerets; (G) leg II, prolateral. Scale lines 1 mm.

January 1983, E.T. Bannigan; AMS KS2668, Tumut, 35°18’S 148°13’E,
30 January 1979, K. Pearce.

Other material examined. New South Wales (males): AMS KS900,
“Bonniemuir”, Adelong area, 29 April 1955, J.D. McMahon; AMS
KS893, Barry, 27 November 1975, Drs. Jones & Redhead; AMS KS891,
“Yewrangara”, 8 kmW. of Bigga, 11 July 1972,1. Chudleigh; AMS KS5202,
64 km W. of Crookwell & 20 km from Wyangla Dam, 6 May 1980, L.E.
Willis; AMS KS 13354, 3 km E. of Binda, 28 March 1983, M. Gray & C.
Horseman; AMS KS888, Bloomfield, N.S.W; AMS KS2720, Cowra, 18
April 1979, M. Henderson; AMS KS898, Crookwell, 22 February 1958,
Clifton’s Pharmacy; AMS KS3077, 10 km N. of Goulburn, May 1979, N.
Vickers; AMS KS884, Holbrook area, 15. iv. 1971, R. Flynn; AMS KS9314,
Humula, 19 May 1982; AMS KS894, Kanangra Walls, near Oberon, 5 June

1956, F.B. Ilann; AMS KS886, Lime kiln s, 20 October 1967, O. Stark; AMS
KS8364, Livingstone State Forest, near Wagga Wagga, 16 September 1981,
Dr Cook; AMS KS897, Nashdale, 10 February 1969, Dixon; AMS KS880,
Oberon, 22 September 1972; AMS KS902, Orange, 20 February 1976,
Bloomfield Hospital; AMS KS903, Orange, 1. ix.1969, F. Freeman; AMS
KS905, Rosewood, 10 May 1972, O. Portors; AMS KS906, Rosewood,
February 1976, G. Portors; AMS KS904, Spring Hill, 20 km E. of Orange,
28 March 1974; AMS KS883, Taralga area, 1977, Goulbum Ambulance;
AMS KS885, Towac, 21 March 1965, J. Maybin; AMS KS6645, Turnkey
Creek, 27 January 1981, J. Dellow; AMS KS889, Tumbarumba, August

1957, R.A. Castle; AMS KS901, Tumorrama, 19 May 1972, M. Blundell;
AMS KS2965, Tumut, 5 May 1979, P.D. Slater; AMS KS8922, Tumut,
6 April 1982, Pearce; AMS KS13424, Wagga Wagga, R. Faulder; AMS
KS887, Yetholme, 19 October 1967, O. Stark; Binalong, 20 February
1975 (ANIC). Australian Capital Territory (male): Campbell, 1974,
D.J. Belford (ANIC).

Diagnosis. CL 8.92-10.71. Larger spiders, leg II without
apophyses/swellings, tibia II and metatarsus II ventrally

Figure 42. Hadronyche marracoonda, female: (A) cheliceral groove
teeth; ( B ) cephalothorax and chelicerae, lateral; (C) sternum,
labium and maxilla; ( D ) cephalothorax and chelicerae, dorsal; (E)
leg III, prolateral, patella, tibia, metatarsus; ( F ) spinnerets; (G)
spermathecae. Scale lines 1 mm.

Gray: Australian funnel-web spiders

333

Figure 43. Hadronyche marracoonda, male ( A,D-K , AMS KS43617; B,C, AMS KS6876): (A) palp, prolateral; (B) bulb; (C) embolus,
distal; ( D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; (El,I) distal patella and tibia
II— H, ventral, /, prolateral; (J, K) metatarsus II— J, ventral, K, prolateral. Scale lines: 1.0 mm, except E-G,J,K 0.3 mm.

334

Records of the Australian Museum (2010) Vol. 62

Table 12. Male morphological data— Hadronyche marracoonda (n =

21).

character

range

mean

character

range

mean

character

ratio

SD

CL

8.92-10.71

9.99

MtlS

29-50

40

CW/CL

0.92

0.026

CW

8.05-10.05

9.23

TalS

15-51

28

CH/CW

0.44

0.037

CH

3.44-4.63

4.10

Ti2S*

6-19 (pO-2)

11

CFW/CL

0.66

0.028

CFW

5.90-7.48

6.64

STC2teeth

12-16

14

CHGW/L

0.20

0.023

ChGL

2.51-3.09

2.83

BulbL

2.90-3.60

3.19

SW/SL

0.79

0.03

ChGW

0.48-0.70

0.56

EmbL

1.91-2.42

2.06

LL/LW

0.89

0.041

ChGCT

12-37

24

BulbW

1.15-1.46

1.32

PLSAPW/L*

0.34

0.026

LL

1.48-1.96

1.69

EmbmidW

0.18-0.31

0.24

BulbW/EmbL

0.65

0.032

LW

1.67-2.16

1.90

PalpTibL

3.98-4.85

4.50

EmbmidW/L

0.12

0.02

CUSP

230-384

280

PalpTibW

1.80-2.25

2.11

PalpTibW/L

0.47

0.02

SL

5.44-6.70

6.06

PalpTibS

2-5

3

BulbL/TibL

0.71

0.032

SW

4.12-5.18

4.77

PalpPatS

0

—

PLSAPW

0.43-0.64

0.54

PalpFemS

0-4

1

PLSAPL

1.43-1.91

1.59

FelS

1-13

8

PalS

2-12 (p2—7)

7

concave,bowed (Fig. 431,K). Differs from infensa and
lamingtonensis group species by having dorsal spines on
femur I, II; from adelaidensis group species by having a long
labium; from//, monaro by lacking metatarsus II apophyseal
swelling, and having numerous tarsus II spines (Table 12);
from H. nimoola by caput not as strongly raised; from H.
tambo by more spinose femur 1 and palpal tibia (Table 12).

Male (holotype) — Size. Carapace length 9.68, width 8.86.
Abdomen length 9.51, width 7.13. — Colour. Basic colour
pattern. — Carapace. Longer than wide, moderately raised.
Height 3.69. Frontal width 5.99. Fovea gently procurved.
Mid-dorsal cephalic setae do not reach fovea. Anterior
strial setae absent. — Eyes. On low tubercle. Eye group
width 2.08. Median ocular quadrangle length 0.90, anterior
width 1.05, posterior width 1.37. Diameters: AME 0.37,
ALE 0.52, PLE 0.43, PME 0.34. — Chelicerae. Cheliceral
groove of moderate width, margins diverging. Groove
length 2.64, middle width 0.55. Cheliceral teeth: 20 central,
running full length of groove; 15 prolateral; 10 retrolateral.
— Labium. Slightly wider than long. Length 1.54, width
1.82. Labiosternal sigilla narrowed centrally. Cuspules
c. 248 cuspules, number moderate to high. — Sternum.
Ovoid, broad. Length 5.74, width 4.84. Posterior sigilla
large, elongate. — Palp. Tegular area slightly wider than
long. Embolus rather short, wide and gently curved; distal
embolus twisted. Embolus hardly offset from tegulum. Bulb
length 3.01, width 1.24. Embolus length 1.91, mid width
0.22. Length of femur 4.17; patella 1.60; tibia 4.23. Width
of tibia 2.02. Spination: tibia 4. Sinuous bristles on distal
femur. — Legs. 4213.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.87

3.28

5.21

6.27

3.77

26.40

2

7.83

3.77

6.71

6.78

4.17

29.26

3

6.97

3.16

5.33

6.11

4.14

25.71

4

8.12

3.44

6.77

7.30

4.59

30.22

Tibia / width 1.72. Metatarsi longer than tibiae. Femora I and
II with dorsal spines. Bristle-like spines on femora III and IV.
Metatarsus I slightly thickened proximally. Tibia II without
apophysis, slightly bowed and ventrally concave, spines
few, grouped proximally. Metatarsus II straight to slightly
bowed and ventrally concave and lacking an apophysis,

rarely weakly sinuous. Coxal hairs unmodified. Tarsal and
distal metatarsal scopulae weakly developed, strongest on
tarsi 3, 4. Tarsal claw teeth legs I, II: superior 15, inferior 4,
trichobothria legs 1,2: tarsus 12,11; metatarsus 16,14; tibia
p6 r7, p6 r7. — Leg spination. Leg I: femur 9 (d3p6), patella
5 (p3), tibia 29 (p2), metatarsus 33, tarsus 19. Leg II: femur
10 (d4p6), patella 4 (p4), tibia 9 (pi), metatarsus 19, tarsus
29. Leg III: femur rd5 (long, bristle-like), patella 9 (p8), tibia
18 (r2 d2), metatarsus 21, tarsus 14. Leg IV: femur r8 (long,
bristle-like), patella pi, tibia 9(r2), metatarsus 21, tarsus
19. — Abdomen. Numerous long bristles dorsally, strongest
anteriorly. Posterior lateral spinnerets with moderately long
apical segment. Lengths: total 3.90; basal segment 1.48,
middle 0.92, apical 1.50; apical width 0.52.

Female (paratype AMS KS10831) — Size. Carapace length
11.64, width 9.35. Abdomen length 10.88, width 8.84. —
Colour. Basic colour pattern. — Carapace. Clearly longer
than wide, strongly raised. Height 4.84; frontal width 8.30.
Cephalic length 8.16. Fovea procurved. Mid-dorsal cephalic
setae do not reach fovea. Anterior strial setae absent.
Anterolateral carapace angle with numerous strong bristles.
— Eyes. Ocular area strongly raised, anteriorly protuberant.
Eye group width 2.62. Diameters: AME 0.33, ALE 0.52, PLE
0.36, PME 0.22. Interdistances: AME-AME 0.44, AME-
ALE 0.34, ALE-PLE 0.26, PLE-PME 0.20, PME-PME
1.30. Median ocular quadrangle length 0.80, anterior width
1.10, posterior width 1.81. — Chelicerae. Cheliceral groove
margins diverge distally, groove wide. Groove length 3.37,
middle width 0.87. Cheliceral teeth: 68 central, in several
rows, occupying full length of groove; 15 prolateral; 11
retrolateral. — Labium. Slightly wider than long, rounded,
anterior margin weakly indented. Length 2.36, width 2.68.
Labiosternal sigilla entire. — Sternum. Ovoid. Length 7.05,
width 5.54. Posterior sigilla long, ovoid: length 0.88, width
0.54. — Palp. Spination: tibia 4, tarsus 9. Trichobothria: tibia
p9 r9, tarsus 13. Tarsal claw with 5 teeth. — Legs. 1423. Legs
I, IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.12

4.51

5.99

5.05

3.06

26.73

2

7.30

4.20

5.33

4.89

3.12

24.84

3

6.17

3.28

3.77

4.39

3.08

20.69

4

7.83

4.06

5.58

5.37

3.42

26.26

Gray: Australian funnel-web spiders

335

<•0 *

Figure 44. Collection records for Hadronyche marracoonda
(squares) and H. tambo (circles).

Tibia / width 2.01. Metatarsus I proximal width 1.38. Coxae
I, II with numerous basally thickened thorn-like setae
anteriorly. Tarsal claw teeth legs I, II: superior 9, 9; inferior
5, 5. Trichobothria legs I, II: tarsus 15, 16; metatarsus 17,
15; tibia p9 rlO, plO r9. — Leg spination. Leg I: femur 0,
patella 0, tibia 3, metatarsus 1, tarsus 7. Leg II: femur 0,
patella pi, tibia 3 (p2), metatarsus 13, tarsus 8. Leg III: femur
0, patella pd6, tibia 7 (p3 r2), metatarsus 19 (p6 r2), tarsus
10. Leg IV: femur 0, patella 0, tibia 1, metatarsus 13, tarsus
15. — Abdomen. Posterior lateral spinnerets with short apical
segment. Lengths: total 5.24; basal segment 2.21, middle
1.31, apical 1.72. Apical segment width 0.88. — Genitalia.
Basal two thirds of spermathecae wide, apical third narrower.
Length 1.09, width 0.72.

Distribution. Southwestern region of the Great Dividing
Range in New South Wales (Fig. 44).

Hadronyche tambo n.sp.

Figures 44-46; Table 13

Etymology. The specific epithet is taken from the Tambo
River in Gippsland, Victoria.

Types. Holotype male: AMS KS8341, 10km north of Bairnsdale, Victoria,
37°49'S 147°37'E, February 1980, C. Brimblecombe. Paratypes. Victoria.
Males: AMS KS10595,3km NW. of Bairnsdale, 13 January 1983, N. Barton;
AMS KS 10561, Butchers Ridge, via Buchan, 37°15'S 148°14'E, December
1982, N. Barton; AMS KS 14336, Granite Rock, NE. of Bairnsdale, 37°45'S
147°39'E, 28 April 1984; K11081 (MV), Nowa Nowa, Vic., 37°44'S
148°06'E, 23 November 1964; AMS KS 16275, Bairnsdale area, 37°50'S
147°37’E, Dec 1985; AMS KS8362, Sarsfield, 37°45'S 147°43'E.

Diagnosis. Cl 8.23-9.59 (male). Differs from H. modesta, H.
meridiana, H.jensenae by leg II without apophysis/swelling
(Fig. 46H-K) and labium long (LL/LW 0.94) (Fig. 45A);
from H. marracoonda by tibia and metatarsus II not bowed
(ventrally concave) (Fig. 461,K): from infensa group species
by presence of spines on femur I, II (but may be absent).

Male (holotype) — Size. Carapace length 9.59, width 8.41.
Abdomen length 9.25, width 6.70. — Colour. Basic colour
pattern. — Carapace. Longer than wide, cephalic area
moderately raised. Height 4.35. Frontal width 6.62. Fovea

procurved. Mid-dorsal cephalic setae almost reach fovea.
Anterior strial setae few to absent. Anterolateral carapace
angle with many strong bristles. — Eyes. Central eye region
raised. Eye group width 2.20. Median ocular quadrangle
length 0.84, anterior width 1.12, posterior width 1.43.
Diameters: AME 0.38, ALE 0.50, PLE 0.37, PME 0.28. —
Chelicerae. Cheliceral groove moderately short and wide,
margins diverging distally. Groove length 2.64, width 0.76.
Cheliceral teeth: 18 central, occupying full length of groove;
8 prolateral; 9 retrolateral. — Labium. About as long as
wide, apical indentation weak to absent. Length 1.84, width
1.88. Labiosternal suture narrowed toward midline. Weak
transverse groove behind cuspules. Cuspules c. 171, number
moderate to low. — Sternum. Ovoid. Length 5.81, width 4.27.
Posterior sigilla elongate, broad. Anterolateral angles of
sternum with weakly grouped bristles. — Palp. Tegular area
wider than long. Embolus of moderate length, gently curved,
weakly to moderately offset from tegulum; shaft strongly
tapered distally. Distal embolus weakly twisted, slender,
flange narrow. Bulb length 3.40, width 1.28. Embolus length
2.33, midwidth 0.19. Length of femur 4.29, patella 1.58,
tibia 4.45. Width of tibia 1.91. Spination: no spines; a few
prolateral tibial bristles, several sinuous bristles on distal
femur. — Legs. 1423. Legs strongly hirsute.

Figure 45. Hadronyche tambo, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) leg II,
prolateral (NB, dorsal femoral spines present or absent); ( D )
cheliceral groove teeth; (E) cephalothorax and chelicerae, dorsal;
(F) spinnerets; (G) palp, prolateral. Scale lines 1 mm.

336

Records of the Australian Museum (2010) Vol. 62

Figure 46. Hadronyche tambo, male ( A,D-K , AMS KS16275; B,C, K11081 [MV]): (A) palp, prolateral; (B) bulb; (C) embolus, distal;
(. D,E ) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; ( H,I) distal patella and tibia II— H,
ventral, /, prolateral; (J,K) metatarsus II— J, ventral, K, prolateral. Scale lines: 1.0 mm, except FJ-K 0.3 mm and G 0.2 mm.

Gray: Australian funnel-web spiders

337

Table 13. Male morphological data —Hadronyche tambo (n = 4).

character

range

mean

character

range

mean

character

ratio

SD

CL

8.23-9.59

8.66

MtlS

13-41

27

CW/CL

0.96

0.017

CW

7.46-8.84

8.12

TalS

5-15

11

CH/CW

0.49

0.017

CH

3.61-4.42

4.08

Ti2S*

4-18

12

CFW/CL

0.74

0.038

CFW

5.80-7.07

6.42

STC2teeth

10-11

11

CHGW/L

0.26

0.035

ChGL

2.22-2.78

2.48

BulbL

3.17-3.77

3.38

SW/SL

0.77

0.028

ChGW

0.53-0.76

0.66

EmbL

2.18-2.65

2.35

LL/LW

0.94

0.04

ChGCT

18-24

20

BulbW

1.14-1.28

1.21

PLSAPW/L*

0.35

0.042

LL

1.43-1.90

1.67

EmbmidW

0.16-0.21

0.19

BulbW/EmbL 0.52

0.029

LW

1.56-1.98

1.76

PalpTibL

3.94-4.76

4.39

EmbmidW/L* 0.08

0.013

CUSP*

149-184

165

PalpTibW

1.66-1.92

1.80

PalpTibW/L

0.41

0.025

SL

4.74-5.99

5.47

PalpTibS

0-1

<1

BulbL/TibL

0.77

0.039

SW

3.74-4.59

4.20

PalpPatS

0-1

<1

PLSAPW

0.40-0.60

0.51

PalpFemS

0

PLSAPL

1.32-1.69

1.44

FelS*

0-1

<1

PalS

0-1

<1

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.27

4.02

6.68

6.29

4.10

29.36

2

7.46

3.77

6.31

5.90

3.94

26.98

3

6.29

3.28

4.39

5.13

4.12

23.21

4

8.04

3.62

6.37

5.96

4.42

28.41

Tibia I width 1.46. Femur I usually without spines; dorsal
spines present or absent on femur n. Patella III with few spines.
Tibia II without apophysis, weakly sinuous; ventral spines
few, scattered, larger proximally. Metatarsi I and II ventrally
flexed basally, slightly concave dorsally; metatarsus II without
apophysis. Anterior coxae with some basally thickened setae
on frontal coxal surfaces. Tarsal scopulae: leg I weak; legs
II-IV moderate to strong. Distal metatarsal scopulae: legs
I, II absent; legs III, IV moderate to strong. Metatarsus IV
weakly curved and dorsally concave distally, ventral scopula
occupying most of distal half of segment. Tarsal claw teeth
legs I, II: superior 11, 11; inf erior 2, 3. Trichobothria legs I,
II: tarsus 14, 13; metatarsus 17, 15; tibia p7 r7, p7 r7. —Leg
spination. Leg I: femur 0, patella 1, tibia 20, metatarsus 25,
tarsus 15. Leg II: femur d4, patella 2, tibia 10, metatarsus 23,
tarsus 17. Leg III: femur 1 (strong bristles present), patella
p2, tibia 13 (p5 r3), metatarsus 23, tarsus 22. Leg IV: femur
0 (strong bristles), patella 0, tibia 4, metatarsus 19, tarsus
27. — Abdomen. Posterior lateral spinnerets with moderately
long apical segment. Lengths: total 3.72; basal segment 1.52;
middle 0.82; apical 1.38. Apical segment width 0.50.

Distribution. Bairnsdale/Buchan region of Gippsland,
Victoria (Fig. 44).

Comments. Dorsal spines are usually absent on leg I and
present on leg II.

Hadronyche monaro n.sp.

Figures 47-49; Table 14

Etymology. The specific epithet refers to the Monaro region
of New South Wales.

Types. Holotype male: AMS KS8744, Sawpit Creek, Kosciuszko National
Park, New South Wales, 28°22'S 152°49'E, J. Gold, February 1982.
Paratypes (all AM). New South Wales. Males : AMS KS6647, Sawpit Creek,

Kosciuszko National Park, 28°22'S 152°49'E, 27 January 1981, J. Robson;
AMS KS1571, Braemer Caravan Park, Lake Eucumbene, 35°58'S 148°39E,
6 December 1977, G. Goodfellow.

Diagnosis. CL 6.23-6.43 (male). Small Atracinae. Differs
from most Hadronyche species by having tibia II unmodified,
but metatarsus II with a small, low midventral apophyseal
swelling (Figs. 47G; 481,K). Differs from H. jensenae by
labium almost as long as wide (Fig. 47A) and distal embolus
not at angle to shaft (Fig. 48B,C).

Figure 47. Hadronyche monaro, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
(D) cheliceral groove teeth; (E) cephalothorax and chelicerae,
dorsal; (F) palp, prolateral; (G) leg II, prolateral. Scale lines 1 mm.

338

Records of the Australian Museum (2010) Vol. 62

Figure 48. Hadronyche monaro, male ( A,D-K , AMS KS1571, RHS; B,C, AMS KS6646): (A) palp, prolateral; (B) bulb; (C) embolus,
distal; ( D,E) distal patella and tibia I—D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; (H,I) distal patella and tibia
II— H, ventral, I, prolateral; ( J, K) metatarsus II—/, ventral, K, prolateral. Scale lines: 0.2 mm, except A 0.3 mm and / 0.1 mm.

Gray: Australian funnel-web spiders

339

Table 14. Male morphological data— Hadronyche monaro (n = 3).

character

range

mean

character

range

mean

character

ratio

CL*

6.23-6.43

6.33

MtlS

23-30

27

CW/CL

0.97

CW

6.08-6.20

6.14

TalS*

7-9

8

CH/CW

0.49

CH

2.95-3.08

3.02

Ti2S*

7-11

9

CFW/CL

0.74

CFW

4.65-4.67

4.66

STC2teeth

10-11

11

CHGW/L

0.21

ChGL

1.85-1.94

1.90

BulbL

2.05-2.19

2.14

SW/SL

0.84

ChGW

0.37-0.41

0.39

EmbL

1.34-1.42

1.38

LL/LW*

0.97

ChGCT

19-26

23

BulbW

0.81-0.93

0.87

PLSAPW/L

0.43

LL

1.02-1.28

1.15

EmbmidW

0.18-0.20

0.19

BulbW/EmbL

0.63

LW

1.07-1.29

1.18

PalpTibL

2.50-3.00

2.85

EmbmidW/L

0.14

CUSP*

143-226

185

PalpTibW

1.46-1.50

1.48

PalpTibW/L

0.52

SL

3.86-4.08

3.96

PalpTibS

0

—

BulbL/TibL

0.75

SW

3.22-3.42

3.32

PalpPatS

0

—

PLSAPW

0.32-0.46

0.39

PalpFemS

0-3

2

PLSAPL

0.76-1.06

0.91

FelS

2-9

6

PalS

2-6 (pi-2)

5

Male (holotype) — Size. Carapace length 6.23, width 6.08.
Abdomen length 7.24, width 5.23. — Colour. Basic colour
pattern. — Carapace. Slightly longer than wide, cephalic
area moderately raised. Height 2.95. Frontal width 4.67.
Fovea procurved, anterior margin weakly notched. Mid¬
dorsal cephalic setae do not reach fovea. Anterior strial
setae absent. Anterolateral carapace angle with small, weak
bristles. — Eyes. Central eye region slightly raised. Eye group
width 1.76. Median ocular quadrangle length 0.66, anterior
width 0.80, posterior width 1.14. Diameters: AME 0.20,
ALE 0.33, PLE 0.20, PME 0.16. — Chelicerae. Cheliceral
groove short, of medium width, margins diverging distally.
Groove length 1.85, middle width 0.41. Cheliceral teeth:
19 central, occupying full length of groove; 9 prolateral;
9 retrolateral. — Labium. Almost as long as wide, not
indented apically. Length 1.28, width 1.29. Labiosternal
sigilla narrowed toward midline. Cuspules c. 226, number
moderate. — Sternum. Ovoid. Length 4.08, width 3.22.
Posterior sigilla oval, broad. — Palp. Tegular area wider than
long. Embolus of moderate length and width; shaft straight
to slightly curved, weakly tapered distally. Distal part of
embolus weakly twisted and flange rather short. Embolus
slightly offset from tegulum. Bulb length 2.19, width 0.93.
Embolus length 1.42, midwidth 0.18. Length of femur 2.86,
patella 1.40, tibia 3.00. Width of tibia 1.46. Spination: femur
2, patella 0, tibia 0. A few sinuous bristles on distal femur.
— Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.04

2.50

4.12

3.77

2.26

17.69

2

4.76

2.38

3.76

3.47

2.26

16.63

3

4.22

1.81

2.61

3.33

2.31

14.28

4

5.03

2.38

3.69

4.22

2.85

18.17

Tibia I width 0.94. Femora I and II with dorsal spines. Leg
I unmodified. Tibia II without apophysis, weakly excavated
distoventrally; ventral spines grouped in proximal half.
Metatarsus II weakly sinuous with a small spined mid-
ventral apophyseal swelling. Metatarsus IV slightly enlarged
distally, concavely arched dorsally. Anterior coxal hairs
normal. Tarsal scopulae: legs I, II weak; legs III, IV moderate
to strong. Distal metatarsal scopulae: legs I—III absent to
weak; leg IV moderately developed, extending to midventral
metatarsus. Tarsal claw teeth legs I, II: superior 11, 10,

inferior 1,0. Trichobothria legs I, II: tarsus 11,8; metatarsus
13, 10; tibia p6 r6, p5 r5. — Leg spination. Leg I: femur d9,
patella 2 (pi vl), tibia 25, metatarsus 23, tarsus 9. Leg II:
femur d6, patella p2, tibia 7, metatarsus 13, tarsus 9. Leg III:
femur d2, patella 12 (plO r2), tibia 17 (p4 r4), metatarsus
25, tarsus 13. Leg IV: femur d4 (bristle-like), patellarl, tibia
6 (pi r3), metatarsus 23, tarsus 21. — Abdomen. Sparsely
covered with weak bristles and hairs. Posterior lateral
spinnerets with medium length apical segment. Lengths: total
2.34; basal segment 1.00, middle 0.58; apical 0.76. Apical
segment width 0.32.

Distribution. East of the Snowy Mountains in the
Eucumbene/Jindabyne region of New South Wales (Fig. 49).

Hadronyche mascordi n.sp.

Figures 49-52; Tables 15, 34

Etymology. The species is named for Ramon Mascord—
author, photographer and student of Australian spiders.

Types. Holotype male: AMS KS2065, Somersby, via Gosford, New South
Wales, 33°21'S 151°17'E, L. Abra, October 1978. Paratypes (all AM).
New South Wales. Males : AMS KS13393, Kulnura, 33°13'S 151°13'E, 25
October 1983; AMS KS13401, Kulnura, 33°13’S 151°13’E, September 1983,
Nelson and Rodgers; AMS KS4102, Somersby, 33°21'S 151°17'E, 3 October

Figure 49. Collection records for Hadronyche monaro (squares)
and H. mascordi (circles)

340

Records of the Australian Museum (2010) Vol. 62

1979, R. Mascord; AMS KS8571, Somersby, 33°21'S 151°17'E, 12 October
1981, L. Abra; AMS KS4108, Somersby, 33°21'S 151°17’E, 3 October 1979,
K. Mascord; AMS KS991, Somersby, 33°21'S 151°17E, November 1956,
J.V. Brown. AMS KS7335, Somersby, 33°21'S 151°17E, 12 September
1956, J. Brown. Females : AMS KS15749, Kariong, near Gosford, 33°26’S
151°18E, L. Abra, October 1984; AMS KS14381, Kulnura, 33°13’S
151°13E, 23 March 1984, M. Gray; AMS KS14757, Kariong, 33°26'S
151°18E, 4 June 1984; AMS KS14234, Kulnura, 33°13’S 151°13E, 10
April 1984, B. Thomas; AMS KS 14233, Kulnura, 33°13’S 15F13E, 10
April 1984, B. Thomas; AMS KS14551, Kulnura, 33°13'S 151°13E, 21
March 1984, E. Thompson; AMS KS14226, Kulnura, 33°13'S 151°13E,
5 April 1984; AMS KS14547, Kariong, 33°26'S 151°18E, 13 April 1984.

Other material examined. New South Wales (males):

AMS KS13394, Kulnura, 25 October 1983; AMS KS13399,
Kulnura, September 1983; AMS KS 13400, Kulnura,
September 1983; AMS KS13377, Somersby, 14 September
1983, K. Gould; AMS KS1929, Somersby, October 1978, L.
Abra; AMS KS23333, Kulnura-Somersby area.

Diagnosis. CL 5.54-6.68 (male). Small Atracinae. Differs
from other atracine species in having a short labium (LW/
LL 0.64) that is fused with the sternum posteromedially
(Figs. 50A, 51 A).

Male (holotype) — Size. Carapace length 6.19, width 5.68.
Abdomen length 6.07, width 4.51. — Colour. Basic colour
pattern, dark brown to black. Abdomen with 4-5 pairs of
small distinct chevron markings on lateral dorsal abdomen.
— Carapace. Slightly longer than wide, strongly raised.
Height 3.01. Frontal width 4.63. Fovea procurved. Mid¬
dorsal cephalic setae numerous, reach fovea. Anterior strial
setae present. Anterolateral angle of carapace with strong
bristles. — Eyes. Sessile. Eye group width 1.74. Median
ocular quadrangle length 0.62, anterior width 0.79, posterior

Figure 50. Hadronyche mascordi, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cheliceral
groove teeth; (D) cephalothorax and chelicerae, dorsal; (E) leg II,
prolateral; ( F ) palp, prolateral; (G) spinnerets. Scale lines 1 mm.

width 1.22. Diameters: AME 0.25, ALE 0.42, PLE 0.33,
PME 0.30. — Chelicerae. Anterodorsal paturon bristles
rather short. Cheliceral groove moderately short and wide,
margins diverging distally. Groove length 1.69, middle
width 0.42. Cheliceral teeth: 7 central, running full length
of groove; 8 prolateral; 6 retrolateral. — Labium. Short—
length 0.98, width 1.39. Labium fused posteromedially with
sternum. Labiosternal sigilla reduced to a weak, transverse
line suture flanked by two small lateral sigilla. Cuspules c.
209, covering most of ventral labium, number moderate.
— Sternum. Ovoid. Raised anteriorly and continuous with
labium. Length 3.81, width 2.82. Posterior sigilla very small,
ovoid to elongate. — Palp. Tegular area slightly wider than
long. Embolus shaft short, tapering, weakly curved. Distal
embolus weakly to moderately twisted, often with a subdistal
“notch”. Embolus moderately offset from tegulum. Bulb
length 2.10, width 0.84. Embolus length 1.34, midwidth
0.15. Length of femur 2.70, patella 1.40, tibia 2.68. Width
of tibia 1.22. Spination: femur 1; few weak sinuous bristles
on distal femur. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.11

2.38

3.81

3.03

1.97

16.30

2

4.88

2.42

3.90

2.69

1.89

15.78

3

4.10

1.91

2.95

3.03

2.13

14.12

4

5.08

2.15

4.04

3.81

2.50

17.58

Tibia / width 0.88. All femora with dorsal spines. Metatarsus

I normal to slightly thickened proximally. Tibia II lacking
apophysis, spines centrally-proximally placed. Metatarsus

II weakly sinuous, with a small mid-ventral swelling.
Coxal hairs normal. Tarsal and distal metatarsal scopulae:
weak to absent on legs I, II; well developed on legs III, IV.
Tarsal claw teeth legs I, II: superior 13, 13; inferior 6, 5.
Trichobothria legs I, II: tarsus 8, 9; metatarsus 12, 10; tibia
p6 r6, p6 r6. — Leg spination. Leg I: femur d4, patella 0,

Figure 51. Hadronyche mascordi, female: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C)
spermathecae; ( D ) spinnerets; (E) cephalothorax and chelicerae,
dorsal; ( F ) leg III, prolateral, patella, tibia, metatarsus; (G)
cheliceral groove teeth. Scale lines 1 mm.

Gray: Australian funnel-web spiders

341

Figure 52. Hadronyche mascordi, male {A,D-K, AMS KS23333; B,C, AMS KS7335): (A) palp, prolateral; ( B ) bulb; (Q embolus, distal;
(. D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; (H,I) distal patella and tibia II— H,
ventral, I, prolateral; ( J,K) metatarsus II— J, ventral, K, prolateral. Scale lines: A,E,F,1, 0.3 mm, B 1.0 mm, D, G,H 0.2 mm and J,K 0.1 mm.

342

Records of the Australian Museum (2010) Vol. 62

Table 15. Male morphological data— Hadronyche mascordi (n = 8).

character

range

mean

character

range

mean

character

ratio

SD

CL*

5.54-6.68

6.27

MtlS

22-43

33

CW/CL

0.97

0.061

CW

5.60-6.26

5.96

TalS

9-24

14

CH/CW*

0.50

0.018

CH

2.83-3.22

3.00

Ti2S*

8-13

10

CFW/CL

0.77

0.036

CFW

4.63-5.33

4.81

STC2teeth

13-14

14

CHGW/L

0.22

0.022

ChGL

1.69-1.86

1.80

BulbL

1.95-2.13

2.06

SW/SL

0.75

0.026

ChGW

0.33-0.46

0.40

EmbL

1.25-1.39

1.32

LL/LW*

0.64

0.051

ChGCT

7-18

12

BulbW

0.84-0.95

0.89

PLSAPW/L*

0.51

0.066

LL

0.8-1.03

0.94

EmbmidW

0.12-0.16

0.14

BulbW/EmbL*

0.68

0.033

LW

1.39-1.58

1.47

PalpTibL

2.68-3.00

2.88

EmbmidW/L

0.11

0.008

CUSP

194-260

223

PalpTibW

1.22-1.41

1.31

PalpTibW/L

0.45

0.01

SL

3.73-4.35

3.95

PalpTibS

0

—

BulbL/TibL

0.71

0.031

SW

2.70-3.05

2.94

PalpPatS

0

—

PLSAPW

0.33-0.41

0.37

PalpFemS

1-3

2

PLSAPL

0.60-0.87

0.75

FelS

2-5

4

PalS*

0

0

tibia 10, metatarsus 30, tarsus 11. Leg II: femur d5, patella
pi, tibia 10, metatarsus 13, tarsus 15. Leg III: femur d2,
patella p4, tibia 10 (p5 r2), metatarsus 19, tarsus 18. Leg
IV: femur d5, patella 0, tibia 6, metatarsus 20, tarsus 16.
— Abdomen. Posterior lateral spinnerets with short apical
segment. Lengths: total 1.99; basal segment 0.76, middle
0.50, apical 0.73. Apical width 0.38.

Female (paratype AMS KS15749) — Size. Carapace length
7.14, width 7.04. Abdomen length 12.50, width 8.78.
— Colour. Basic colour pattern. — Carapace. Almost as
wide as long; strongly raised, broad frontally. Height 3.81;
frontal width 6.05. Cephalic length 5.44. Fovea procurved.
Mid-dorsal cephalic setae reach fovea. Anterior strial setae
present. Anterolateral carapace angle with a few bristles.
— Eyes. Central eye region slightly raised. Eye group width
2.11. Diameters: AME 0.22, ALE 0.46, PLE 0.34, PME
0.28. Interdistances: AME-AME 0.37, AME-ALE 0.20,
ALE-PLE 0.26, PLE-PME 0.14, PME-PME 0.95. Median
ocular quadrangle length 0.73, anterior width 0.91, posterior
width 1.45. — Chelicerae. Cheliceral groove margins diverge
distally. Groove length 2.31, middle width 0.65. Cheliceral
teeth: 22 central in a single row occupying full length of
groove; 8 prolateral; 6 retrolateral. — Labium. Short (length
1.09, width 1.67), fused with sternum as in male; anterior
margin not indented. Cuspules distributed over most of
ventral labium. — Sternum. Ovoid, fused anteriorly with
labium, surface strongly convex. Length 4.62, width 3.81.
Posterior sternal sigilla small, narrow and elongate: length
0.61, width 0.21. — Palp. Spination: tibia 0 (strong bristles),
tarsus 6. Tarsal claw with 8 teeth. Trichobothria: tibia p5 r6,
tarsus 10. — Legs. 4123. Legs I and IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.66

2.99

4.11

3.32

2.21

18.29

2

5.10

2.79

3.88

3.18

2.18

17.13

3

4.22

1.97

2.48

2.89

2.24

13.80

4

5.58

2.58

3.89

3.77

2.48

18.30

Tibia / width 1.20. Metatarsus I proximal width 0.86. Coxal
setae unmodified. Tarsal claw teeth legs I, II: superior 10,
10; inferior 5, 5. Trichobothria legs I, II: tarsus 11, 11;
metatarsus 13,10; tibia p6 r6. — Leg spination. Leg I: femur
0, patella 0, tibia 1, metatarsus 6, tarsus 5. Leg II: femur 0,
patella 0, tibia 2, metatarsus 11, tarsus 7. Leg III: femur 0,

patella p4, tibia 8 (p4 r2), metatarsus 23 (p6 r3), tarsus 13.
Leg IV: femur 0, patella 0, tibia 3, metatarsus 21, tarsus
21. — Abdomen. Sparse cover of weak hairs and bristles.
Posterior lateral spinnerets with short apical segment.
Lengths: total 2.58, basal segment 1.53, middle 0.82, apical
0.82. Apical segment width 0.55. — Genitalia. Spermathecae
short, bluntly conical, lateral margins sloping medially; wide
basally, separated by about half basal width. Length 0.75,
width 0.68.

Distribution. Kulnura/Somersby region, Central Coast, New
South Wales (Fig. 49).

Comments. A female spider from Kulnura, NSW was
collected (apparently dead) in her burrow with 14 young
(KS 14551). These “spiderlings” were remarkable because
of their large size—their mean carapace length was 4.0 mm,
compared with 8.0 mm for the female.

Hadronyche nimoola n.sp.

Figures 53-56; Tables 16, 34

Etymology. The specific epithet is taken from an aboriginal
word for a steep slope, a reference to the caput profile of
this species.

Types. Holotype male: AMS KS867, Batemans Bay, New South Wales,
35°43'S 150°13'E, J.R. Mylott, 3 March 1973. Paratypes (all AM). New
South Wales Males : AMS KS2276 Benandarah State Forest, 35°39'S
150° 15'E, 30 November 1978-4 January 1979, C. Horseman; AMS KS863,
Bermagui, 36°25’S 150°04'E, K. Walker; AMS KS 12124, Bondi State
Forest, nr. Bombala, 37°06’S 149°15E, 26 January 1981, G. Gowing etal .;
AMS KS2665, Bowral, 34°28'S 150°25E, 7 February 1979, G. Goodfellow;
AMS KS2081, Crookwell, 34°27’S 149°28F, 28 November 1978, L.E.
Willis; AMS KS862, Narooma, 36°12'S 150°08F, 14 February 1971,
B. Plunkett-Cole; AMS KS88199, Exeter, 34°36’S 150°19'E, 21 January
2004; AMS KS3876, Narooma, 36°13'S 150°08'E, January 1979, B.H.
Plunkett-Cole. Females : AMS KS4664, Bundanoon, 34°39'S 150°18'E,
February, 1980; AMS KS 16446, Bermagui, 36°25'S 150°04'E, K. Walker;
AMS KS12139, Bondi State Forest, nr. Bombala, 37°06’S 149°15F, 26
January 1981, G. Gowing et al .; AMS KS6163, Bowral, 34°28'S 150°25F,
October 1980, H. Styles; AMS KS6825, Grabben Gullen, 34°32’S 149°24F,
16 December 1980, R. Chown; AMS KS4506, Moruya, 35°54’S 150°06F,
January 1980; AMS KS3625, Narooma, 36°12'S 150°08'E, 15 June 1971,
J. Cobcroft.

Gray: Australian funnel-web spiders

343

Other material examined. New South Wales (males): AMS KS2032,
Benandarah State Forest, 5 October 1978-2 November 1978, C. Horseman;
AMS KS867, Bateman’s Bay, 3 March 1973; AMS KS12150, Bondi State
Forest, near Bombala, 26 January 1981, G. Gowing et al.; AMS KS3460,
Yass River, 30 July 1966, A.B. Thompson; AMS KS6163, Bowral, October
1980, H. Styles; AMS KS13806, Bundanoon, 5 February 1984, W. Nooijen;
AMS KS6825, Grabben Gullen, 16 December 1980, R. Chown; AMS
KS7567, Grabben Gullen, 23 October 1980, R. Chown; AMS KS2117,
Kioloa State Forest, 2 November 1978-30 November 1978, C. Horseman;
AMS KS2293, Kioloa State Forest, 30 November 1978-4 January 1979,
C. Horseman; AMS KS1163, Mittagong, 13 January 1978, M. Johnson;
AMS KS6275, Moss Yale, 27 December 1980, J. Kelly; AMS KS868 &
869, Nerrigundah, December 1945, V. Haskell; AMS KS861, Narooma,
16 November 1974, B. Plunkett-Cole. New South Wales (females): AMS
KS1865, Bega, 24 January 1976, H.W. Kemp; AMS KS1867, Bermagui,
May 1964, K. Walker; AMS KS 11996, Bondi State Forest, near Bombala, 14
November 1980, G. Gowing et ah', AMS KS1872, Bowral, H.H. Florence;
AMS KS1873, Bundanoon, 8 January 1952, N.C. de Meyrick; Merimbula,
5 June 1964 (MY); AMS KS4777, Mittagong, June 1937, A. Livingstone;
AMS KS2961, Narooma, 29 April 1979, A. Brown. Australian Capital
Territory (male): AMS KS 13824, Tidbinbilla Nature Reserve, 9 March
1978, P. Ormay.

Diagnosis. CL 5.85-7.22 (male). Small atracine spiders
without leg II apophyses. Differs from most Hadronyche
spp. by caput being high and wide, almost bulbous (CH/CW:
male 0.53 [Fig. 53A], female 0.61 [Fig. 54B]), and having
relatively few STC teeth (7-9) on tarsi I, II; from H. mascordi
by having an entire labiosternal sigilla; from lamingtonensis
group species by having a long central cheliceral tooth row.

Male (holotype) — Size. Carapace length 7.01, width 7.30.
Abdomen length 8.44, width 6.03. — Colour. Basic colour

and chelicerae, lateral; (B) sternum, labium and maxilla; (C)
cephalothorax and chelicerae, dorsal; ( D ) palp, prolateral; (E) leg
II, prolateral (NB, dorsal femoral spines present or absent); (F)
cheliceral groove teeth; (G) spinnerets. Scale lines 1 mm.

pattern. — Carapace. About as wide as long, strongly raised.
Height 3.61. Frontal width 5.99. Fovea broad, procurved,
anterior margin weakly indented. Mid-dorsal cephalic
setae small, closely spaced, reach, or almost reach, fovea.
Anterior strial setae absent. Anterolateral angle of carapace
with weak bristles. — Eyes. Sessile to slightly raised in
central eye region. Eye group width 2.32. Median ocular
quadrangle length 0.78, anterior width 1.05, posterior
width 1.60. Diameters: AME 0.33, ALE 0.47, PLE 0.37,
PME 0.25. — Chelicerae. Cheliceral groove broad, margins
diverging distally. Groove length 2.28, middle width 0.52.
Cheliceral teeth: 16 central, running full length of groove;
9 prolateral; 11 retrolateral. — Labium. Wider than long.
Length 1.44, width 1.66. Labiosternal sigilla narrowed in
midline. Cuspules c. 234, number moderate. — Sternum.
Ovoid, broad. Setae weak. Length 4.35, width 3.65. Posterior
sigilla elongate, ovoid. — Palp. Tegular area wider than
long. Embolus rather short, strongly tapered distally, weakly
curved. Distal part moderately twisted, embolic groove
sometimes rather wide. Embolus weakly to moderately
offset from tegulum. Bulb length 2.43, width 0.99. Embolus
length 1.61, midwidth 0.18. Length of femur 3.50, patella
1.45, tibia 3.49. Width of tibia 1.64. Spination: femur with
7 sinuous bristle-like spines, tibia with 2 bristle-like spines.
— Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.23

3.01

4.80

4.26

2.65

20.95

2

5.66

2.87

4.43

3.90

2.46

19.32

3

4.82

2.34

3.28

3.62

2.65

16.71

4

6.10

2.88

4.92

4.74

3.15

21.79

Tibia I width 1.15. Femur I with few or no dorsal spines;
femur II usually with spines, occasionally none; femora III
and IV with spines. Legs I and II unmodified, spines small,
scattered. Metatarsus IV thickened distally, concavely arched

Figure 54. Hadronyche nimoola, female: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C)
spermathecae; ( D ) cephalothorax and chelicerae, dorsal; ( E )
cheliceral groove teeth; ( F ) spinnerets; (G) leg III, prolateral,
patella, tibia, metatarsus. Scale lines 1 mm.

344

Records of the Australian Museum (2010) Vol. 62

Figure 55. Hadronyche nimoola, male ( A,D-K , AMS KS88199;
B,C, AMS KS3876): (A) palp, prolateral; (B) bulb; (Q embolus,
distal; ( D,E ) distal patella and tibia I— D, ventral, E, prolateral;
(EG) metatarsus I— F, ventral, G, prolateral; ( H,1 ) distal patella and
tibia II — H, ventral, I, prolateral; (J,K) metatarsus II — J, ventral, K,
prolateral. Scale lines: 0.2 mm, except B 0.5 mm and E,G 0.3 mm.

Gray: Australian funnel-web spiders

345

Table 16. Male morphological data— Hadronyche nimoola (n = 17).

character

range

mean

character

range

mean

character

ratio

SD

CL*

5.85-7.22

6.62

MtlS

6-25

17

CW/CL*

1.01

0.032

CW

6.07-7.46

6.72

TalS

6-22

13

CH/CW*

0.53

0.029

CH*

3.20-4.03

3.56

Ti2S*

4-18

10

CFW/CL*

0.83

0.042

CFW

4.81-6.23

5.5

STC2*teeth

7-9

8

CHGW/L

0.24

0.041

ChGL

1.97-2.56

2.17

BulbL

2.14-2.70

2.38

SW/SL*

0.88

0.034

ChGW

0.42-0.78

0.51

EmbL

1.30-1.75

1.54

LL/LW

0.85

0.053

ChGCT

9-26

17

BulbW

0.90-1.40

0.99

PLSAPW/L

0.45

0.029

LL

1.06-1.60

1.28

EmbmidW

0.12-0.18

0.15

BulbW/EmbL

0.64

0.053

LW

1.34-1.68

1.49

PalpTibL

2.65-3.77

3.21

EmbmidW/L

0.10

0.012

CUSP

160-281

219

PalpTibW

1.44-1.74

1.58

PalpTibW/L

0.49

0.031

SL

3.82-4.67

4.12

PalpTibS

0-2

<1

BulbL/TibL

0.74

0.029

SW

3.25-4.14

3.63

PalpPatS

0

—

PLSAPW

0.38-0.49

0.42

PalpFemS

0-4

1

PLSAPL

0.82-1.20

0.95

FelS

0-5

<1

PalS

0-3

<1

dorsally. Anteromedial hairs on coxae I, II usually thickened
basally. Tarsal and distal metatarsal scopulae moderately
developed on legs I, II; strongly developed on legs III, IV.
Metatarsi IV scopulae extend to mid ventral area. Tarsal
claw teeth legs I, II: superior 7, 8: inf erior 0,0. Trichobothria
legs I, II: tarsus 11, 10, metatarsus 13, 13; tibia p6 r6, p6
r6. — Leg spination. Leg I: femur 0, patella pi, tibia 11 (p3),
metatarsus 25, tarsus 22. Leg II: femur d5, patella pi, tibia
17, metatarsus 32, tarsus 23. Leg III: femur d6, patella 33
(p26 r7), tibia 22 (p7, r2, d2), metatarsus 40, tarsus 27. Leg
IV: femur d7, patella r5, tibia 8 (rl d2), metatarsus 22, tarsus
35. — Abdomen. Sparse, fine hair cover, bristles lacking or
very weak. Posterior lateral spinnerets with moderately
short apical segment. Lengths: total 2.88; basal segment
1.32; middle 0.58; apical 0.98. Apical segment width 0.48.

Female (paratype AMS KS4664) — Size. Carapace length
8.74, width 8.65. Abdomen length 12.24, width 8.30. —
Colour. Basic colour pattern. — Carapace. About as long as
wide, cephalic area broad and very strongly raised. Height
5.08; frontal width 7.84. Cephalic length 6.90. Fovea wide,
deeply procurved, anterior margin indented. Mid-dorsal
cephalic setae variable, may or may not reach fovea. Anterior
strial setae absent. Anterolateral carapace angle with several
weak bristles. — Eyes. Eye region sessile. Eye group width
2.76. Diameters: AME 0.28, ALE 0.47, PLE 0.38, PME
0.24. Interdistances: AME-AME 0.55, AME-ALE 0.43,
A.L.E-PLE 0.40, PLE-PME 0.26, PME-PME 1.36. Median
ocular quadrangle length 0.88, anterior width 1.17, posterior
width 1.88. — Chelicerae. Cheliceral groove wide, margins
diverging distally. Groove length 2.83, middle width 0.74.
Cheliceral teeth: 36 central, occupying full length of groove
in irregular double row; 11 prolateral; 9 retrolateral. —
Labium. Slightly wider than long, anterior margin weakly
or not indented. Length 2.10, width 2.46. Cuspules occupy
anterior two-thirds of labium. Labiosternal sigilla entire,
narrow. — Sternum. Broad, subcircular. Length 5.78,
width 5.08. Anterolateral angles with a few short, basally
thickened, bristle-like setae. Posterior sigilla long, ovoid:
length 1.31, width 0.49. — Palp. Spination: tibia 1, tarsus
8. Trichobothria: tibia p6 r5, tarsus 14. Tarsal claws with 6
teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.11

3.36

4.18

3.28

2.12

19.05

2

5.45

3.20

3.62

3.03

2.26

17.56

3

4.58

2.36

2.59

2.83

2.35

14.71

4

6.15

3.77

4.63

3.98

2.74

21.27

Tibia /width 1.48. Metatarsus I proximal width 1.03. Coxae
I, II with basally thickened and thorn-like setae anteriorly.
Tarsal claw teeth legs I, II: superior 7, 7; inferior 2, 2.
Trichobothria legs I, II: tarsus 11,11, metatarsus 12,11; tibia
p6 r5, p6 r5. — Leg spination. Leg I: femur 0, patella 0, tibia
1, metatarsus 10, tarsus 7. Leg II: femur 0, patella 0, tibia 1,
metatarsus 14, tarsus 7. Leg III; femur 0, patella pd 20, tibia
12 (p6 r3), metatarsus 33 (pl4 r6), tarsus 15. Leg IV: femur
0, patella 0, tibia 1, metatarsus 14, tarsus 7. — Abdomen.
Posterior lateral spinnerets with short apical segment.
Lengths: total 3.95; basal segment 1.79, middle 0.87, apical
1.29. Apical segment width 0.69. — Genitalia. Spermathecae
short, digitiform, well separated basally (by slightly more
than basal spermatheca width); diverge toward rounded and
slightly enlarged apices. Length 0.87, width 0.44.

Distribution. Southern highland and south coastal regions
of New South Wales (Fig. 56).

Figure 56. Collection records for Hadronyche nimoola (squares)
and H. modesta (circles).

346

Records of the Australian Museum (2010) Vol. 62

Hadronyche modesta (Simon)

Figures 56-59; Tables 17, 34

Atrax modesta Simon, 1891: 302. Hogg, 1901: 272. Main,
1985:41.

Hadronyche modesta. -Gray, 1988: 114 (transferred from
Atrax).

Types. Holotype female: MNHN, Paris (not seen). Melbourne, Victoria.

Other material examined. Victoria (males): Ellinbank via Warragul, 11
May 1962, J. Copley (MV); AMS KS8471, Emerald, 3 April 1978; AMS
KS8472, Emerald, 20 April 1979; Leongatha, 6 June 1963,1. Bissett (MV);
AMS KS7292, Back Creek Rd., Gembrook State Forest, near Gembrook,
Victoria, L. Gibson, 10 March 1981); Mirboo North 15 April 1966, G.B.
Burr (MV); 6 km NE. of Mirboo North, 26.iii.1967, J. Seebeck (MV);
AMS KS8473, Mount Eliza, 23 April 1979; AMS KS8930, Mount Tassie,
Strezelecki Ranges, 25 March 1982, R. Waters; AMS KS8933, Tanjil South,
near Moe, February 1982, N. Barton; AMS KS8474, Thorpdale, 10 April
1980; Wright, SE of Emerald, September 1951, C. Oke (MV); Yalloum,
6 February 1963, J. Irving (MV); Yarra Junction, 4 May 1964, J. Mooney
(MV); Yarra Junction 20 April 1964, J. Mooney (MV). Victoria (females):
AMS KS10775, Cockatoo, 10 October 1982, ? McKelvey; AMS KS8477,
Ferny Creek, 6 October 1979; AMS KS12618, Hawthorn, 23 January 1983,

M. Harvey; AMS KS8476, Milford, 26 October 1976; AMS KS8339,
Mirboo North, Apr 1980; AMS KS8932, Moe area, October 1981, N. Barton;
AMS KS8470, Mount Evelyn, Lilydale Shire, 5 June 1979; AMS KS8931,
Newborough, near Moe, March 1982, N. Barton; AMS KS8926, Sassafras,
14 February 1982, N. Wentworth; AMS KS10560, Warragul, January 1983,

N. Barton; AMS KS3386, Yinnar, 9 July 1979, N. Barton.

Diagnosis. CL 5.48-7.30 (male). Small atracine spiders with
leg II apophyseal swellings (Figs. 57G, 59H-K). Differs from
H. meridiana and H. jensenae by having a short, straight
embolus but with the distal part neither twisted nor angled,
respectively (Fig. 59B,C). Differs from other Hadronyche
spp. with leg II apophyses/swellings by having a relatively
short labium (LL/LW 0.71) (Fig. 57E).

Figure 57. Hadronyche modesta, male: (A) cephalothorax and
chelicerae, dorsal; ( B ) cephalothorax and chelicerae, lateral; (C)
cheliceral groove teeth; (D) spinnerets; (E) sternum, labium and
maxilla; ( F ) palp, prolateral; (G) leg II, prolateral. Scale lines 1 mm.

Male (AMS KS8475, Lilydale, 14 May 1980). —Size.
Carapace length 6.40, width 6.10. Abdomen length 5.85,
width 6.12. — Colour. Basic colour pattern. — Carapace.
Slightly longer than wide, cephalic area moderately raised.
Height 2.68, frontal width 4.16. Fovea slightly procurved.
Mid-dorsal cephalic setae few, do not reach fovea. Anterior
strial setae absent. Anterolateral carapace angle with a
few very weak bristles. — Eyes. Central eye region raised.
Eye group width 1.82. Median ocular quadrangle length
0.64, anterior width 0.86, posterior width 1.14. Diameters:
AME 0.28, ALE 0.32, PLE 0.22, PME 0.14. —Chelicerae.
Cheliceral groove margins subparallel to weakly divergent.
Groove length 2.06, middle width 0.34. Cheliceral teeth:
23 central, occupying full length of groove; 8 prolateral,
9 retrolateral. Central tooth row occasionally shorter. —
Labium. Wider than long, apically indented. Length LOO,
width 1.38. Labiosternal sigilla narrowed toward midline.
Slight transverse groove behind cuspules. Cuspules c. 123,
number low-moderate. — Sternum. Ovoid. Length 4.01,
width 3.28. Posterior sigilla elongate, narrow. — Palp.
Tegular area wider than long. Embolus short and broad,
almost straight, base moderately offset from tegulum; distal
embolus straight, hardly twisted. Bulb length 2.26, width
1.03. Embolus length 1.37, midwidth 0.20. Length of femur
3.06, patella 1.36, tibia 3.18. Width of tibia 1.71. Spination:
femur 3, patella 4, tibia 0. Distal femur spined, sinuous
bristles few to absent. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.16

2.69

4.82

4.45

2.75

20.87

2

5.91

2.61

4.82

4.20

2.75

20.29

3

5.22

2.23

3.76

4.20

2.92

17.33

4

6.32

2.72

5.24

5.44

3.48

23.20

Figure 58. Hadronyche modesta, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
(D) cheliceral groove teeth; (E) cephalothorax and chelicerae,
dorsal; ( F ) spermathecae; (G) leg III, prolateral, patella, tibia,
metatarsus. Scale lines 1 mm.

Gray: Australian funnel-web spiders

347

Figure 59. Hadronyche modesta, male (KS7292): (A) palp, prolateral; ( B ) bulb; (C) embolus, distal; (D,E) distal patella and tibia I— D,
ventral, E, prolateral; (F, G ) metatarsus I— F, ventral, G, prolateral; (FI, I) distal patella and tibia II— H, ventral, I, prolateral; (J,K) metatarsus
II— J, ventral, K, prolateral. Scale lines: 0.3 mm, except B 0.5 mm and F,I,J 0.2 mm.

348

Records of the Australian Museum (2010) Vol. 62

Table 17. Male morphological data— Hadronyche modesta (n = 26).

character

range

mean

character

range

mean

character

ratio

SD

CL*

5.48-7.30

6.24

MtlS

20-49

36

CW/CL

0.96

0.026

CW

5.28-6.85

5.97

TalS*

15-41

28

CH/CW

0.44

0.025

CH

2.36-3.03

2.64

Ti2S

5-19

13

CFW/CL

0.67

0.015

CFW

3.73^4.96

4.20

STC2*

7-10

8

CHGW/L

0.17

0.011

ChGL

1.76-2.24

2.01

BulbL

1.89-2.38

2.13

SW/SL

0.81

0.027

ChGW

0.30-0.40

0.31

EmbL

1.12-1.41

1.28

LL/LW*

0.71

0.055

ChGCT

8-23

19

BulbW

0.91-1.15

1.03

PLSAPW/L*

0.51

0.053

LL

0.76-1.15

0.95

EmbmidW

0.15-0.24

0.17

BulbW/EmbL*

: 0.80

0.048

LW

1.13-1.52

1.37

PalpTibL

2.53-3.59

3.06

EmbmidW/L*

0.15

0.014

CUSP*

104-160

129*

PalpTibW

1.39-1.83

1.55

PalpTibW/L*

0.52

0.023

SL

3.50-4.70

3.93

PalpTibS

0-5

1

BulbL/TibL

0.70

0.024

SW

2.84-3.79

3.19

PalpPatS*

3-9

5

PLSAPW

0.27-0.52

0.36

PalpFemS

3-7

5

PLSAPL

0.53-92

0.70

FelS

3-8

5

PalS

0-4 (pO)

2

Tibia I width 1.14. Femora I and II with dorsal spines. Leg
I unmodified, tibial and metatarsal ventral spines numerous.
Patella I and II lack prolateral spines. Ventral tibia II weakly
swollen and spinose proximally-centrally; with a few
enlarged central spines; distal tibia II concave ventrally and
lacking spines (other than apical pair). Metatarsus II sinuous,
with a small, spined, midventral apophysis. Coxal setae
unmodified. Scopulae legs I-IV: tarsus, weak to moderate;
distal metatarsus, all weak. Tarsal claw teeth legs I, II:
superior 9, 8; inferior 0, 2. Trichobothria legs I, II: tarsus 9,
8; metatarsus 9, 9; tibia p5 r5, p6 r5. — Leg spination. Leg
I: femur d4, patella 1, tibia 30, metatarsus 41, tarsus 32. Leg
II: femur d4, patella 2, tibia 12, metatarsus 32, tarsus 32.
Leg III: femur d2 (plus strong bristles), patella p9, tibia 15
(pi r4), metatarsus 24, tarsus 31. Leg IV: femur 0 (strong
bristles present), patella 0, tibia 12 (r3), metatarsus 21, tarsus
32. — Abdomen. Posterior lateral spinnerets with short apical
segment. Lengths: total 2.01; basal segment 0.94; middle
0.40; apical 0.67. Apical segment width 0.31.

Female (MV K11082), Hoddles Creek, Victoria, 37°50’S
145°36'E, S. Johnson, 18 October 1981). — Size. Carapace
length 7.48, width 7.11. Abdomen length 10.74, width 7.21.
— Colour. Basic colour pattern. Anterodorsal abdominal
sigilla prominent. — Carapace. Slightly longer than wide,
strongly raised. Height 3.60. Frontal width 6.02. Cephalic
length 5.57. Fovea strongly procurved. Mid-dorsal cephalic
setae few, reach only two-thirds way to fovea. Anterior
strial setae absent. Anterolateral carapace angle with a
few hairs, no bristles. — Eyes. Central eye region weakly
raised. Eye group width 1.97. Diameters: AME 0.25, ALE
0.43, PLE 0.27, PME 0.18. Interdistances: AME-AME
0.37, AME-ALE 0.26, ALE-PLE 0.28, PLE-PME 0.30,
PME-PME 0.84. Median ocular quadrangle length 0.71,
anterior width 0.81, posterior width 1.24. — Chelicerae.
Paturon with sparse anterodorsal bristle cover. Cheliceral
groove moderately wide, margins diverging distally. Groove
length 2.65, middle width 0.68. Cheliceral teeth: 18 central,
in single row occupying full length of groove, occasionally
row shorter; 11 prolateral; 9 retrolateral. — Labium. Wider
than long, apically shallowly indented. Length 1.43, width
2.01. Cuspules grouped on anterior half. Labiosternal sigilla
slightly narrowed toward midline. — Sternum. Broad, ovoid.

Length 5.08, width 4.38. Posterior sigilla elongate, length
1.02, width 0.39. — Palp. Spination: tibia 1-3 ventral bristle¬
like spines, tarsus 9. Trichobothria: tibia p5 r4, tarsus 7.
Tarsal claw with 5 teeth. — Legs. 1423.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.08

4.04

5.29

4.11

3.01

23.53

2

6.29

3.13

4.74

3.64

2.41

20.21

3

4.83

2.72

3.01

3.11

2.62

16.29

4

6.30

3.70

4.82

4.32

3.05

22.19

Tibial width 1.40. Metatarsus I proximal width 0.88. Coxal
setae unmodified. Tibia I, II with no to few spines. Tarsal
claw teeth legs I, II: superior 7, 8; inferior 0,0. Trichobothria
legs I, II: tarsus 10, 9; metatarsus 10, 9; tibia p6 r6, p6 r5.
—Leg spination. Leg I: femur 0, patella 0, tibia 0, metatarsus

6, tarsus 12. Leg II: femur 0, patella 0, tibia 0, metatarsus

7, tarsus 16. Leg III: femur 0, patella pdl3, tibia 5(p2 rl),
metatarsus 17(pd4 rd2) tarsus 19. Leg IV: femur 0, patella
0, tibia 0, metatarsus 12, tarsus 24. — Abdomen. Sparse
cover of weak hairs and bristles. Posterior lateral spinnerets
with short apical segment. Length: total 3.16; basal segment
1.50, middle 0.67, apical 0.99. Apical segment width 0.56.
— Genitalia. Spermathecae digitiform, short, apical third
separated from basal two thirds by a slight constriction.
Length 0.85, width. 0.41.

Distribution. Melbourne/Dandenong Range region to
eastern Gippsland, Victoria (Fig. 56).

Hadronyche meridiana Hogg

Figures 60-63; Tables 18, 34

Hadronyche meridiana Hogg, 1902: 122. Original descrip¬
tion by Hogg (1901, 274) as mis-identified male of H.
cerberea.

Types. Holotype male: BMNH 1903.2.10.6 (NHM). Mt. Macedon, Victoria,
37°23'S 144°35'E, H.R. Hogg.

Other material examined. New South Wales (males): AMS KS4503,
Adelong,22July 1980,M.Pearce;AMSKS917,Kunama, lOJanuary 1971;
AMS KS916, “Loch Haven”, Wantagong, 20 June 971, R. Flynn; AMS
KS915, Wantagong, 1 April 1971, R. Hunter; AMS KS913, Wondalga, 23
September 1948, C.H. Jagoe. New South Wales (females): AMS KS4973,

Gray: Australian funnel-web spiders

349

Adelong, 10 January 1980, M. Pearce; AMS KS6053, Rosewood, 20
October 1980, D.M. Fordham. Victoria (males): AMS KS8488, Baranduda,
13 October 1978, A. McDonald; AMS KS10771, Lake Bolac, June 1982,
H. Pamaby; Avenel, 6 September 1963 (MV); Benalla, 20 October 1968
(MV); Euroa, 19 June 1965 (MV); Pranjip, Longwood area, 4 July 1965
(MV); AMS KS8484, Trawool, near Seymour, 15 August 1980, P May
Williams; Violet Town, 5 February 1980, A. Kube (MV); AMS KS8485,
Willowmavin, near Kilmore, 29 April 1974. Victoria (females): Rutherglen
Research Institute; AMS KS13651-3, Shepparton, 15 January 1984, M.
Gardener; Violet Town, 5 February 1980, A. Kube (MV); AMS KS3399,
Hall’s Gap, 26 March 1974, M. Gray.

Diagnosis. CL 5.44-7.04. Small atracine spiders with weak
apophyseal swellings on leg II (Figs. 60F, 62H-J). Differs
from H. modesta and H. jensenae by having embolus apically
twisted (Fig. 62B,C). Differs from other Hadronyche spp.
with leg II apophyses/swellings by having a relatively short
labium (LL/LW 0.78) (Fig. 60D).

Male (AMS KS8490), Avenel, Victoria, 36°54’S 144°12'E,
May 1981). — Size. Carapace length 6.92, width 6.64.
Abdomen length 7.17, width 5.23. — Colour. Basic colour
pattern. — Carapace. Slightly longer than wide, cephalic area
moderately raised. Height 3.12. Frontal width 5.45. Fovea
straight to slightly procurved. Mid-dorsal cephalic setae do
not reach fovea. Anterior strial setae present. Anterolateral
angle of carapace with weak bristles. — Eyes. Central eye
region slightly raised. Eye group width 1.86. Median ocular
quadrangle length 0.75, anterior width 0.96, posterior
width 1.24. Diameters: AME 0.32, ALE 0.46, PLE 0.43,
PME 0.20. — Chelicerae. Cheliceral groove rather narrow,
margins subparallel. Groove length 2.35, middle width 0.38.
Cheliceral teeth: 7 central (6 in basal half, 1 in distal half);
12 prolateral; 12 retrolateral. Central tooth row distribution

Figure 60. Hadronyche meridiana, male: (A) spinnerets; ( B )
cephalothorax and chelicerae, lateral; (C) cheliceral groove teeth;
( D ) sternum, labium and maxilla; (E) cephalothorax and chelicerae,
dorsal; (F) leg II, prolateral; (G) palp, prolateral. Scale lines 1 mm.

varies from basal area only to full length of groove. —
Labium. Wider than long, surface flattened. Length 1.20,
width 1.58. Labiostemal sigilla narrowed centrally. Cuspules
c. 134, number low to moderate. — Sternum. Ovoid. Length
4.20, width 3.24. Posterior sigilla ovoid. — Palp. Tegular area
slightly wider than long. Embolus of moderate length, rather
broad, weakly curved and tapered, with distal part twisted.
Embolus base weakly to moderately offset from tegulum.
Bulb length 2.51, width 0.95. Embolus length 1.67, midwidth
0.30. Length of femur 3.24, patella 1.27, tibia 3.30. Width
of tibia 1.48. Spination: femur 5, patella 1, tibia 3. Sinuous
bristles and spines present on distal femur. — Legs. 4123.
Legs I, 2 subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.15

2.87

4.59

4.48

2.99

21.08

2

5.82

2.81

4.84

4.43

3.05

20.95

3

5.01

2.40

3.52

4.16

3.09

18.18

4

6.03

2.68

4.89

5.09

3.35

22.04

Tibia I width 1.07. Femora I and II with dorsal spines; bristle¬
like spines on femora II, IV. Metatarsus I slightly thickened
near middle, large ventral spines grouped here. Metatarsus
II sinuous with spined midventral apophysis. Tibia II with
weak, spined apophyseal swelling mid ventrally, ventral
spines grouped with 2-4 central spines enlarged. Tibia II
weakly excavated distally. Anterior coxal hairs normal.
Tarsal scopulae: leg I weak; legs II-IV moderate to strong.
Distal metatarsal scopulae: legs I, II weak to absent, legs III,
IV moderate to strong. Tarsal claw teeth legs I, II: superior
12, 11; inferior 2, 2. Trichobothria legs I, II: tarsus 9, 9;
metatarsus 11,8; tibia p5 r5, p6 r5. — Leg spination. Leg I:
femur d5, patella vl, tibia 26, metatarsus 33, tarsus 13. Leg
II: femur d4, patella pi, tibia 9, metatarsus 20, tarsus 13. Leg
III: femur d4 (bristle like), patella p7, metatarsus 17, tarsus
20. Leg IV: femur d5 (bristle like), patella 0, tibia 2 (dl),
metatarsus 10, tarsus 19. — Abdomen. Sparsely covered with
weak bristles and hairs with a pair of larger bristles above
pedicel. Posterior lateral spinnerets with a rather short apical
segment. Lengths: total 2.84; basal segment 1.24; middle
0.74; apical 0.86. Apical segment width 0.46.

Figure 61. Hadronyche meridiana, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cheliceral
groove teeth; (D) spinnerets; (E) cephalothorax and chelicerae,
dorsal; ( F ) leg III, prolateral, patella, tibia, metatarsus; (G)
spermathecae. Scale lines 1 mm.

350

Records of the Australian Museum (2010) Vol. 62

Figure 62. Hadronyche meridiana, male (. A,D-K , AMS KS8485; B,C, Pranjip, Victoria, [MV]): (A) palp, prolateral; (B) bulb; (C) embolus,
distal; ( D,E) distal patella and tibia I—D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; ( H,I) distal patella and tibia
II— H, ventral, /, prolateral; ( J ) metatarsus II, prolateral; (K) femur II, prolateral. Scale lines: 0.2 mm, except B 0.5 mm and D,I,J 0.3 mm.

Gray: Australian funnel-web spiders

351

Table 18. Male morphological data— Hadronyche meridiana (n = 12).

character

range

mean

character

range

mean

character

ratio

SD

CL*

5.44-7.04

6.37

MtlS

18-37

27

CW/CL

0.98

0.035

CW

5.51-6.64

6.26

TalS

7-22

14

CH/CW

0.50

0.026

CH

2.68-3.53

3.10

Ti2S

8-20

12

CFW/CL

0.73

0.047

CFW

4.15-5.45

4.68

STC2teeth

9-12

10

CHGW/L

0.16

0.011

ChGL

1.96-2.48

2.20

BulbL

2.11-2.54

2.68

SW/SL

0.78

0.028

ChGW

0.27-0.42

0.35

EmbL

1.36-1.67

1.54

LL/LW*

0.78

0.02

ChGCT

3-16

10

BulbW

0.83-1.00

0.93

PLSAPW/L

0.47

0.055

LL

0.96-1.24

1.09

EmbmidW

0.15-0.30

0.20

BulbW/EmbL* 0.60

0.033

LW

1.19-1.58

1.38

PalpTibL

2.74-3.30

3.07

EmbmidW/L

0.14

0.019

CUSP*

117-191

157

PalpTibW

1.26-1.53

1.41

PalpTibW/L

0.46

0.024

SL

3.52-4.40

4.02

PalpTibS

2-7

5

BulbL/TibL

0.77

0.031

SW

2.89-3.40

3.15

PalpPatS

1-2

1

PLSAPW

0.31-0.46

0.35

PalpFemS

5-7*

6

PLSAPL

0.68-0.98

0.70

FelS

3-9

6

PalS

0-7 (p0-3)

4

Female (K11083 MV), Strathbogie Range, Victoria, 36°56'S
145°38'E, 7 March 1982) —Size. Carapace length 7.83,
width 7.07. Abdomen length 11.02, width 7.48. — Colour.
Basic colour pattern. Abdominal chevrons distinct, well
separated. — Carapace. Slightly longer than wide, strongly
raised. Height 4.01; frontal width 6.60. Cephalic length
5.60. Fovea wide, procurved anterior margin indented. Mid¬
dorsal cephalic setae do not reach fovea. Anterior strial setae
absent basally. Anterolateral carapace angle with a few weak
bristles. — Eyes. Central eye region raised. Eye group width
2.11. Diameters: AME 0.27, ALE 0.41, PLE 0.34, PME
0.23. Interdistances: AME-AME 0.36, AME-ALE 0.25,
ALE-PLE 0.21, PLE-PME 0.17, PME-PME 0.96. Median
ocular quadrangle length 0.70, anterior width 0.89, posterior
width 1.43. — Chelicerae. Cheliceral groove margins diverge
weakly distally. Groove length 2.65, middle width 0.54.
Cheliceral teeth: 26 central, in single row occupying full
length of groove; 13 prolateral; 11 retrolateral. Central tooth
row length variable. — Labium. Wider than long, anterior
margin slightly concave, sides rounded. Length 1.55, width
2.11. Cuspules occupying anterior two-thirds of labium.
Labiosternal sigilla narrowed at midline. — Sternum. Ovoid.
Length 5.10, width 4.01. Posterior sternal sigilla elongate,
removed from margins: length 0.99, width 0.34. — Palp.
Spination: tibia 0-1 (strong bristles present), tarsus 8.
Trichobothria: tibia p6 r6, tarsus 8. Tarsal claw with 4 teeth.
— Legs. 4123. Legs I, IV subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.92

2.82

4.59

3.60

2.37

19.30

2

5.10

2.62

3.94

3.47

2.38

17.51

3

4.60

2.55

2.84

3.12

2.52

15.91

4

5.71

2.96

4.08

4.09

2.92

19.76

Tibia / width 1.29. Metatarsus I proximal width 0.95. Coxae
I, II with a few basally thickened setae frontally. Tibia I, II
with few to no spines. Tarsal claw teeth legs I, II: superior
7, 7; inferior 3, 3. Trichobothria legs I, II: tarsus 9, 10;
metatarsus 14, 12; tibia p6 r6, p6 r5. — Leg spination. Leg
I; femur 0, patella 0, tibia 0, metatarsus 5, tarsus 4. Leg II:
femur 0, patella 0, tibia 0, metatarsus 7, tarsus 5. Leg III:
femur 0, patella p7, tibia 9 (p5 r2), metatarsus 15 (p5 rd2),
tarsus 12. Leg IV: femur 0, patella 0, tibia 0, metatarsus 10,
tarsus 14. — Abdomen. Posterior lateral spinnerets stout,

apical segment short. Lengths, total 3.07; basal segment
1.43; middle 0.69; apical 0.95. Apical segment width 0.56.
— Genitalia. Spermathecae very short and broad. Apical
area narrower than expanded central region. Spermatheca
length 0.82, width 0.68.

Distribution. Western side of Great Dividing Range in
Southern New South Wales and northern Victoria (Fig. 63).
This species is provisionally recorded from the Grampian
Ranges on the basis of a single female from the Hall’s Gap
area—male records are needed to confirm this.

Comments. In some males a few mid row or distal teeth
may be missing, making keying such individuals difficult.

Hadronyche jensenae n.sp.

Figures 63-65; Table 19

Etymology. The species is named for my wife, Greta Jensen
Gray.

Types. Holotype male: K11085 (MV), Woodside, Victoria, 38°32'S
146°53'E, 15 August 1966. Paratypes. Victoria. Males : K11086 (MV),
Woodside, data as for holotype; K11087 (MV), Morwell, 38°14'S 146°24'E„
28 August 1967.

Figure 63. Collection records for Hadronyche meridiana (squares)
and H. jensenae (circles).

352

Records of the Australian Museum (2010) Vol. 62

Table 19. Male morphological data— Hadronyche jensenae (n = 4).

character

range

mean

character

range

mean

character

ratio

SD

CL*

4.72-6.60

5.77

MtlS

14-27 (pO)

19

CW/CL

0.99

0.029

CW

4.84-6.60

5.72

TalS

6-10

8

CH/CW

0.45

0.03

CH

2.0-3.16

2.60

Ti2S*

5-7 (pO)

6

CFW/CL

0.70

0.043

CFW

3.36-4.85

4.06

STC2teeth

10-11

11

CHGW/L

0.16

0.013

ChGL

1.66-2.18

1.94

BulbL

1.25-2.34

2.17

SW/SL

0.81

0.022

ChGW

0.27-0.34

0.31

EmbL

1.15-1.36

1.28

LL/LW*

0.79

0.075

ChGCT

15-19

17

BulbW

0.86-1.09

0.99

PLSAPW/L

0.47

0.055

LL

0.78-1.29

1.02

EmbmidW

0.18-0.21

0.19

BulbW/EmbL* 0.77

0.022

LW

1.06-1.43

1.28

PalpTibL

2.71-3.09

2.82

EmbmidW/L

0.16

0.006

CUSP*

112-169

145

PalpTibW

1.26-1.52

1.40

PalpTibW/L*

0.50

0.003

SL

3.11-4.15

3.64

PalpTibS

1-2

2

BulbL/TibL*

0.77

0.053

SW

2.43-3.41

2.95

PalpPatS

1-3

2

PLSAPW

0.29-0.35

0.34

PalpFemS

0-3

1

PLSAPL

0.64-0.82

0.73

FelS

2-5

4

PalS*

0

0

Other material. AMS KS112636, Vic. Dept. Agric., Bairnsdale (locality
unknown), 20 June 1978.

Diagnosis. CL 4.72-6.60 (male). Small atracine spiders.
Differ from other atracine species by having a short, broad
embolus with distal part set at distinct angle to shaft (Fig.
65B,C).

Male (holotype) — Size. Carapace length 5.74, width 5.70.
Abdomen length 6.5, width 4.7. — Colour. Basic colour

Figure 64. Hadronyche jensenae, male: (A) cephalothorax and
chelicerae, dorsal; ( B ) cephalothorax and chelicerae, lateral; (C)
cheliceral groove teeth; ( D ) sternum, labium and maxilla; (E) leg
II, prolateral; (F) spinnerets; (G) palp, prolateral. Scale lines 1 mm.

pattern; abdomen partly discoloured or damaged in all
specimens. — Carapace. About as wide as long, cephalic area
moderately raised. Height 2.65; frontal width 4.21. Fovea
procurved. Mid-dorsal cephalic setae few, do not reach fovea.
Anterior strial setae absent. Anterolateral carapace angle with
weak hairs only. — Eyes. Central eye region slightly raised.
Eye group width 1.57. Median ocular quadrangle length
0.60, anterior width 0.74, posterior width 1.10. Diameters:
AME 0.20; ALE 0.35; PLE 0.29; PME 0.23. —Chelicerae.
Groove margins diverge weakly distally. Groove length 2.00,
middle width 0.30. Cheliceral teeth. 15 central, occupying
full length of groove; 9 prolateral; 12 retrolateral. — Labium.
Much wider than long, rectangular, apically weakly indented.
Length 0.92, width 1.25. Labiosternal sigilla broad. Cuspules
c. 160, number moderate to low. — Sternum. Ovoid. Length
3.59, width 2.98. Posterior sternal sigilla ovoid to elongate;
middle sigilla small. — Palp. Tegular area slightly wider than
long. Embolus very short and broad, base strongly offset
from tegulum. Distal part of embolus not twisted, but angled
strongly at about 45° to shaft. Bulb length 2.23, width 1.01.
Embolus length 1.31, midwidth 0.20. Tibia short, strongly
incrassate proximally. Length of femur 2.71, patella 1.12,
tibia 2.71. Width of tibia 2.01. Spination: femur 6; patella 1;
tibia d2. Distal femur with spines, without sinuous bristles.
— Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.34

2.52

3.94

3.46

2.38

17.64

2

4.79

2.30

3.74

3.19

2.38

16.40

3

4.08

2.14

2.89

3.03

2.24

14.38

4

5.17

2.28

4.08

3.90

2.82

18.25

Tibia I width 0.94. Femora I, II with dorsal spines.
Metatarsus I slightly thickened in proximal half, spines
strong, scattered. Tibia II without apophysis, ventral spines
few, grouped proximally with c. two larger spines toward
middle; distoventral tibia weakly concave. Metatarsus II
weakly to moderately sinuous with small, apophysis mid-
ventrally. Tarsi III, IV slightly swollen. Anterior coxal hairs
unmodified. Tarsal and distal metatarsal scopulae: legs I,
II weak; legs III, IV moderate. Tarsal claw teeth legs I, II:
superior 11, 11; inferior 0, 0. Trichobothria legs I, II: tarsus
8,7; metatarsus 6,6; tibia p5 r5, p5 r5. — Leg spination. Leg
I: femur d5, patella 0, tibia 18, metatarsus 19, tarsus 10. Leg

Gray: Australian funnel-web spiders

353

Figure 65. Hadronyche jensenae, male ( A,D-K , AMS KS112636; B,C, 11086 [MV]): (A) palp, prolateral; (B) bulb; (C) embolus, distal;
(D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; ( H,I) distal patella and tibia II— H,
ventral, /, prolateral; ( J,K) metatarsus II— J, ventral, K, prolateral. Scale lines: 0.2 mm, except B 0.5 mm and D,F,G 0.3 mm.

354

Records of the Australian Museum (2010) Vol. 62

II: femur d4, patella 0, tibia 5, metatarsus 11, tarsus 9. Leg
III: femur dl, patella pdlO, tibia 10 (r2), metatarsus 17 (p5
r2), tarsus 8. Leg IV: femur dl-2 (strong bristles present),
patella 0, tibia 3 (rdl), metatarsus 10, tarsus 11. — Abdomen.
Bristles stronger anteriorly with a prominent frontal pair.
Posterior lateral spinnerets with rather short apical segment.
Lengths: total 1.90; basal segment 0.84; middle 0.42; apical
0.64. Apical segment width 0.34.

Distribution. Central Gippsland, Victoria (Fig. 63).

Hadronyche pulvinator (Hickman)

Figures 66, 67

Atrax pulvinator Hickman, 1927: 70. Main, 1985: 41.
Hadronyche pulvinator. -Gray, 1988: 114 (Transferred from
Atrax).

Types. Holotype female: QVM 1957-15-20 Type 17. The Cascades, Hobart,
Tasmania, 42°54'S 147°17’E, V.V. Hickman.

Diagnosis. CL 6.15 (female). Small atracine spiders. Differs
from H. venenata having central cheliceral teeth in a short,
basal row (Fig. 66D), and relatively short labium and
spinnerets (Fig. 66A,G).

Redescription of holotype female. Size. Carapace length
6.15, width 5.64. Abdomen length 10.20, width 8.50. —
Colour. Basic colour pattern. — Carapace. Slightly longer
than wide, strongly raised. Height 2.71. Frontal width 4.94.
Cephalic length 4.69. Fovea procurved, anterior margin
slightly indented. Mid-dorsal cephalic setae do not reach
fovea. Anterior strial setae absent. Anterolateral carapace
angle with a few very weak bristles. — Eyes. Central eye
region very weakly raised. Eye group width 1.72. Diameters:

Figure 66. Hadronyche pulvinator, female: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) leg III,
prolateral, patella, tibia, metatarsus; (D) cheliceral groove teeth;
(E) spermathecae; (F) cephalothorax and chelicerae, dorsal; (G)
spinnerets. Scale lines 1 mm.

' 250 km'

Figure 67. Collection records for Hadronyche pulvinator.

AME 0.20, ALE 0.34, PLE 0.25, PME 0.22. Interdistances:
AME-AME 0.30, AME-ALE 0.17, ALE-PLE 0.20, PLE-
PME 0.07, PME-PME 0.82. Median ocular quadrangle
length 0.50, anterior width 0.70, posterior width 1.25.
— Chelicerae. Paturon with a medial line of long, weak
bristles, remaining anterodorsal bristles shorter. Groove
narrow, margins diverge slightly distally. Groove length
1.91, middle width 0.50. Distal half of groove transversely
ridged. Cheliceral teeth: 7 central, in basal third of groove; 6
prolateral; 6 retrolateral, absent from distal third of margin.
— Labium. Wider than long. Length 1.22, width 1.48.
Anterior margin indented apically, lateral margins rounded.
Cuspules grouped anterocentrally. Labiosternal sigilla
narrowed centrally. — Sternum. Broad, ventrally domed.
Length 3.85, width 3.44. Posterior sigilla ovoid: length 0.76,
width 0.44. — Palp. Spination: tarsus 3. Trichobothria: tibia
p4 r5, tarsus 4. Tarsal claw with 2-3 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

3.94

1.89

2.50

1.97

1.60

11.90

2

3.36

1.89

2.13

1.79

1.46

10.63

3

3.44

1.64

1.89

1.64

1.48

10.09

4

4.14

2.26

2.46

2.50

1.84

13.20

Tibia I width 1.10. Metatarsus I short, proximal width 0.83.
Coxal hairs unmodified. Tibia I, II without spines. Tarsal
claw teeth legs I, II: superior 5,5; inferior 2,1. Trichobothria
legs I, II: tarsus 7, 6; metatarsus 7, 7; tibia p4 r5, p5 r4. —
Leg spination. Leg I: femur 0, patella 0, tibia 0 (2 bristles
distally), metatarsus 3; tarsus 2. Leg II: femur 0, patella
0, tibia 0, metatarsus 3, tarsus 2. Leg III: femur 0, patella
pd9, tibia 6 (p3 rl), metatarsus 12 (p6 r2), tarsus 5. Leg IV:
femur 0, patella pi, tibia 2 (pi), metatarsus pv8, tarsus 6 —
Abdomen. Sparse cover of fine hairs and bristles. Posterior
lateral spinnerets with short apical segment. Lengths: total
1.87; basal segment 0.91, middle 0.36, apical 0.60. Apical
segment width 0.34. — Genitalia. Spermathecae short,
weakly bulbous apically. Length 0.68, width 0.41.

Distribution. Known only from the Cascades area, Hobart,
Tasmania (Fig. 67).

Comments. This species is known only from the holotype
female, collected more than 80 years ago. Sporadic collecting
efforts at the type locality have failed to locate additional
specimens, leading to real concern about the status of the
species. Collecting activities have mostly been restricted to

Gray: Australian funnel-web spiders

355

visual (daytime) searching in bushland areas in the Cascades
region, mainly targeting logs and rocks, often near creek
banks. A regular program of short-term pitfall trapping
and ground/litter slicing may be more successful in finding
specimens of this elusive species. These techniques were
use to good effect in trapping males and locating the in-litter
burrows of females of H. flindersi in South Australia

infensa species group

Description (Figs. 87-89, 91-93). Medium to large sized
funnel web spiders. Male femora I, II without spines.
Labium relatively long (LL/LW 0.85-0.96) (cf. short labium
in lamingtonensis group species). Central cheliceral tooth
row long (uniquely short, basal in H. kaputarensis ). Tibia II
without apophysis—either more or less sinuous (sometimes
slightly thickened proximally) with ventral spines clustered in
proximal half; or more cylindrical with clustered to scattered
spines. Metatarsus II without apophysis, cylindrical to weakly
sinuous, somet im es slightly swollen mid-ventrally. Male palp
with few spines (tibia 0-3, patella 0-1, femur 0-2). Embolus
with shaft broad or narrow, moderately curved and weakly to
not tapered. Distal part of embolus weakly to strongly twisted
(Figs. 77B, 81B)—twisting often extended back along shaft
as a deep, longitudinal fold, well developed in species with
wider, weakly tapered emboli (Fig. 86B,C). Ratio of bulb
length to palpal tibia length 0.78-0.95 (i.e., usually greater
than in cerberea group species). Posterior lateral spinnerets
with moderately long apical segment (PLSAPW/L 0.32-0.37).

maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cheliceral
groove teeth; (D) palp, prolateral; (E) cephalothorax and chelicerae,
dorsal; ( F ) leg II, prolateral; (G) spinnerets. Scale lines 1 mm.

Included species: H. infensa (Hickman), H. valida
(Rainbow & Pulleine), H. orana n.sp., H. lynabrae n.sp., H.
macquariensis n.sp., H. levittgreggae n.sp., H. kaputarensis
n.sp., H. walkeri n.sp.

Distribution. Coast and highlands from Hawkesbury River
region of mid-eastern New South Wales to southeastern
Queensland.

Comments. These species are distributed across a very
dissected coastal and highland landscape and it is likely that
many more species await recognition—for example males
from the Barrington Tops and Mount Banda Banda regions
probably represent new species.

Hadronyche infensa (Hickman)

Figures 68-71; Tables 20, 34

Atrax infensus Hickman, 1964: 108. Main, 1985: 41.
Hadronyche infensa- Gray, 1988: 114 (transferred from
Atrax).

Types. Holotype male: AMS KS953 (old catalogue number K68385).
Toowoomba, Queensland, 27°33'S 151°57'E, P. Walker, 26 January 1963.
Allotype female: AMS KS954 (old catalogue number K68386). Data as
for holotype.

Figure 69. Hadronyche infensa, female: (A) spinnerets; ( B )
cephalothorax and chelicerae, lateral; (C and D ) spermathecae
variation; ( E ) cephalothorax and chelicerae, dorsal; (E) cheliceral
groove teeth; (G) leg III, prolateral, patella, tibia, metatarsus; (H)
sternum, labium and maxilla. Scale lines 1 mm,

356

Records of the Australian Museum (2010) Vol. 62

Figure 70. Hadronyche infensa, male (AMS KS4716, RHS except B,C LHS): (A) palp, prolateral; (B) bulb; (C) embolus, distal; (D,E)
distal patella and tibia I— D, ventral, E, prolateral; (EG) metatarsus I— F, ventral, G, prolateral; ( H,I) distal patella and tibia II— H, ventral,
/, prolateral; (J,K) metatarsus II— J, ventral, K, prolateral. Scale lines: 0.3 mm, except B 1.0 mm and G,J 0.2 mm.

Gray: Australian funnel-web spiders

357

Table 20. Male morphological data— Hadronyche infensa (n = 13).

character

range

mean

character

range

mean

character

ratio

SD

CL

9.04-11.48

10.35

MtlS

25-52

37

CW/CL

0.93

0.038

CW

8.70-10.58

9.58

TalS

24-51

36

CH/CW

0.42

0.031

CH

3.43^.84

4.02

Ti2S

11-29 (p0-l) 17

CFW/CL

0.66

0.022

CFW

5.95-7.41

6.8

STC2teeth

11-13

12

CHGW/L

0.18

0.015

ChGL

2.58-3.38

2.73

BulbL

3.82-4.86

4.37

SW/SL

0.75

0.034

ChGW

0.46-0.60

0.52

EmbL

2.82-3.58

3.15

LL/LW

0.93

0.04

ChGCT

8-26

15

BulbW

1.22-1.64

1.47

PLSAPW/L

0.30

0.035

LL

1.48-1.94

1.71

EmbmidW

0.35-0.49

0.41

BulbW/EmbL 0.47

0.049

LW

1.58-1.98

1.83

PalpTibL

4.41-5.78

5.00

EmbmidW/L* 0.13

0.013

CUSP

320-494

383

PalpTibW

1.91-2.34

2.11

PalpTibW/L

0.42

0.029

SL

4.93-6.89

5.74

PalpTibS

0-1

<1

BulbL/TibL*

0.88

0.044

SW

3.76-5.00

4.49

PalpPatS

0

—

PLSAPW

0.49-0.74

0.58

PalpFemS

0

—

PLSAPL

1.69-2.24

1.78

FelS

0

PalS

0-5 (p0-l)

3

Other material examined. Queensland (males): QMB S672, Bellthorpe
West, 5 December 1979, G. Hodges; QMB S392, Royce Reserve, near
Toowoomba, 24 October 1975-30 April 1976, G.B. & S.R. Monteith; AMS
KS8790, Buderim, January 1982; QMB S147, Casey Creek via Imbil,
31 December 1974-27 March 1975, G.B. & S.R. Monteith; QMB S173,
Cooran Plateau via Traveston, 31 December 1974-27 March 1975, G.B.
& S.R. Monteith; QMB S396, Elginvale, 12 December 1976-26 March
1977, G.B. & S.R. Monteith; QMB S158, Flaxon, 30 November 1973,
J.J. Roth; QMB S168, Gallangowan, 10 November 1974-29 December
1974, G.B. & S.R. Monteith; QMB S376, Mount Glorious, 7 March 1978,
T. Hillier; QMB SI65, Kingaroy, 23 November 1977, L. Briskey; QMB
S673, Maleny, 4.X.1979,1. Joyce; QMB S152, Mount Nebo, 26 October
1972, D. Dale; QMB S171, Maleny, 23 November 1973, M. Erskine-Wyse;
QMB S153, Mapleton, 2 December 1975, P. Allsop; QMB S395, Mistake
Mountains, 10 October 1976-9 January 1977, G.B. & S.R. Monteith; QMB
S160, Montville, 7 November 1977, J. Channon; QMB S670, Montville, 23
November 1979, D. Roberts; QMB S674, Nambour, 17 November 1979, R.
Cook; QMB S379, Nambour, 16 November 1978, S. Martin; QMB S671,
Mount Nebo, 22 December 1979, D. Webster; QMB S188, Mount Nebo, 28
April 1979, A. Rozetelds; QMB S170, Mount Tennisonwoods, near Mount
Glorious, 12 November 1975-27 January 1976, G.B. & S.R. Monteith;
QMB S151, Toowoomba, 15 January 1952, T. Passlow; QMB S167, Upper
Yarraman State Forest via Maidenwell, 17 July 1974—10 November 1974,

G. B. &S.R. Monteith; AMS KS4716, Toowoomba, 27°36'S 151°57'E, 12Jan
1980, P. Walker. Queensland (females): AMS KS13423, Cunninghams Gap;
QMB S376, Mount Glorious, 7 March 1978, T. Hillier; QMB S374, Maleny;
AMS KS990, Toowoomba, 20v.l972, J. Cann; AMS KS4596, Toowoomba,
February 1979, P. Walker. New South Wales (males): Tenterfield, 29°03'S
152°01F, (AMS); Tooloom, 28°37'S 152°25’E, (AMS).

Diagnosis. CL 9.04-11.48 (male). Differs from//, walkeri
and H. macquariensis by embolus base strongly offset from
tegulum (Fig. 70B) and greater length of bulb (BulbL/Tib
L 0.88) (Figs. 68D, 70A); differs from H. levittgreggae,

H. valida, H. orana and H. kaputarensis by more slender
embolus shaft (EmbmidW/L 0.13) and from H. lynabrae
by less pronounced distal embolic twisting (Fig. 70B,C).

Redescription of holotype male. Size. Carapace length
10.78, width 9.38. Abdomen length 9.86, width 7.48. —
Colour. Basic colour pattern. Abdomen often with a small
anterodorsal pale patch flanked by small sigilla. — Carapace.
Longer than wide, cephalic area moderately raised. Height
4.18; frontal width 7.15. Fovea narrow, procurved. Mid¬
dorsal cephalic setae reach fovea. Anterior strial setae
present. Anterolateral angle of carapace with numerous
bristles. — Eyes. Central eye region weakly raised. Eye
group width 2.31. Median ocular quadrangle length 0.88,

anterior width 0.97, posterior width 1.44. Diameters: AME
0.31, ALE 0.49, PLE 0.42, PME 0.24. —Chelicerae. Dorsal
paturon with oblique ridges proximally. Cheliceral groove
with margins diverging distally. Groove length 3.03, middle
width 0.52. Cheliceral teeth: 22 central, occupying full length
of groove; 14 prolateral; 10 retrolateral. — Labium. Wider
than long, weakly indented apically. Length 1.67, width 1.97.
Labiosternal sigilla broad, entire. Cuspules c. 379, number
high. — Sternum. Ovoid. Length 6.15, width 4.83. Posterior
sigilla long, ovoid. — Palp. Tegular area wider than long.
Embolus moderately long and wide, base strongly offset from
tegulum. Embolus moderately long and broad, shaft curved
and hardly tapered; distal embolus moderately twisted with
twisting extended back along shaft as a rolled margin. Bulb
length 4.42, width 1.43. Embolus length 3.26, midwidth 0.38.
Length of femur 4.62, patella 2.38, tibia 4.90. Width of tibia
1.94. Spines absent. Tibia with several strong ventral bristles.
Several sinuous bristles on distal femur. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.87

4.39

6.66

5.78

3.60

29.30

2

8.08

3.91

6.46

6.10

3.63

28.08

3

7.04

3.57

5.17

5.81

3.52

25.11

4

8.33

4.15

6.50

6.57

4.28

29.83

Tibia I width 1.58. Femora I and II without spines. Metatarsus

I slightly thickened proximally with larger ventral spines
grouped in proximal half. Tibia II without apophysis, most
ventral spines loosely grouped proximocentrally. Metatarsus

II weakly sinuous proximally, otherwise unmodified. Coxae
I and II with anteromedial setae weakly thickened, sparse.
Scopulae: tarsi (legs I-IV), moderate to strong; distal
metatarsi, absent to weak (legs I, II), moderate (legs III,
IV). Tarsal claw teeth legs I, II; superior 14,12; inferior 4, 3.
Trichobothrialegs I, II: tarsus 13,12: metatarsus 17,13; tibia
p6 r7, p7 r8. — Leg spination. Leg I; femur 0, patella 0, tibia
33, metatarsus 38, tarsus 27. Leg II: femur 0, patella p2, tibia
19 (pi) metatarsus 24, tarsus 35. Leg III: femur 0, patella p5,
tibia 12 (p2 r2), metatarsus 31, tarsus 33. Leg IV: femur 0,
patella 0, tibia 10 (rl), metatarsus 22, tarsus 30. — Abdomen.
Posterior lateral spinnerets with apical segment moderately
long. Lengths: total 4.48; basal segment 1.70; middle 1.05;
apical 1.73. Apical segment width 0.56.

358

Records of the Australian Museum (2010) Vol. 62

Figure 71. Collection records for Hadronyche infensa (squares)
and H. orana (circles).

Redescription of allotype female. Size. Carapace length
11.63, width 10.27. Abdomen length 15.98, width 10.88.
— Colour. Basic colour pattern. — Carapace. Longer than
wide, strongly raised. Height 4.76; frontal width 8.77.
Cephalic length 8.76. Fovea procurved, narrow. Mid-dorsal
cephalic setae numerous, reach back to fovea. Anterior
strial setae very few basally. Anterolateral carapace angle
with several bristles. — Eyes. Eye region sessile or very
slightly raised centrally. Eye group width 2.52. Diameters:
AME 0.29, ALE 0.58, PLE 0.41, PME 0.26. Interdistances:
AME-AME 0.46, AME-ALE 0.31, ALE-PLE 0.29, PLE-
PME 0.10, PME-PME 1.22. Median ocular quadrangle
length 0.97, anterior width 1.09, posterior width 1.70. —
Chelicerae. Cheliceral groove moderately wide, margins
diverging distally. Groove length 3.40, middle width 0.82.
Cheliceral teeth: 67 central, in several rows, occupying full
length of groove; 15 prolateral; 12 retrolateral. — Labium.
Slightly wider than long. Length 2.31, width 2.52. Anterior
margin weakly indented. Cuspules spread in broad v-shape
over anterior three quarters of labium. — Sternum. Ovoid.
Length 5.44, width 4.22. Posterior sigilla moderately small,
elongate: length 0.68, width 0.27. — Palp. Spination: tibia
4, tarsus 8. Trichobothria: tibia p8 r7, tarsus 7. Tarsal claw
with 7 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.30

4.69

6.12

4.86

3.06

27.03

2

7.45

4.52

5.44

4.59

2.89

23.89

3

6.43

3.64

4.49

4.59

3.06

22.21

4

8.23

4.42

5.98

5.66

3.40

27.69

Tibia I width 2.07. Metatarsus I proximal width 1.43. Coxae
I, II with basally thickened setae anteriorly. Tarsal claw teeth
legs I, II: superior 10, 10; inferior 4, 4. Trichobothria legs

I, II: tarsus 12, 12; metatarsus 16, 14; tibia p8 r7, p7 r7. —
Leg spination. Leg I: femur 0, patella 0, tibia 3, metatarsus

II, tarsus 7. Leg II: femur 0, patella pi, tibia 2, metatarsus
12, tarsus 9. Leg III: femur 0, patella pd8, tibia 6 (p2 rl),
metatarsus 18 (p6 rd2), tarsus 12. Leg IV: femur 0, patella 0,

tibia 4 (rl), metatarsus 15, tarsus 13. — Abdomen. Posterior
lateral spinnerets with slender apical segment of moderate
length. Lengths: total 4.28; basal segment 1.70, middle
0.88, apical 1.60. Apical segment width 0.65. — Genitalia.
Spermathecae digitiform, usually elongate. Length 1.41,
width 0.44.

Distribution. Southeastern Queensland to upper north¬
eastern new South Wales (Fig. 71).

Hadronyche orana n.sp.

Figures 71-74; Tables 21, 34

Etymology. The specific epithet refers to the Orana region
of New South Wales, part of the distribution of this species.

Types. Holotype male: AMS KS7811, Mt. Deba near Coolah, New South
Wales, 31°52'S 149°39’E, 1 June 1981, P. Esdaile. Paratypes (AMS). New
South Wales. Males-. AMS KS5288, Hargraves, 32°44’S 149°27'E, 11 May
1980, N. Mattick; AMS KS967, Mandurama, 33°38’S 149°04’E, 20 June
1976, P. Bremner; AMS KS968, Molong, 33°05’S 148°51'E, 12 July 1969,
R.H. Keeling; AMS KS5203, Mt. Deba, nr. Coolah, 31°52’S 149°39’E, 8
May 1980, S. Esdaile; AMS KS10974, “Stonehenge” on Cassilis Road,
Mudgee area, 3 May 1983, E.H. Elward; AMS KS7563, Warrumbungle
Mtns., 31°19’S 149°00’E, 10 November 1981, C. Martin. Females : AMS
KS1071, “Hill Top” near Hargraves, 32°44’S 149°27'E, 30 June 1978, N.
Mattick; AMS KS4053, Binnaway, 31°33’S 149°23’E, September 1979,
R. England; AMS KS4476, Dunedoo area, 32°00'S 149°23'E, 15 January
1980; AMS KS3429, Hargraves, 32°44'S 149°27'E, 18 May 1979; AMS
KS8593, Mudgee, 32°34'S 149°35’E, 4 January 1982, P Schiemer; AMS
KS977, Warrumbungle Mtns, 231°19'S 149°00'E, 3 April 1967; AMS
KS5980, Munghom Gap Nature Reserve, nr. Mudgee, 32°24’S 149°47’E,

21 August 1980, M. Gray & C. Horseman.

Other material examined. New South Wales (males): AMS KS8793,
Borenore, nr. Orange, December 1981, G. Scarvell; AMS KS7691, Clergate,
15 May 1981; AMS KS964, Hargraves, 10 October 1977, J. Sibley; AMS
KS2963, Hargraves, May 1979, Gillman; AMS KS965, Hargraves area,

22 October 1977, L.M. Mattick; AMS KS960, Mudgee, December 1977,
M.P.P.B.; AMS KS961, Mudgee, 23 November 1973, M.P.P.B.; AMS
KS 10975, Mudgee, 5 May 1983; AMS KS976, “Tara”, Warrumbungle
Mountains, 28 November 1977, E. Edmonson; AMS KS966, Orange district,
AMS KS 14262, Siding Springs in Warrumbungle Mountains, Apr 1984, D.
Rowell. New South Wales (females): AMS KS8746, Hargraves, January
1982; AMS KS978, Warrumbungle Mountains; AMS KS6219, Munghorn
Gap Nature Reserve, nr. Mudgee, 21 August 1980, M. Gray & C. Horseman.

Diagnosis. CL 7.76-9.72 (male). Differs from H. levitt-
greggae and H. valida by having few tibiae II ventral
spines (3-7) (Fig. 74D,H); from//, infensa, H. walkeri, H.
macquariensis, H. lynabrae by having a slender, weakly
twisted embolic shaft (EmbmidW/L 0.06) (Fig. 74B,C);
and from H. kaputarensis by having a long central cheliceral
tooth row.

Male (holotype) — Size. Carapace length 7.76, width 7.41.
Abdomen length 8.04, width 6.57. — Colour. Basic colour
pattern. — Carapace. Slightly longer than wide, cephalic
area moderately raised. Height 3.57. Frontal width 5.93.
Fovea procurved, anterior margin indented. Mid-dorsal
cephalic setae almost reach fovea. Anterior strial setae absent.
Anterolateral carapace angle with a few bristles of moderate
size. — Eyes. Central eye region slightly raised. Eye group
width 2.16. Median ocular quadrangle length 0.78, anterior
width 1.02, posterior width 1.48. Diameters; AME 0.30,
ALE 0.51, PLE 0.36, PME 0.23. — Chelicerae. Cheliceral
groove of medium width, margins diverging distally.

Gray: Australian funnel-web spiders

359

Table 21. Male morphological data —Hadronyche orana (n = 11).

character

range

mean

character

range

mean

character

ratio

SD

CL

7.76-9.72

8.77

MtlS*

10-16

13

CW/CL

0.93

0.042

CW

7.41-8.87

8.14

TalS*

4-10

6

CH/CW

0.48

0.03

CH

3.32-4.48

3.88

Ti2S*

3-7

5

CFW/CL

0.70

0.031

CFW

5.59-6.87

6.13

STC2teeth

10-15

11

CHGW/L

0.21

0.036

ChGL

2.19-2.65

2.35

BulbL

3.21-3.77

3.48

SW/SL

0.81

0.04

ChGW

0.43-0.63

0.50

EmbL

2.19-2.63

2.39

LL/LW

0.90

0.065

ChGCT

11-26

19

BulbW

1.14-1.36

1.23

PLSAPW/L

0.35

0.025

LL

1.29-1.76

1.54

EmbmidW

0.14-0.19

0.16

BulbW/EmbL 0.52

0.02

LW

1.60-1.90

1.71

PalpTibL

3.74_4.42

4.06

EmbmidW/L* 0.06

0.009

CUSP

246-390

309

PalpTibW

1.59-1.90

1.76

PalpTibW/L

0.43

0.02

SL

4.83-5.74

5.28

PalpTibS

0-1

<1

BulbL/TibL

0.86

0.035

SW

3.96-4.86

4.28

PalpPatS

0

—

PLSAPW

0.33-0.68

0.55

PalpFemS

0

—

PLSAPL

1.44-1.86

1.63

FelS

0

—

PalS

0-lp

(«1)

Groove length 2.19, middle width 0.54. Cheliceral teeth:
18 central, occupying full length of groove; 8 prolateral;
9 retrolateral. — Labium. Wider than long, with long hairs
laterally. Weakly indented apically. Length 1.32, width
1.60. Shallowly grooved behind cuspules. Labiosternal
sigilla entire. Cuspules c. 309, number moderate to high.
— Sternum. Ovoid. Length 5.00, width 4.02. Posterior sigilla
ovoid, elongate. — Palp. Tegular area a little wider than long.
Embolus shaft long, slender, weakly tapering distally, gently

Figure 72. Hadronyche orana, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cheliceral
groove teeth; (D) spinnerets; (E) cephalothorax and jaw, dorsal; (F)
leg II, prolateral; (G) palp, prolateral. Scale lines 1 mm.

curved. Distal embolus slender, only slightly twisted, flange
elongate. Basal embolus strongly offset from tegulum. Bulb
length 3.57, width 1.24. Embolus length 2.55, midwidth
0.17. Length of femur 4.04, patella 1.93, tibia 4.22. Width
of tibia 1.77. Spination: spines absent; weak sinuous bristles
on distal femur. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.01

3.35

5.19

4.98

3.13

23.66

2

6.44

3.20

4.90

5.00

3.24

22.58

3

5.63

2.75

4.02

5.16

3.62

21.88

4

7.22

3.18

5.36

6.55

4.22

26.53

and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) leg
III, prolateral, patella, tibia, metatarsus; ( D ) cephalothorax and
chelicerae, dorsal; ( E ) spermathecae; (F) cheliceral groove teeth;
(G) spinnerets. Scale lines 1 mm.

360

Records of the Australian Museum (2010) Vol. 62

Figure 74. Hadronyche orana, male ( A,D-K , AMS KS960; B,C, AMS KS5203): (A) palp, prolateral; (B) bulb; (C) embolus, distal; (D,E)
distal patella and tibia I— D, ventral, E, prolateral; (EG) metatarsus I— F, ventral, G, prolateral; (//,/) distal patella and tibia II— H, ventral,
I, prolateral; ( J,K) metatarsus II—/, ventral, K, prolateral. Scale lines: 0.2 mm, except A,B 1.0 mm and D,E,EI,1 0.3 mm.

Gray: Australian funnel-web spiders

361

Tibia I width 1.48. Femora I and II lacking dorsal spines.
Metatarsus I slightly thickened proximally. Tibia II and
metatarsus II unmodified. Tibia I and II with few ventral
spines, scattered. Tarsus I, II with few spines. Anterior coxal
hairs normal. Tarsal scopulae: legs I, II weak; legs III, IV
moderate to strongly developed. Distal metatarsal scopulae:
legs I, II absent; legs III, IV moderate. Tarsal claw teeth legs

I, II: superior 10, 10; inferior 4, 4. Trichobothria legs I, II:
tarsus 13, 11; metatarsus 15, 14; tibia p7 r7, p7 r7. — Leg
spination. Leg I: femur 0, patella 0, tibia 6, metatarsus 13,
tarsus 6. Leg II: femur 0, patella pi, tibia 3, metatarsus

II, tarsus 9. Leg III: femur 0, patella p3, tibia 10 (pi r2),
metatarsus 23, tarsus 15. Leg IV: femur 0, patella 0, tibia 4
(r2), metatarsus 19, tarsus 21. — Abdomen. Posterior lateral
spinnerets with apical segment of moderate length. Lengths:
total 4.65; basal segment 1.68, middle 1.35, apical 1.62.
Apical segment width 0.59.

Female (paratype AMS KS2071) — Size. Carapace length
8.57, width 8.67. Abdomen length 14.21, width 11.22.
— Colour. Basic colour pattern. — Carapace. About as
long as wide, cephalic area strongly raised. Height 3.85;
frontal width 7.99. Cephalic length 7.11. Fovea procurved.
Mid-dorsal cephalic setae numerous, extend back to fovea.
Anterior strial setae numerous. Strial, cephalic and marginal
setae numerous. Anterolateral carapace angle with several
weak bristles. — Eyes. Central eye region slightly raised.
Eye group width 2.52. Diameters: AME 0.31, ALE 0.60,
PLE 0.41, PME 0.31. Interdistances: AME-AME 0.44,
AME-ALE 0.31, ALE-PLE 0.28, PLE-PME 0.16, PME-
PME 1.09. Median ocular quadrangle length 0.80, anterior
width 1.02, posterior width 1.67. — Chelicerae. Cheliceral
groove margins diverge distally. Groove length 3.06, middle
width 0.75. Cheliceral teeth: 64 central occupying full
length of groove in 2-3 irregular rows; 11 prolateral; 10
retrolateral. — Labium. As long as wide, anterior margin
not indented. Length 2.10, width 2.12. Cuspules occupying
anterior half of labium. Labiosternal sigilla entire, narrowed
centrally. — Sternum. Ovoid, strongly convex. Length
6.24, width 4.82. A few strong bristle-like setae grouped
at anterolateral corners. Posterior sternal sigilla elongate:
length 1.02, width 0.44. — Palp. Spination: tibia 5, tarsus
11. Trichobothria: tibia p7 r7, tarsus 14. Tarsal claws with
7 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.75

3.77

5.30

4.39

2.58

22.79

2

6.19

3.40

4.76

4.08

2.65

21.08

3

5.44

3.13

3.74

4.01

2.82

19.14

4

6.90

3.67

5.20

4.92

2.92

23.61

Tibia I width 1.60. Metatarsus I proximal width 1.05. Coxae
I, II with thorn-like setae frontally. Tarsal claw teeth legs I,
II: superior 10-11, 10; inferior 5, 4. Trichobothria legs I, II:
tarsus 15, 14; metatarsus 15, 14; tibia p7 r7, p8 r8. — Leg
spination. Leg I: femur 0, patella 0, tibia 4, metatarsus 12,
tarsus 9. Leg II: femur 0, patella pi, tibia 4 (pi), metatarsus
13, tarsus 10. Leg III; femur 0, patella p6 rl, tibia 12 (p5 r2),
metatarsus 23 (p3 r3), tarsus 14. Leg IV: femur 0, patella 0,
tibia 5 (rl), metatarsus 18, tarsus 24. — Abdomen. Posterior
lateral spinnerets with short apical segment. Lengths; total
3.74; basal segment 1.60, middle 0.95, apical 1.36. Apical
segment width 0.82. — Genitalia. Spermathecae short, apical
third narrower than basal two thirds. Length 1.02, width 0.48.

Distribution. Western Great Dividing Range from the
Warrumbungles Range south to the Mudgee region, New
South Wales (Fig. 71).

Hadronyche valida (Rainbow & Pulleine)

Figures 75-78; Tables 22, 34

Atrax valida Rainbow & Pulleine, 1918: 165.

Anepsiada ventricosa Rainbow & Pulleine, 1918: 167.
Gray, 1984: 441 (inferred synonymy as juvenile of Atrax
validus). First synonymized after Gray in Main, 1985: 42.
Atrax validus- Main, 1985: 41

Hadronyche valida.-G ray, 1988: 114 (Transferred from
Atrax).

Types. Holotype female: AMS KS955 (old catalogue number K40961).
Tamborine Mountain, Queensland, 27°55'S 153°11'E, 5 October 1912.
Paratype females: AMS KS956-9, data as above.

Other material examined. Queensland (males): QMB S163, Binna Burra,
Lamington Plateau, November 1973, R.J. Raven; QMB S157, Lamington
Plateau, 13 April 1974, R.J. Raven; QMB S390, Lower Albert River,
Lamington National Park, 1 November 1975-4 April 1976, G.B. & S.R.
Monteith; AMS KS13629, Mount Tamborine, 5 December 1983; QMB

5175, Eagle Heights, Mount Tamborine, 17 August 1973, G. Walker; QMB

5176, Springbrook, October 1976, J. Mainwaring; QMB S381, Tamborine
North, 26 March 1979, J. Aagaad; AMS KS988, Tyalgum, 7 January 1972,
H.G. Suttolk; AMS KS989, Tuntable Lalls, January 1975, McGovern;
QMB S164, Warrie National Park, Lamington Plateau, 15 November
1971, Balwin. New South Wales (males): AMS KS13539, Huonbrook,
5 December 1983, G. Watson; AMS KS987, Jiggi, November 1962, L.J.
Hartridge; QMB S183, Brindle Creek, nr. Wiangaree, 2 July 1975-15
November 1975, G.B. & S.R. Monteith; AMS KS13500, Coffee Camp,
nr. Lismore, 9 December 1983, C. McQueen; QMB S393, Mount Clunie,
nr. Woodenbong, 22 Lebruary 1976-8 May 1976, G.B. & S.R. Monteith;
QMB S391, Mount Warning, 16 November 1975-7 March 1976, G.B. &
S.R. Monteith; AMS KS4255, Lismore, 14 December 1979; AMS KS13407,
Nimbin, 23 November 1983, A. Lrame; AMS KS6824, Mullumbimby, 29
December 1980, W. Dawes; AMS KS2431, Murwillumbah, 11 January
1979, J. Morris; AMS KS22460, Murwillumbah, 28°20'S 153°24'E, 08
Dec 1989, A.W. Tucker.

Diagnosis. CL 8.73-10.61 (male). Differs from H. infensa,
H. walkeri, H. macquariensis and H. lynabrae by having
embolus moderately slender (EmbmidW/L 0.09), weakly
curved and twisted (Fig. 77A,B,C); differs from H. orana
and H. levittgreggae by more numerous spines on metatarsus
I (23-46) and tibia II (13-25) (Table 22); and from H.
kaputarensis by presence of long central cheliceral tooth
row (Fig. 75G).

Redescription of holotype female. Size. Carapace length
10.40, width 9.28. Abdomen length 13.60, width 9.86. —
Colour. Basic colour pattern. — Carapace. Longer than wide,
cephalic area strongly raised. Height 4.83, frontal width 8.33.
Cephalic length 7.82. Fovea strongly procurved. Mid-dorsal
cephalic setae numerous, reach back to fovea. Anterior strial
setae numerous. Strial and marginal setae numerous, weak.
Anterolateral carapace angle with many weak bristles. —
Eyes. Eye region slightly raised centrally. Eye group width
2.92. Diameters: AME 0.36, ALE 0.68, PLE 0.50, PME
0.31. Interdistances: AME-AME 0.42, AME-ALE 0.31,
ALE-PLE 0.34, PLE-PME 0.17, PME-PME 1.29. Median
ocular quadrangle length 0.95, anterior width 1.22, posterior
width 1.93. — Chelicerae. Cheliceral groove margins diverge
distally. Groove length 3.16, middle width 0.82. Cheliceral
teeth: 43 central, in irregular double row occupying full

362

Records of the Australian Museum (2010) Vol. 62

Table 22. Male morphological data —Hadronyche valida (n = 13).

character

range

mean

character

range

mean

character

ratio

SD

CL

8.73-10.61

9.60

MtlS

31-56

45

CW/CL

0.91

0.031

CW

7.92-9.57

8.71

TalS

23-46

34

CH/CW

0.41

0.025

CH

3.13-3.94

3.58

Ti2S

13-25 (pO-2) 21

CFW/CL

0.64

0.034

CFW

5.61-7.00

6.08

STC2teeth

10-12

11

CHGW/L

0.19

0.013

ChGL

2.42-2.98

2.64

BulbL

3.59^1.25

4.00

SW/SL

0.76

0.043

ChGW

0.43-0.56

0.50

EmbL

2.49-3.06

2.96

LL/LW

0.93

0.03

ChGCT

11-52

27

BulbW

1.16-1.43

1.26

PLSAPW/L

0.37

0.05

LL

1.36-1.72

1.56

EmbmidW

0.20-0.30

0.26

BulbW/EmbL 0.43

0.053

LW

1.53-1.84

1.67

PalpTibL

4.12-4.86

4.51

EmbmidW/L* 0.09

0.014

CUSP

265-393

325

PalpTibW

1.79-2.13

1.94

PalpTibW/L

0.43

0.029

SL

5.30-6.39

5.58

PalpTibS

0-3

1

BulbL/TibL

0.89

0.035

SW

4.16-4.59

4.24

PalpPatS

0

—

PLSAPW

0.43-0.72

0.59

PalpFemS

0

—

PLSAPL

1.36-1.87

1.61

FelS

0

—

PalS

0-2 (pi)

1

length of groove; 11 prolateral; 9 retrolateral. — Labium.
Wider than long, anterior margin not indented. Length
2.07, width 2.58. Cuspules occupying anterior two thirds
of labium. Labiosternal sigilla narrow, entire. — Sternum.
Ovoid. Length 6.94, width 5.32. Posterior sternal sigilla
elongate: length 1.50, width 0.54. — Palp. Spination: tibia 3
(bristle-like), tarsus 9. Trichobothria: tibia p8 r8, tarsus 13.
Tarsal claw with 7 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.89

4.76

5.98

4.93

2.85

26.41

2

6.94

4.28

5.61

4.56

2.79

24.18

3

6.53

3.67

4.42

4.59

3.06

22.27

4

7.89

4.66

6.19

5.78

3.50

28.02

Tibia I width 2.01. Metatarsus I proximal width 1.41. Coxae
I, II with basally thickened to thorn-like setae anteriorly.

Figure 75. Hadronyche valida, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cephalo-
thorax and chelicerae, dorsal; (D) leg II, prolateral; (E) spinnerets;
(F) palp, prolateral; (G) cheliceral groove teeth. Scale lines 1 mm.

Tarsal claw teeth legs I, II: superior 10, 9; inferior 4, 4.
Trichobothria legs I, II: tarsus 12,12; metatarsus 15,12; tibia
p8 r8, p8 r8. —Leg spination. Leg I: femur 0, patella 0, tibia

2, metatarsus 11, tarsus 10. Leg II: femur 0, patella 0, tibia

3, metatarsus 11, tarsus 7. Leg III: femur 0, patella pi, tibia
10 (p3 r3), metatarsus 19 (p2 rl), tarsus 14. Leg IV: femur 0,
patella 0, tibia 4 (r2), metatarsus 15, tarsus 20. — Abdomen.
Posterior lateral spinnerets with apical segment of moderate
length. Lengths: total 3.80; basal segment 1.46, middle

Figure 76. Hadronyche valida, female: (A) cheliceral groove teeth;
(B) cephalothorax and chelicerae, lateral; (C) sternum, labium and
maxilla; ( D ) spinnerets; (E) cephalothorax and chelicerae, dorsal;
( F) leg III, prolateral, patella, tibia, metatarsus; (G) spermathecae.
Scale lines 1 mm.

Gray: Australian funnel-web spiders

363

Figure 77. Hadronyche valida, male (. A,D-K , AMS KS22460; B,C, AMS KS6824): (A) palp, prolateral; (B) bulb; (C) embolus, distal;
( D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; ( H,I) distal patella and tibia II— H,
ventral, I, prolateral; ( J,K) metatarsus II—J, ventral, K, prolateral. Scale lines: 0.3 mm, except A,B 1.0 mm and/,/0.2 mm.

364

Records of the Australian Museum (2010) Vol. 62

0.90, apical 1.43. Apical segment width 0.68. — Genitalia.
Spermathecae digitiform, long and thin, straight except for
apical region which may be angled slightly. Apical quarter
of spermathecae slightly enlarged and set off from basal part
by a weak constriction. Length 1.71, width 0.48.

Male (S175 (QMB), Eagle Heights, Mt. Tamborine,
Queensland, 27°54'S 153°12'E, G. Walker, 17 August
1973). — Size. Carapace length 8.72, width 8.12. Abdomen
length 9.35, width 7.13. — Colour. Basic colour pattern.
— Carapace. Longer than wide, moderately raised. Height
3.61. Frontal width 5.70. Fovea procurved. Mid-dorsal
cephalic setae reach fovea. Anterior strial setae numerous.
Antero-lateral angle of carapace with numerous weak
bristles. — Eyes. Sessile to slightly raised. Eye group width
2.17. Median ocular quadrangle length 0.86, anterior width
0.96, posterior width 1.38. Diameters: AME 0.33, ALE 0.52,
PLE 0.37, PME 0.20. — Chelicerae. Dorsal paturon slightly
ridged proximally. Cheliceral groove narrow, margins
diverging distally. Groove length 2.42, middle width 0.50.
Cheliceral teeth: 14 central, running full length of groove;
9 prolateral; 12 retrolateral. — -Labium. Slightly wider than
long. Length 1.60, width 1.69. Labiosternal sigilla normal,
complete. Cuspules c. 336, number high. — Sternum. Ovoid.
Length 5.40, width 4.16. Posterior sigilla large, elongate.
— Palp. Tegular area wider than long. Embolus long and
moderately slender, weakly curved but shaft hardly tapered.
Distal embolus weakly to moderately twisted with twisting
extended back along shaft as a shallow longitudinal fold.
Basal embolus offset from tegulum. Bulb length 3.92, width
1.30. Embolus length 2.88, midwidth 0.27. Length of femur
4.19, patella 1.80, tibia 4.35. Width of tibia 1.80. Spination:
tibia with 2-4 bristle-like spines. Sinuous bristles on distal
femur weakly developed. — Legs. 4213. Legs I, II subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.05

3.69

5.66

5.54

3.20

25.14

2

6.81

3.69

5.90

5.58

3.20

25.38

3

6.35

3.36

4.92

5.54

3.61

23.78

4

7.91

3.85

6.47

6.85

4.22

29.30

Tibia I width 1.48. All femora lack dorsal spines. Metatarsus
I weakly thickened proximally, larger ventral spines grouped
proximally. Tibia II without apophysis but slightly thickened
proximally and weakly excavated distoventrally; spines
scattered, largest grouped proximally. Metatarsus II slightly
bent at base, no apophysis, spines scattered. Anterior coxal
hairs fine, unmodified. Tarsal scopulae weak on legs I, II,
strong on legs III, IV. Distal metatarsal scopulae absent to
weak legs I, II; moderately strong legs III, IV. Tarsal claw
teeth legs I, II: superior 11, 11; inferior 3, 3. Trichobothria
legs I, II: tarsus 15, 11; metatarsus 12, 13; tibia p7 r7, p7
r6. — Leg spination. Leg I: femur 0, patella pi, tibia 32 (pi),
metatarsus 56, tarsus 41. Leg II: femur 0, patella pO, tibia 23
(p2), metatarsus 41, tarsus 29. Leg III: femur 0, patella rl,
tibia 13 (p3 r2), metatarsus 31, tarsus 27. Leg IV: femur 0,
patella rl, tibia 11 (r4), metatarsus 20, tarsus 34 — Abdomen.
Posterior lateral spinnerets: apical segment of moderate
length. Lengths: total 3.91; basal segment 1.60; middle 0.75;
apical 1.56. Apical segment width 0.70.

Distribution. Eastern Border Ranges area of northern New
South Wales and southern Queensland (Fig. 78).

Figure 78. Collection records for Hadronyche valida (squares) and
H. lynabrae (circles).

Hadronyche lynabrae n.sp.

Figures 78-81; Tables 23, 34

Etymology. The species is named for Lyn Abra who sent
many funnel-web and other spiders to the Australian Museum
while working at the Australian Reptile Park.

Types. Holotype male: AMS KS8379, Wollombi near Maitland, New
South Wales, 32°40'S 151°04'E, November 1981. Paratypes (all AM).
New South Wales. Males : AMS KS13462, Dharug National Park, Mill
Creek, 33°21'S 15F05'E, September 1983, G.P. Clancy; AMS KS16449,
Dharug National Park, Mill Creek, August 1983, 33°21'S 151°05'E, G.P
Clancy; AMS KS5741, Morriset district, 33°06'S 151°30’E, 24 July 1980;
AMS KS1065, Stratford, 32°06’S 151°56'E, 26 June 1956, L. Davenport,
AMS KS1066, Newcastle, 32°55'S 151°47’E, 23 May 1955, J. Kennewell;
AMS KS1064, Weston, 32°48’S 151°27E, 7 November 1966. Females'.
AMS KS 13463, Mill Creek, Dharug National Park. 33°21'S 151°05E,
August 1983, G.P. Clancy. AMS KS1063, Kurri Kurri, 32°48'S 151°29'E,
September 1966, C. Dew.

Diagnosis. CL 7.68-8.20 (male). Differs from other infens a
group species by presence of swollen (boat-shaped) tarsi
III, IV and strong proximal curvature and distal twisting of
embolus (Fig. 81B,C); differs from lamingtonensis group
spp. by presence of full row of central cheliceral teeth (Fig.
79D) and longer PLS apical segment (Fig. 79C).

Male (holotype) — Size. Carapace length 7.73, width 7.46.
Abdomen length 8.04, width 6.70. — Colour. Basic colour
pattern. — Carapace. Slightly longer than wide, cephalic area
moderately raised. Height 3.44. Frontal width 5.16. Fovea
procurved. Mid-dorsal cephalic setae reach fovea. Anterior
strial setae present. Anterolateral carapace angle with several
weak bristles. — Eyes. Central eye region slightly raised.
Eye group width 1.92. Median ocular quadrangle length
0.74, anterior width 0.96, posterior width 1.34. Diameters:
AME 0.31, ALE 0.47, PLE 0.36, PME 0.26. —Chelicerae.
Cheliceral groove of medium width, margins diverging
distally. Groove length 2.72, middle width 0.42. Cheliceral
teeth: 22 central occupying full length of groove; 11

Gray: Australian funnel-web spiders

365

Table 23. Male morphological data—

-Hadronyche lynab

me (n = 6).

character

range

mean

character

range

mean

character

ratio

SD

CL

7.68-8.20

7.79

MtlS

13-29

22

CW/CL

0.94

0.038

CW

6.80-7.88

7.35

TalS

11-24

18

CH/CW

0.45

0.019

CH

3.06-3.44

3.29

Ti2S*

8-13

11

CFW/CL

0.70

0.04

CFW

5.10-6.12

5.45

STC2teeth

10-13

11

CHGW/L

0.16

0.013

ChGL

2.38-2.72

2.52

BulbL

3.13-3.28

3.20

SW/SL

0.76

0.029

ChGW

0.35-0.44

0.41

EmbL

2.28-2.38

2.32

LL/LW

0.87

0.049

ChGCT

18-23

21

BulbW

1.02-1.16

1.10

PLSAPW/L

0.36

0.035

LL

1.22-1.46

1.32

EmbmidW

0.24-0.30

0.27

BulbW/EmbL

0.47

0.027

LW

1.33-1.63

1.52

PalpTibL

3.28-3.61

3.38

EmbmidW/L

0.12

0.013

CUSP

250-333

283

PalpTibW

1.52-1.70

1.61

PalpTibW/L

0.48

0.015

SL

4.47-4.88

4.68

PalpTibS

0

—

BulbL/TibL*

0.95

0.013

SW

3.24-3.88

3.57

PalpPatS

0-1

1

PLSAPW

0.36-0.73

0.48

PalpFemS

0-1

1

PLSAPL

1.04-2.04

1.34

FelS

0

PalS

0-3 (p0-l)

1

prolateral; 11 retrolateral. — Labium. A little wider than long,
apical indentation weak to absent. Length 1.31, width 1.52.
Labiosternal sigilla narrowed toward midline. Cuspules c.
316, number moderate-high. — Sternum. Ovoid. Length 4.76,
width 3.62. Posterior sigilla long, ovoid. — Palp. Tegular
area small, wider than long. Embolus moderately long;
shaft broad and strongly curved proximally, hardly tapered.
Distal shaft of embolus very strongly twisted (flange reflexed
almost 180°) with twisting extended back along shaft as a
deep longitudinal fold. Basal embolus strongly offset from
tegulum. Bulb length 3.20, width 1.11. Embolus length 2.32,
midwidth 0.29. Length of femur 3.49, patella 1.07, tibia 3.44.
Width of tibia 1.64. Spination: femur 1, patella 1, tibia 0;
spines bristle-like. Several strong sinuous bristles on distal
femur. — Legs. 4123.

Figure 79. Hadronyche lynabrae, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
( D ) cheliceral groove teeth; (E) cephalothorax and chelicerae,
dorsal; ( F ) palp, prolateral; (G) leg II, prolateral. Scale lines 1 mm.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.81

3.57

5.33

5.06

2.98

23.75

2

6.36

3.36

5.17

4.82

3.03

22.74

3

5.86

2.65

3.67

4.54

3.45

20.17

4

7.04

3.02

5.09

5.83

4.02

25.00

Tibia I width 1.20. Femora I and II lack dorsal spines.
Some anterior setae on coxae I, II thickened basally Leg I
unmodified. Tibia II without apophysis; ventral spines few,
with 3-5 grouped proximally. Distoventral tibia slightly

chelicerae, lateral; ( B ) cheliceral groove teeth; (C) sternum, labium
and maxilla; ( D ) spermathecae; (E) spinnerets; (E) cephalothorax
and chelicerae, dorsal; (G) leg III, prolateral, patella, tibia,
metatarsus. Scale lines 1 mm.

366

Records of the Australian Museum (2010) Vol. 62

Figure 81. Hadronyche lynabrae, male ( A,D-K , AMS KS52488; B,C, AMS KS1005): (A) palp, prolateral; (B) bulb; (C) embolus, distal;
( D,E ) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; (77,7) distal patella and tibia II—77,
ventral, 7, prolateral; (J,K) metatarsus II— J, ventral, K, prolateral. Scale lines: 0.3 mm, except B 1.0 mm and K 0.2 mm.

Gray: Australian funnel-web spiders

367

concave. Metatarsus II weakly sinuous to straight. Tarsi III,
IV swollen (boat-shaped) and longer than tarsi I, II. Tarsal
scopulae: leg I weak; legs II-IV moderate to strong. Distal
metatarsal scopulae legs I-IV, absent to weak. Tarsal claw
teeth legs I, II: superior 13, 13; inferior 1, 0. Trichobothria
legs I, II: tarsus 10, 9; metatarsus 12, 11; tibia p7 r7, p6 r7.
—Leg spination. Leg I: femur 0, patella 2 (pi vl), tibia 23,
metatarsus 29, tarsus 24. Leg II: femur 0, patella pi, tibia

13, metatarsus 23, tarsus 23. Leg III: femur 0, patella 15
(pi4 rl), tibia 12 (pi r2), metatarsus 23, tarsus 18. Leg IV:
femur 0 (strong bristles), patella 0, tibia 9 (r2), metatarsus

14, tarsus 27. — Abdomen. Posterior lateral spinnerets with
apical segment of moderate length. Lengths: total 2.90; basal
segment 1.10; middle 0.60; apical 1.20. Apical segment
width 0.36.

Female (paratype AMS KS13463) — Size. Carapace length
9.45, width 8.18. Abdomen length 13.60, width 10.20. —
Colour. Basic colour pattern. — Carapace. Longer than wide,
strongly raised. Height 4.49; frontal width 7.41. Cephalic
length 7.28. Fovea procurved. Mid-dorsal cephalic setae
reach fovea. Anterior strial setae present. Anterolateral
carapace angle with few bristles. — Eyes. Central eye region
slightly raised. Eye group width 2.18. Diameters: AME 0.27,
ALE 0.47, PLE 0.37, PME 0.22. Interdistances: AME-AME
0.32, AME-ALE 0.37, ALE-PLE 0.27, PLE-PME 0.13,
PME-PME 1.09. Median ocular quadrangle length 0.68,
anterior width 0.86, posterior width 1.52. — Chelicerae.
Cheliceral groove margins diverge weakly distally. Groove
length 3.26, middle width 0.67. Cheliceral teeth: 58 central
occupying full length of groove; 16 prolateral; 11 retrolateral.
— Labium. Wider than long, apical indentation weak to
absent. Length 1.73, width 2.13. Cuspules distributed in a
broad V-shape over central labium. Labiostemal sigilla entire.
— Sternum. Ovoid. Length 6.02, width 4.42. Posterior sternal
sigilla ovoid to elongate; length 1.05, width 0.51. — Palp.
Spination: tibia 2 (bristle-like), tarsus 5. Tarsal claw with 4
teeth. Trichobothria: tibia p6 r6, tarsus 10. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.56

3.88

4.59

3.67

2.11

20.81

2

5.64

3.62

3.94

3.23

2.14

18.57

3

4.96

2.89

3.06

3.11

2.65

16.67

4

6.44

3.57

4.28

4.42

3.16

21.87

Tibia I width 1.50. Metatarsus I proximal width 1.03. Coxa
I with a few thorn shaped setae frontally. Legs I, II with few
spines. Tarsal claw teeth legs I, II: superior 6, 6; inferior 2
(small), 0. Trichobothria legs I, II: tarsus 10,10; metatarsus
12, 10; tibia p7 r7, p6 r7. — Leg spination. Leg I: femur 0,
patella 0, tibia 0, metatarsus 5, tarsus 6. Leg II: femur 0,
patella 0, tibia 0, metatarsus 5, tarsus 7. Leg III. femur 0,
patella p 14, tibia 8 (p3 r2), metatarsus 16 (p6 r2), tarsus 14.
Leg IV: femur 0, patella 0, tibia 2, metatarsus 15, tarsus 29.
— Abdomen. Posterior lateral spinnerets with apical segment
of moderate length. Lengths: total 3.89; basal segment 1.70,
middle 0.85, apical 1.33. Apical segment width 0.61. —
Genitalia. Spermathecae widen markedly towards apices;
narrow bases well separated by c. 1.5x spermatheca width.
Spermatheca length 1.50, width 0.73.

Distribution. From the Hawkesbury River region near
Sydney, north to the southeastern foothills of the Barrington
Tops massif, New South Wales (Fig. 78).

Hadronyche kaputarensis n.sp.

Figures 82, 83; Table 24

Etymology. The specific epithet is taken from Mount.
Kaputar, the type locality.

Types. Holotype male: AMS KS1378, Mount Kaputar National Park, New
South Wales, 30 o ll'S 150°09'E, R. Cronin, November-December 1977.

Diagnosis. CL 9.72 (male). Differs from other infensa group
species by having the central tooth row of cheliceral groove
short and basal (Fig. 82D) and male femora I, II with a dorsal
row of bristles (Fig. 82F).

Male (holotype) — Size. Carapace length 9.72, width 8.97.
Abdomen length 10.40, width 7.96. — Colour. Basic colour
pattern. — Carapace. Longer than wide, cephalic area
moderately raised. Height 4.14, frontal width 6.49. Fovea
procurved. Mid-dorsal cephalic setae stop short of fovea.
Anterior strial setae absent. Anterolateral carapace angle with
moderately strong bristles. — Eyes. Central eye region raised.
Eye group width 2.12. Median ocular quadrangle length
0.76, anterior width 0.89, posterior width 1.43. Diameters:
AME 0.23, ALE 0.39, PLE 0.22, PME 0.12. —Chelicerae.
Cheliceral groove margins subparallel to slightly divergent
distally. Groove length 2.99, middle width 0.48. Cheliceral
teeth: 6 central, occupying proximal third of groove; 10
prolateral; 12 retrolateral. — Labium. Slightly wider than
long, apically very weakly indented. Length 1.68, width 1.75.

Figure 82. Hadronyche kaputarensis, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
(D) cheliceral groove teeth; (E) cephalothorax and chelicerae,
dorsal; (F) leg II, prolateral; (G) palp, prolateral. Scale lines 1 mm.

368

Records of the Australian Museum (2010) Vol. 62

Table 24. Male morphological data-

-Hadronyche kaputarensis (n = 1).

character

character

character

ratio

CL

9.72

MtlS

40

CW/CL

0.92

CW

8.97

TalS

21

CH/CW

0.46

CH

4.14

Ti2S

35 (pi)

CFW/CL

0.67

CFW

6.49

STC2teeth

14

CHGW/L

0.19

ChGL

2.99

BulbL

3.42

SW/SL

0.71

ChGW

0.48

EmbL

2.53

LL/LW

0.96

ChGCT*

7

BulbW

1.22

PLSAPW/L

0.34

LL

1.68

EmbmidW

0.19

BulbW/EmbL

0.48

LW

1.75

PalpTibL

4.49

EmbmidW/L*

0.08

CUSP

254

PalpTibW

1.87

PalpTibW/L

0.42

SL

5.86

PalpTibS

0

BulbL/TibL

0.76

SW

4.15

PalpPatS

0

PLSAPW

0.57

PalpFemS

0

PLSAPL

1.68

FelS

0

PalS

7 (pi)

Labiosternal sigilla entire, broad. Cuspules c. 254, number
moderate. — Sternum. Ovoid, long. Length 5.86, width 4.15.
Posterior sigilla large, ovoid. — Palp. Tegular area wider
than long. Embolus moderately long and slender, weakly
offset from tegulum; shaft gently curved. Distal embolus
weakly to moderately twisted. Bulb length 3.42, width 1.22.
Embolus length 2.43, midwidth 0.19. Length of femur 4.53,
patella 1.97, tibia 4.49. Width of tibia 1.87. Spination: femur
0; patella 0; tibia 2-3 prolateral bristle-like spines. Distal
femur with several strong, sinuous bristles. — Legs. 4123.
Legs I and II subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.32

3.94

6.97

6.43

3.64

29.30

2

8.02

3.88

6.97

6.46

3.74

29.07

3

6.97

3.20

5.47

6.10

3.96

25.70

4

8.64

3.40

7.14

7.73

4.41

31.32

Tibia I width 1.43. Femora I and II without dorsal spines,
but with long dorsal bristles; 1-2 small prolateral dorsal
or retrolateral spines may be present. Proximal metatarsus
I weakly swollen, ventral spines largest proximally. Tibia
and metatarsus I with numerous spines. Tibia II without
apophysis; with numerous ventral spines, most grouped in

and H. macquariensis (squares).

proximal half; ventral tibia gently concave. Metatarsus II
weakly sinuous, ventral spines evenly distributed. Coxal
setae unmodified. Tarsal scopulae: legs I, II weak; III, IV
moderate-strong. Distal metatarsal scopulae: legs I, II absent;
III, IV weak-moderate. Tarsal claw teeth legs I, II: superior
14, 14; inferior 2, 2. Trichobothria legs I, II: tarsus 11, 10;
metatarsus 11, 11; tibia p5 r5, p6 r6. —Leg spination. Leg
I: femur pl-2, patella 7 (pi v6), tibia 65 (pi), metatarsus
40, tarsus 21. Leg II: femur plrl, patella 2 (pi), tibia 35
(pi), metatarsus 33, tarsus 24. Leg III: Femur 0 (long,
strong bristle), patella p4, tibia 20 (p5 r5), metatarsus 33,
tarsus 21. Leg IV: Femur 0 (long bristle), patella 0, tibia 11
(r3), metatarsus 25, tarsus 26. — Abdomen. Posterior lateral
spinnerets with apical segment of moderate length. Lengths:
total 4.23; basal segment 1.53; middle 1.02; apical 1.68.
Apical segment width 0.57.

Distribution. Known only from the type locality (Fig. 83).

Hadronyche macquariensis n.sp.

Figures 83-86; Tables 25, 34

Etymology. The specific epithet refers to the Port Macquarie
region of New South Wales, where this species is common.

Types. Holotype male: AMS KS1315, Taree, New South Wales, 31°54'S
152°27'E, 31 January 1978. Paratypes (all AM). New South Wales. Males :
AMS KS836, Bellingen, 30°26’S 152°54’E, 10 October 1948, G.H. Hewitt;
AMS KS4515, Burgess Beach, near Forster, 32°1 l'S 152°32'E, 15 January
1980, G. Sanders; AMS KS832, Taree, 31°54'S 152°27’E, 21 February 1974;
AMS KS838, Timmsvale, 30°12'S 152°52’E, February 1956, H.J. Stokes;
AMS KS13531,Coffs Harbour, 30° 18’S 153°07'E, 19 December 1983, C.S.
Martin; AMS KS829, Pt. Macquarie, 31°25'S 152°55'E, 26 December 1960,
D.H. Thomson. Females : AMS KS1165, Port Macquarie, 31°25'S 152°55'E,
28 April 1954, C.M. Edwards; AMS KS7565, Allyn River, 10 December
1980, G. Anderson; AMS KS 13589, Coffs Harbour, 7 December 1983; AMS
KS13351, Gloucester, 31°59'S 151°58’E, September 1983, A. D’Ombrain;
AMS KS8775, Kerewong State Forest, nr. Taree, 31°35'S 152°33'E, 28
March 1982, H. Pamaby; AMS KS834, Pt. Macquarie, 31°25'S 152°55’E,
February 1973; AMS KS3229, Taree, 31°54’S 152°27'E, 10 May 1975.

Other material examined. New South Wales (males): AMS KS840,
Bellingen, G.H. Hewitt; AMS KS8794, Blackhead, February 1982; AMS
KS 10797, Bowraville, 28 February 1983, D. Channels; AMS KS833,
Taree, 4 March 1972, K. Walters; AMS KS6457, Tullymorgan, December
1980, Ambulance Station; AMS KS6827, Wardell, 14 January 1981, M.

Gray: Australian funnel-web spiders

369

Kajewski; AMS KS13591, Woolgoola, near Red Rock, 5 December 1983,
J.W. Kramer; AMS KS911, Chatsworth Island, near. Grafton, 20 December
1975, A. Mills; AMS KS13590, Coffs Harbour, 7 December 1983; AMS
KS14036, Coffs Harbour, 18 March 1984, C.S. Martin; AMS KS8743,
Evans Head, 29 January 1982, Y. Bridges; AMS KS 10679, Forster, 25
January 1983, A. D’Ombrain; AMS KS839, Halliday’s Point, near Manning,
25 February 1976, Manning Council; Boambee, October 1974 (ANIC);
AMS KS912, Kororo, March 1965, D.M. Lee; AMS KS837, Kranbach,
Wallamba River, 40 km from Taree, 22 March 1959, J. Bidner; AMS
KS1111, Lome State Forest, near Taree, 16 January 1978, D. Milledge;
AMS KS1112, Lome State Forest, near Taree, 17 January 1978, D. Milledge;
AMS KS10680, Maclean, 31 December 1982; AMS KS10769, Maclean,
12 October 1982; AMS KS13530, Moonee Beach, near Coffs Harbour,
19 December 1983, C.S. Martini AMS KS13532, Mullaway, near Coffs
Harbour, 19 December 1983, C.S. Martin; AMS KS826, Port Macquarie,
February 1976, A. Proudman; AMS KS827, Port Macquarie, 5 March 1976;
AMS KS13814, Smiths Lake, near Myall Lakes, 16 January 1984, J. Rawle;
AMS KS828, Taree, 18 March 1974. New South Wales (females): AMS
KS5201, Nambucca Heads, 16 May 1980, S. Johnson; AMS KS841, Port
Macquarie, February 1955, W. Cleland; AMS KS3227, Taree, 10 May 1975.

Diagnosis. CL 8.74-11.22 (male). Differs from H.
levittgreggae, H. valida, H. orana and H. kaputarensis by
having a shorter, broader embolus (EmbmidW/L 0.16) (Fig.
86B); from H. infens a by weak offset of basal embolus from
tegulum; from H. walkeri by ventral tibia II only weakly
concave, and spines loosely grouped (Fig. 86H,I); from H.
lynabrae by embolus less strongly twisted (Fig. 86B) and
tarsus I more strongly spinose (30-63 spines).

Male (holotype) — Size. Carapace length 9.32, width
9.04. Abdomen length 11.42, width 8.30. — Colour. Basic
colour pattern. — Carapace. Slightly longer than wide,
cephalic area moderately raised. Height 3.57. Frontal width
6.07. Fovea procurved, rather narrow. Mid-dorsal cephalic
setae numerous, reach fovea. Anterior strial setae present.
Anterolateral carapace angle with numerous strong bristles.

Figure 84. Hadronyche macquariensis, male: (A) sternum,
labium and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C)
spinnerets; (D) palp, prolateral; (E) cephalothorax and chelicerae,
dorsal; (F) cheliceral groove teeth; (G) leg II, prolateral. Scale
lines 1 mm.

— Eyes. Central eye region raised. Eye group width 2.37.
Median ocular quadrangle length 0.93, anterior width 1.16,
posterior width 1.64. Diameters: AME 0.33, AFE 0.57, PEE
0.39, PME 0.30. — Chelicerae. Dorsal paturon obliquely
ridged proximally. Cheliceral groove margins divergent.
Groove length 2.93, middle width 0.53. Cheliceral teeth:
18 central, occupying full length of groove; 10 prolateral;
9 retrolateral. — Labium. Slightly wider than long, not
or weakly apically indented. Fength 1.68, width 1.74.
Fabiosternal sigilla narrowed toward midline. Cuspules c.
274, number moderate-high. — Sternum. Ovoid. Fength 5.95,
width 4.32. Posterior sigilla ovoid, long. — Palp. Tegular area
wider than long. Embolus of moderate length, base weakly
offset from tegulum. Embolus shaft curved, broad and blade¬
like, not tapered. Distal part of embolus strongly twisted and
grooved, apical flange reflexed. Bulb length 3.90, width 1.37.
Embolus length 2.80, midwidth 0.40. Length of femur 4.39,
patella 1.76, tibia 4.67. Width of tibia 2.21. Spination: none,
sinuous bristles on distal femur. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.45

3.85

7.01

6.94

3.80

30.05

2

7.85

4.02

6.97

6.40

3.77

29.01

3

7.41

3.37

5.78

6.83

4.08

27.47

4

8.77

3.77

7.07

8.09

4.49

32.19

Tibia I width 1.43. Femora I and II without spines. Leg
I with numerous tibial and metatarsal ventral spines.
Metatarsus I slightly thickened proximally, proximal ventral
spines largest. Tibia II without apophysis, ventrally weakly
concave; ventral spines loosely grouped proximocentrally,
fewer spines distally. Metatarsus II unmodified. Coxal hairs
unmodified. Tarsal scopulae: legs I, II weak; legs III, IV

Figure 85. Hadronyche macquariensis, female: (A) sternum,
labium and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C)
cheliceral groove teeth; (D) spermathecae; (E) cephalothorax and
chelicerae, dorsal; ( F) leg III, prolateral, patella, tibia, metatarsus;
(G) spinnerets. Scale lines 1 mm.

370

Records of the Australian Museum (2010) Vol. 62

Figure 86. Hadronyche macquariensis, male (A,D-J, AMS KS10679; B,C, AMS KS4515): (A) palp, prolateral; ( B ) bulb; (C) embolus,
distal; ( D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; (H,I) distal patella and tibia
II— H, ventral, I, prolateral; (7), metatarsus II, prolateral. Scale lines: 0.3 mm, except A,B,1 1.0 mm.

Gray: Australian funnel-web spiders

371

Table 25. Male morphological data—

-Hadronyche macquariensis (n =

15).

character

range

mean

character

range

mean

character

ratio

SD

CL

8.74-11.22

9.79

MtlS

34-76

53

CW/CL

0.92

0.029

CW

8.15-10.09

9.05

TalS*

30-63

43

CH/CW

0.42

0.033

CH

3.28—4.51

3.79

Ti2S

7-38 (p0-l)

23

CFW/CL

0.64

0.037

CFW

5.10-6.83

5.99

STC2teeth

10-15

12

CHGW/L

0.18

0.02

ChGL

2.58-3.32

2.82

BulbL

3.26-4.16

3.64

SW/SL

0.77

0.04

ChGW

0.43-0.66

0.51

EmbL

2.21-2.80

2.54

LL/LW

0.93

0.038

ChGCT

18-34

25

BulbW

1.22-1.64

1.39

PLSAPW/L

0.32

0.052

LL

1.36-1.96

1.64

EmbmidW

0.33-0.48

0.40

BulbW/EmbL

0.55

0.037

LW

1.60-2.06

1.76

PalpTibL

3.85-5.03

4.65

EmbmidW/L*

0.16

0.014

CUSP

231-346

283

PalpTibW

1.93-2.42

2.16

PalpTibW/L

0.46

0.02

SL

5.20-6.33

5.77

PalpTibS

0-3

1.40

BulbL/TibL

0.78

0.035

SW

3.94-4.92

4.47

PalpPatS

0

—

PLSAPW

0.48-0.88

0.60

PalpFemS

0-2

1

PLSAPL

1.67-2.36

1.86

FelS

0

PalS

0-13

3

moderate-strong. Distal metatarsal scopulae: legs I, II absent;
legs III, IV moderate. Tarsal claw teeth legs I, II: superior
12, 11; inferior 3, 2. Trichobothria legs I, II: tarsus 14, 13;
metatarsus 14, 14; tibia p9 r7, p8 r8. — Leg spination. Leg
I: femur 0, patella 1, tibia 44, metatarsus 57, tarsus 41. Leg
II: femur 0, patella 0, tibia 26, metatarsus 44, tarsus 49. Leg
III: femur 0, patella 3, tibia 26, metatarsus 44, tarsus 49.
Leg IV: femur 0, (long bristles present), patella 0, tibia 6
(rl), metatarsus 26, tarsus 47. — Abdomen. Posterior lateral
spinnerets: apical segment moderately long. Lengths: total
5.08, basal segment 1.72; middle 1.15; apical 2.21. Apical
segment width 0.71.

Female (paratype AMS KS1165) — Size. Carapace length
11.25, width 10.03. Abdomen length 14.80, width 11.80.
— Colour. Basic colour pattern. — Carapace. Longer than
wide, cephalic area strongly raised. Height 5.13; frontal width
8.64. Cephalic length 8.09. Fovea procurved, anterior margin
indented. Mid-dorsal cephalic setae numerous, extend back
to fovea. Anterior strial setae numerous. Strial and marginal
setae numerous. Anterolateral carapace angle with many weak
bristles. — Eyes. Eye region sessile to weakly raised centrally.
Eye group width 2.75. Diameters: AME 0.39, ALE 0.62, PLE
0.54, PME 0.34. Interdistances: AME-AME 0.34, AME-ALE
0.27, ALE-PLE 0.20, PLE-PME 0.10, PME-PME 1.19.
Median ocular quadrangle length 0.88, anterior width 1.10,
posterior width 1.78. — Chelicerae. Cheliceral groove margins
diverge distally. Groove length 3.26, middle width 0.80.
Cheliceral teeth: 34 central occupying full length of groove
in an irregular double row; 13 prolateral; 11 retrolateral. —
Labium. Slightly wider than long, anterior margin very weakly
indented. Length 2.07, width 2.28. Cuspules occupying
anterior three quarters of labium. Labiostemal sigilla entire.
— Sternum. Ovoid, moderately wide. Length 6.80, width 5.68.
Posterior sternal sigilla ovoid: length 1.09, width 0.54. — Palp.
Spination: tibia 4, tarsus 8. Trichobothria: tibia p6 r7, tarsus
10. Tarsal claws with 6 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

8.09

4.56

6.12

5.17

3.06

27.00

2

7.31

4.08

5.51

4.79

2.92

23.61

3

6.53

3.74

4.39

4.85

3.40

22.91

4

8.19

4.08

5.98

6.09

3.74

28.08

Tibia I width 2.14. Metatarsus I proximal width 1.43.
Coxae I, II with setal bases slightly thickened, otherwise
unmodified. Tarsal claw teeth legs I, II: superior 9,9; inferior
3, 3. Trichobothria legs I, II: tarsus 12,12; metatarsus 12,12;
tibia p8 r8, p9 r8. — Leg spination. Leg I: femur 0, patella 0,
tibia 2, metatarsus 10, tarsus 6. Leg II: femur 0, patella pi,
tibia 4 (pi), metatarsus 14, tarsus 9. Leg III: femur 4 (pd3,
rdl) patella7 (p6 rl), tibia 10 (p3 r2), metatarsus 20 (p6 r2),
tarsus 14. Leg IV: femur 0, patella 0, tibia 8 (r2), metatarsus
21, tarsus 24 — Abdomen. Sparse cover of weak hairs and
bristles. Posterior lateral spinnerets with apical segment of
medium length. Lengths: total 4.86; basal segment 2.11,
middle 0.88, apical 1.94. Apical segment width 0.82. —
Genitalia. Spermathecae rather short, broad, separated
basally by a spermathecal width. Length 1.43, width 0.75.

Distribution. Mid northeastern region of New South Wales
(Fig. 83).

Hadronyche walkeri n.sp.

Figures 87-90; Tables 26, 34

Etymology. The species is named for Mr Pat Walker, North
Queensland Naturalist and Photographer.

Types. Holotype male: S389 (QMB). Devils Pinch, north of Armidale,
New South Wales, 33°09'S 150°17'E, C. Anderson, 1 February 1979.
Paratypes. New South Wales. Males:Q MB S382 (QMB), Armidale, 30°30'S
151°39’E, 17 May 1979, K. McWilliam. Others all AM: AMS KS936, Ben
Lomond, 30°00'S 151°39'E, February 1963, M. Trudgeon; AMS KS943,
“Wongwibinda”, 48km E. of Guyra, 30°12’S 151°40'E, December 1952,
M. Wyndham; AMS KS944, Bullock Creek, Point. Lookout, New England
National Park, 30°29’S 152°30’E, 25 November 1969, M. Gray; AMS
KS4455, Walcha, 30°58'S 151°35E, 11 January 1980, C. Easton; AMS
KS44614, Guyra, 30°13'S 151°40E, May 1971, D.I. Clay; AMS KS936,
Ben Lomond Public School, 30°01'S 151°40E, February 1963, M. Trudgeon.
Females: AMS KS15750, Styx River near Jeogla, 30°35'S 152°08'E, 8 June
1979; AMS KS13805, Ulong, nr. Dorrigo, 30°13’S 152°53’E, January 1980,
C. Martin; AMS KS7524, Lower Wattle Flat, Pt. Lookout, New England
National Park, 30°29'S 152°30’E, 12 May 1981, G. Hunt; AMS KS14037,
Dorrigo, 30°19'S 152°43'E, 18 March 1984, C. Martin.

Other material examined. New South Wales (males): AMS KS937, Ben
Lomond, February 1963, M. Trudgeon; QMB S675, Black Mountain, 28
February 1977, Snell; AMS KS940, Glen Innes, February 1952, ? Every;
QMB S384,42 km E. of Guyra, January 1972; AMS KS945, Point. Lookout,

372

Records of the Australian Museum (2010) Vol. 62

New England National Park, 26 November 1969, M. Gray; QMB S386,
“Newholme”, near Armidale, 2 February 1974, Jen ki ns. New South Wales
(females): AMS KS13596, Dorrigo, 22 December 1983; AMS KS14384,
Ulong, nr. Dorrigo, 26 April 1984; AMS KS7525-7, Lower Wattle Flat,
Point Lookout area, New England National Park, 12 May 1981, G. Hunt;
AMS KS14038, Dorrigo, 18 March 1984, C. Martin.

Diagnosis. CL 7.34-10.54 (male). Differs from H. infensa
and H. macquariensis by tibia II ventrally sinuous/concave
with strong ventral spines clustered proximocentrally (Figs.
87G; 89H,I); from H. infensa by basal embolus weakly offset
from tegulum (Fig. 89B);from H. lynabrae by tarsi III, IV
not swollen, and embolus less twisted; and from H. valida,

H. orana, H. kaputarensis and H. levittgreggae by greater
width of embolus (EmbmidW/L 0.14) (Fig. 89B).

Male (holotype) — Size. Carapace length 8.69, width 8.43.
Abdomen length 8.84, width 6.80. — Colour. Basic colour
pattern. Posterior abdominal chevrons often joined across
abdomen. — Carapace. A little longer than wide, cephalic
area moderately raised. Height 3.63. Frontal width 5.82.
Fovea procurved. Mid-dorsal cephalic setae almost reach
fovea. Anterior strial setae few to absent. Anterolateral
carapace angle with a few strong bristles. — Eyes. Central eye
region strongly raised. Eye group width 2.02. Median ocular
quadrangle length 0.80, anterior width 0.92, posterior width

I. 42. Diameters: AME 0.64, ALE 0.80, PLE 0.62, PME
0.50. — Chelicerae. Cheliceral groove margins divergent.
Groove length 2.52, middle width 0.46. Cheliceral teeth:
32 central, occupying full length of groove; 13 prolateral;
12 retrolateral. — Labium. Slightly wider than long, weakly
apically indented. Length 1.68, width 1.72. Labiosternal
sigilla entire. Slight transverse groove behind cuspules.
Cuspules c. 344, number high. — Sternum. Ovoid, long.
Length 5.51, width 4.02. Posterior sigilla ovoid, elongate.
— Palp. Tegular area wider than long. Embolus rather short,

Figure 87. Hadronyche walkeri, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cephalo-
thorax and chelicerae, dorsal; ( D ) spinnerets; (E) cheliceral groove
teeth; ( F ) palp, prolateral; (G) leg II, prolateral. Scale lines 1 mm.

not or very weakly offset from tegulum. Embolus shaft gently
curved, broad, not tapered. Distal part of embolus moderately
twisted with twisting extended back along shaft as a deep
longitudinal fold; distal flange moderately reflexed. Bulb
length 3.32, width 1.40. Embolus length 2.28, midwidth
0.39. Length of femur 4.10, patella 1.76, tibia 3.98. Width
of tibia 2.08. Spination: tibia with 2 prolateral bristle like
spines, absent elsewhere. Distal femur with several sinuous
bristles. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.46

3.53

5.78

5.49

3.44

25.70

2

7.01

3.57

5.74

5.33

3.36

25.01

3

6.32

2.99

4.56

5.54

3.67

23.08

4

7.75

3.48

5.77

6.80

4.32

28.12

Tibia / width 1.52. Femora I and II without spines. Metatarsus
I slightly thickened proximally, proximal ventral spines
largest. Tibia II without apophysis, moderately sinuous
(concave distoventrally), ventral spines rather large, grouped
proximally. Distal tibia II concave ventrally, lacking spines
(except apically). Metatarsus II weakly sinuous-straight,
strong ventral spines centrally. Coxal hairs unmodified.
Tarsal scopulae: leg I weak; legs II-IV moderate to strong.
Distal metatarsal scopulae: leg I absent; legs II-IV weak
to moderate. Tarsal claw teeth legs I, II: superior 12, 12;
inferior 2,2. Trichobothria legs I, II: tarsus 11,9; metatarsus
15, 14; tibia p8 r7, p8 r8. — Leg spination. Leg I: femur 0,
patella 5 (pi), tibia 34, metatarsus 25, tarsus 27. Leg II:
femur 0, patella 0, tibia 13, metatarsus 23, tarsus 21. Leg
III: femur 0, patella p6, tibia 9 (p2 r2), metatarsus 22, tarsus
22. Leg IV: femur 0 (long bristles present), patella 0, tibia 8

Figure 88. Hadronyche walkeri, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) sperma-
thecae; (D) cephalothorax and chelicerae, dorsal; (E) spinnerets;
(F) cheliceral groove teeth; (G) leg III, prolateral, patella, tibia,
metatarsus. Scale lines 1 mm.

Gray: Australian funnel-web spiders

373

Figure 89. Hadronyche walkeri, male. (A,D-K, AMS KS44614; B,C, AMS KS936): (A) palp, prolateral; ( B ) bulb; (C) embolus, distal; (D,E)
distal patella and tibia I —D, ventral, E, prolateral; ( F,G ) metatarsus I —F, ventral, G, prolateral; ( H,I) distal patella and tibia II —H, ventral
(image foreshortened), I, prolateral; ( J,K) metatarsus II —J, ventral, K, prolateral. Scale lines: 0.3 mm, except B 1.0 mm and D-G 0.2 mm.

374

Records of the Australian Museum (2010) Vol. 62

Table 26. Male morphological data— Hadronyche walkeri (n = 11).

character

range

mean

character

range

mean

character

ratio

SD

CL

7.34-10.54

8.77

MtlS

23-55

36

CW/CL

0.91

0.032

CW

6.60-9.28

8.01

TalS

8-38

21

CH/CW

0.41

0.031

CH

2.91-3.85

3.34

Ti2S

10-29

20

CFW/CW

0.65

0.034

CFW

5.50-6.31

5.65

STC2teeth

10-15

12

CHGW/L

0.17

0.027

ChGL

2.21-2.86

2.50

BulbL

2.63-3.40

3.02

SW/SL

0.77

0.025

ChGW

0.31-0.55

0.42

EmbL

1.76-2.40

2.05

LL/LW

0.92

0.053

ChGCT

5-46

23

BulbW

1.15-1.40

1.25

PLSAPW/L

0.35

0.034

LL

1.11-1.69

1.44

EmbmidW

0.22-0.39

0.27

BulbW/EmbL* 0.62

0.04

LW

1.31-1.72

1.56

PalpTibL

3.28-4.49

3.93

EmbmidW/L*

0.14

0.02

CUSP

144-344

240

PalpTibW

1.72-2.11

1.94

PalpTibW/L

0.49

0.03

SL

4.26-5.83

5.17

PalpTibS

0-2

<1

BulbL/TibL

0.78

0.037

SW

3.37-4.49

4.02

PalpPatS

0

—

PLSAPW

0.37-0.60

0.49

PalpFemS

0

—

PLSAPL

1.07-1.70

1.41

FelS

0

—

PalS*

0-13 (pO-2)

6

(rl), metatarsus 22, tarsus 27 — Abdomen. Posterior lateral
spinnerets: apical segment moderately long. Lengths: total
4.44; basal segment 1.72, middle 1.04, apical 1.68. Apical
segment width 0.60.

Female (paratype AMS KS15750) — Size. Carapace length
12.34, width 10.13. Abdomen length 15.30, width 11.70.
— Colour. Basic colour pattern. — Carapace. Longer than
wide, moderately raised. Height 5.24. Frontal width 8.84.
Cephalic length 8.67. Fovea strongly procurved. Mid¬
dorsal cephalic setae numerous, reach fovea. Anterior strial
setae numerous. Strial setae well developed. Anterolateral
carapace angle with several moderately strong bristles. —
Eyes. Eye region sessile to weakly raised centrally. Eye group
width 2.70. Diameters: AME 0.34, ALE 0.54, PLE 0.55,
PME 0.35. Interdistances: AME-AME 0.34, AME-ALE 0.29,
ALE-PLE 0.24, PLE-PME 0.14, PME-PME 1.17. Median
ocular quadrangle length 0.95, anterior width 1.05, posterior
width 1.84. — Chelicerae. Cheliceral groove margins diverge
distally. Groove length 3.58, middle width 0.82. Cheliceral
teeth: 79 central, in two to three rows occupying full length
of groove; 15 prolateral; 12 retrolateral. — Labium. Almost
as long as wide, anterior margin broadly indented. Length
2.43, width 2.68. Cuspules occupying anterior half of
labium. Labiosternal sigilla narrowed to divided in midline.
— Sternum. Ovoid. Length 7.55, width 5.71. Posterior sigilla
elongate: length 1.73, width 0.61. — Palp. Spination: tibia 6,
tarsus 7. Trichobothria: tibia p8 r8, tarsus 14. Tarsal claws
with 9 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus

Tarsus

Total

1

7.46

3.53

5.78

5.49

3.44

25.70

2

7.01

3.57

5.74

5.33

3.36

25.01

3

6.32

2.99

4.56

5.54

3.67

23.08

4

7.75

3.48

5.77

6.80

4.32

28.12

Tibia I width 2.04. Metatarsus I proximal width 1.43. Coxae
I, II with frontal setae thickened basally. Tarsal claw teeth
legs I, II: superior 12, 11; inferior 4, 4. Trichobothria legs I,
II: tarsus 15, 16; metatarsus 20, 20; tibia p7 r8, p9 rlO. —
Leg spination. Leg I: femur 0, patella 0, tibia 4, metatarsus
14, tarsus 8. Leg II: femur 0, patella 0, tibia 7, metatarsus
16, tarsus 9. Leg III: femur 0, patella pd4, tibia 7 (p2 r2),
metatarsus 25 (pd4 rd3) tarsus 18. Leg IV: femur 0, patella
0, tibia 6 (r2), metatarsus 18 (rl), tarsus 25. — Abdomen.

Posterior lateral spinnerets with long apical segment.
Lengths: total 6.76; basal segment 2.62, middle 1.63, apical
2.53. Apical segment width 0.75. — Genitalia. Spermathecae
straight, digitiform, rather long, distal third slightly enlarged;
well separated (by about twice the width of a spermatheca).
Spermatheca length 1.53, width 0.48.

Distribution. Great Dividing Range, New South Wales from
the Armidale region east to the Dorrigo region.

Comments. Specimens from the Barrington region may
belong to this species.

Hadronyche levittgreggae n.sp.

Figures 90-93; Tables 27, 34

Etymology. The species is named in recognition of the
late Vera Levitt-Gregg, former Associate of the Australian
Museum, naturalist and funnel-web worker.

Types. Holotype male: AMS KS 13528, Annangrove, New South Wales,
33°39'S 150°56'E, P.G. Nield, November 1978. Paratypes (all AM). New
South Wales. Males : AMS KS2076, Fairs Creek, Cattai area, 33°33'S
150°55'E, 9 November 1978, R.H. Eastment; AMS KS 1062, Dural, 33°40’S
151°01'E, 5 June 1977; AMS KS2066, Kellyville, 33°43’S 150°57’E,
6 November 1978, M.J. Fletcher; AMS KS1059, Kenthurst, 33°41’S

Figure 90. Collection records for Hadronyche walkeri (squares)
and H. levittgreggae (circles).

Gray: Australian funnel-web spiders

375

Table 27. Male morphological data — Hadronyche levittgreggae (n = 11).

character

range

mean

character

range

mean

character

ratio

SD

CL

8.72-10.30

9.69

MtlS

27-43 (pO-2)

34

CW/CL

0.95

0.027

CW

8.61-10.17

9.15

TalS

10-20

13

CH/CW

0.42

0.019

CH

3.57-4.06

3.86

Ti2S

19-46 (pO-2)

31

CFW/CL

0.69

0.023

CFW

5.99-7.14

6.67

STC2teeth

12-14

13

CHGW/L

0.20

0.017

ChGL

2.57-2.99

2.70

BulbL

4.62-5.44

4.86

SW/SL

0.72

0.031

ChGW

0.48-0.59

0.53

EmbL

3.53—4.15

3.80

LL/LW

0.96

0.04

ChGCT

15-27

22

BulbW

1.21-1.36

1.26

PLSAPW/L

0.37

0.043

LL

1.50-1.78

1.67

EmbmidW

0.21-0.28

0.25

BulbW/EmbL* 0.34

0.021

LW

1.66-1.84

1.73

PalpTibL

4.51-5.66

5.26

EmbmidW/L*

0.07

0.005

CUSP

210-345

285

PalpTibW

1.82-2.45

1.99

PalpTibW/L*

0.38

0.033

SL

5.49-6.41

5.92

PalpTibS*

2-3

2

BulbL/TibL

0.93

0.058

SW

3.77-4.59

4.30

PalpPatS

0

—

PLSAPW

0.39-0.69

0.56

PalpFemS

0-1

<1

PLSAPL

1.27-1.73

1.51

FelS

0

0

PalS

3-10 (pi—3)

6

150°59'E, 12 November 1969, A. Adamson; AMS KS7336, Lower Portland,
Hawkesbury River, 33°25'S 150°53'E, February 1980; AMS KS6274,
Tennyson, nr. Windsor, 33°31'S 150°44'E, 8 August 1980, M. Gray & R.
McDonald; AMS KS23457, Galston, 33°39'S 151°03'E, 07 Nov 1990, T.
Dixen. Females: AMS KS1061, Annangrove, 33°39'S 150°56'E, R. Redfern,
January 1976; AMS KS9036, Dural/Kenthurst area, 33°38'S 151°01’E, 6
May 1982, M. Gray; AMS KS6273, Tennyson, near. Windsor, 33°31'S
150°44E, 8 August 1980, M. Gray & R. McDonald; AMS KS2964, Arcadia,
33°37’S 151°02E, 10 May 1979, Sharpies.

Other material examined. New South Wales (males): AMS KS1060,
Kenthurst, 1969, A. Adamson; AMS KS6073, Kenthurst, 26 October 1980;
AMS KS8378, Kenthurst, 3 November 1981, L. Taylor; AMS KS 10795,
Kenthurst, 7 December 1982, L. Millard.

Figure 91. Hadronyche levittgreggae, male: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
( D ) cephalothorax and chelicerae, dorsal; (E) leg II, prolateral;
(F) palp, prolateral; (G) cheliceral groove teeth. Scale lines 1 mm.

Diagnosis. CL 8.72-10.30 (male)Differs from H. infensa,
H. lynabrae, H. macquariensis and H. walkeri by relatively
long, slender embolus (EmbmidW/L 0.07); from//, orana by
strongly spinose tibiaII(19-46 spines) (Fig. 93H,I); from//.
valida by longer, narrower embolus (EmbL 3.53-4.15) (Fig.
93B); and from H. kaputarensis by long central cheliceral
tooth row.

Male (holotype) — Size. Carapace length 8.94, width 8.61.
Abdomen length 8.85 width 6.97. — Colour. Basic colour
pattern. — Carapace. Slightly longer than wide, moderately
raised. Height 3.84. Frontal width 6.36. Fovea procurved,
narrow. Mid-dorsal cephalic setae reach fovea. Anterior strial
area with few setae. Anterolateral angle of carapace with
weak bristles. — Eyes. Sessile to slightly raised. Eye group
width 2.32. Median ocular quadrangle length 0.84, anterior

Figure 92. Hadronyche levittgreggae, female: (A) sternum,
labium and maxilla; ( B ) cephalothorax and chelicerae, lateral;
(C) spinnerets; (D) cephalothorax and chelicerae, dorsal; (E)
spermathecae; ( F) cheliceral groove teeth; (G) leg III, prolateral,
patella, tibia, metatarsus. Scale lines 1 mm.

376

Records of the Australian Museum (2010) Vol. 62

Figure 93. Hadronyche levittgreggae, male (. A,D-K , AMS KS23457; B,C, AMS KS2066): (A) palp, prolateral; (B) bulb; (C) embolus,
distal; ( D,E) distal patella and tibia I— D, ventral, E, prolateral; (F,G) metatarsus I— F, ventral, G, prolateral; ( H,I) distal patella and tibia
II— H, ventral, /, prolateral; (J,K) metatarsus II— J, ventral (image foreshortened), K, prolateral. Scale lines: 1.0 mm, except A,F,G 0.3
mm and K 0.2 mm.

Gray: Australian funnel-web spiders

377

width 1.07, posterior width 1.53. Diameters: AME 0.26, ALE
0.46, PLE 0.48, PME 0.25. — Chelicerae. Cheliceral groove
margins weakly divergent. Groove length 2.57, middle
width 0.51. Cheliceral teeth: 15 central, running full length
of groove; 11 prolateral; 9 retrolateral. — Labium. Slightly
wider than long. Length 1.50, width 1.70. Labiosternal
sigilla entire. Cuspules c. 262, number moderate to high.
— Sternum. Ovoid; length 5.67, width 4.19. Posterior
sigilla ovoid, elongate. — Palp. Tegular area slightly wider
than long. Embolus shaft long, slender and gently curved,
with little distal taper. Distal embolus moderately twisted.
Embolus base offset from tegulum. Bulb length 4.68, width
1.24. Embolus length 3.76, midwidth 0.21. Tibia long.
Length of femur 4.00, patella 1.78, tibia 5.00. Width of tibia
1.86. Spination: tibia 3 (bristle-like). Sinuous bristles on
distal femur. — Legs. 4123. Legs I, II subequal.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.30

3.40

6.27

5.78

3.44

26.19

2

6.90

3.75

6.38

5.79

3.44

26.04

3

6.11

3.28

5.17

5.56

3.90

24.02

4

7.38

3.57

6.56

6.89

4.43

28.83

Tibia I width 1.44. Femora I, II lack dorsal spines. Femora III,
IV with long dorsal bristles. Metatarsus I slightly thickened
proximally, larger ventral spines grouped proximally.
Tibia II sinuous, weakly concave distoventrally, ventral
spines grouped proximally. Metatarsus II slightly sinuous,
apophysis absent. Coxal hairs normal. Tarsal scopulae: legs
I, II weak; legs III, IV strong. Distal metatarsal scopulae
weak. Tarsal claw teeth legs I, II: superior 14; inferior III,
IV. Trichobothria legs I, II: tarsus 13,12; metatarsus 15, 17;
tibia p8 r7, p8 r7. — Leg spination. Leg I: femur 0, patella
7 (pi), tibia 65, metatarsus 30, tarsus 11. Leg II: femur 0,
patella 1-2 (p0-l), tibia 28, metatarsus 21, tarsus 12. Leg III:
femur 0, patella p5, tibia 7 (p2 rl), metatarsus 23, tarsus 17.
Leg IV: femur 0, patella rl, tibia 4 (rl), metatarsus 16, tarsus
17 — Abdomen. Posterior lateral spinnerets with moderately
long apical segment. Lengths: total 3.91; basal segment 1.64,
middle 0.88, apical 1.39; apical width 0.58.

Female (paratype AMS KS1061) — Size. Carapace length
10.80, width 9.28. Abdomen length 11.42, width 8.74. —
Colour. Basic colour pattern. — Carapace. Longer than
wide, cephalic area strongly raised. Height 5.02, frontal
width 8.01. Cephalic length 7.51. Fovea narrow, procurved.
Mid-dorsal cephalic setae numerous, extend back to fovea.
Anterior strial setae present. Anterolateral carapace angle
with several weak bristles. — Eyes. Central eye region weakly
raised. Eye group width 2.62. Diameters: AME 0.34, ALE
0.62, PLE 0.46, PME 0.29. Interdistances: AME-AME 0.55,
AME-ALE 0.25, ALE-PLE 0.26, PLE-PME 0.15, PME-
PME 1.32. Median ocular quadrangle length 0.80, anterior
width 1.16, posterior width 1.85. — Chelicerae. Cheliceral
groove wide, margins diverge distally. Groove length 2.94,
middle width 0.85. Cheliceral teeth: 44 central occupying
full length of groove in 2-3 irregular rows; 14 prolateral;
10 retrolateral. — Labium. Almost as long as wide, anterior
margin not indented. Length 2.14, width 2.21. Cuspules
occupying anterior three quarters of labium. Labiosternal
sigilla narrow, entire. — Sternum. Ovoid. Length 8.02, width
5.20. Posterior sternal sigilla elongate: length 1.19, width
0.46. — Palp. Spination: tibia 2-3, tarsus 9. Trichobothria:
tibia p7 r7, tarsus 20. Tarsal claws with 8 teeth. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

7.48

4.05

5.70

4.52

2.80

24.55

2

6.54

3.79

5.18

4.24

2.62

22.37

3

5.92

3.28

4.01

4.25

2.99

20.45

4

7.49

4.05

5.51

5.47

3.52

26.04

Tibia I width 1.90. Metatarsus I proximal width 1.21. Coxae
I, II with basally thickened setae frontally. Tarsal claw teeth
legs I, II: superior 9, 9; inferior 4, 4. Trichobothria legs I,
II: tarsus 16, 13; metatarsus 20,17; tibia p8 r8, p8 r7. — Leg
spination. Leg I: femur 0, patella 0, tibia 1, metatarsus 10,
tarsus 8. Leg II: femur 0, patella pi, tibia 1-2, metatarsus
12, tarsus 7. Leg III: femur 0, patella p4, tibia 6 (p2 r2),
metatarsus 14 (p4 rl), tarsus 14. Leg IV: femur 0 patella 0,
tibia 2 (rl), metatarsus 13, tarsus 16. — Abdomen. Posterior
lateral spinnerets with moderately short apical segment.
Lengths: total 4.50; basal segment 1.83, middle 1.29, apical
1.43. Apical segment width 0.76. — Genitalia. Spermathecae
long and thin, digitiform, separated basally but gently curved
so that apices are contiguous. Length 2.58, width 0.54.

Distribution. Hawkesbury River region, northwest of
Sydney, New South Wales (Fig. 90).

lamingtonensis species group

Description. (Figs. 3G, 94-102; Tables 28-31). Small
to medium sized atracine spiders (CL 5.0-8.0). Males
without leg II apophyses; femora I, II typically without
dorsal spines (rarely a bristle-like spine on femur II).
Middle haematodocha usually exposed between tegulum
and subtegulum. Carapace broad, strongly raised (CW/CL
0.48-0.51). Cheliceral groove narrow, central teeth few,
basal. Posterior lateral spinnerets with short apical segment
(PLSAPW/L 0.53-0.63). Labium relatively short (LL/LW
0.74-0.83), cuspule number usually low (44-102), but
moderate in H. raveni (mean = 213). Sternum moderately
wide. Several species have swollen or “boat-shaped” tarsi III
and IV. Palpal patella wider than femur; tibia without spines,
rather short and basally broad.

Included species. Hadronyche lamingtonensis n.sp., H.
annachristiae n.sp., H. raveni n.sp., H. monteithi n.sp., H.
anzses Raven.

Distribution. Northeastern New South Wales to northeastern
Queensland.

Comments. The morphology of the disjunct north
Queensland species, H. anzses, clearly justifies its inclusion
within this species group. Swollen (“boat-shaped”) tarsi III
and IV are present in several species (H. annachristiae, H.
anzses, H. monteithi and, more weakly, in H. raveni). This
character provides an additional taxonomic link between H.
anzses and its southern relatives. Two undescribed species
are known from the Border Ranges region of NSW.

The lamingtonensis group species probably represent the
remnants of a former more continuous distribution in the
Tertiary rainforests of eastern Australia.

Note on Anepsiada ventricosa (Rainbow & Pulleine),

AMS KS9624, old catalogue number K40965. This species
was described for a small female specimen from Cloncurry,

378

Records of the Australian Museum (2010) Vol. 62

central Queensland (Rainbow & Pulleine, 1918). The label
data was subsequently found to be in error, resulting in
the correction of the locality to Mount Tamborine in the
Border Ranges area of southeast Queensland The species
was proposed for synonymy as a juvenile of Atrax valida
(Gray, 1984). However, a recent re-examination of the type
indicates that it is a female, and suggests that it belongs to
the lamingtonensis species group of Hadronyche. At present,
no other female specimens of this species group are known
and the type specimen is in poor condition. If these findings
are confirmed by additional female material, the synonymy
of this species with A. valida (after Gray, in Main, 1985),
and its status, will need to be reviewed.

Hadronyche raveni n.sp.

Figures 94-96; Table 28

Etymology. The species is named for Dr Robert Raven,
Senior Curator of Arachnology at the Queensland Museum
and a distinguished spider researcher.

Types. Holotype male: QMB S161 (QMB). Bouloumba Creek, Conondale
Range, Queensland, 26°31'S 152°39'E, G.B. and S.R. Monteith, 29
November 1974-22 February 1975. Paratypes (QMB). Queensland. Males:
QMB S179, 19 April-23 August 1975; QMB SI80, 22 February-14 April
1975: other data as for holotype.

Diagnosis. CL 5.82-5.94 (male). Small atracine spiders.
Differs from other lamingtonensis group species by having
cheliceral groove margins diverging distally (Fig. 94C)
and 2-4 times as many labial cuspules (202-224); from H.

Figure 94. Hadronyche raveni, male: (A) cephalothorax and
chelicerae, dorsal; ( B ) cephalothorax and chelicerae, lateral; (C)
cheliceral groove teeth; ( D ) leg II, prolateral; (E) sternum, labium
and maxilla; (F) palp, prolateral; (G) spinnerets; (H) palpal organ.
Scale lines 1 mm.

annachristiae and H. monteithi by the short, broad embolus;
from H. lamingtonensis by tibia II with few spines (2-6)
(Fig. 95D); and from H. anzses by having a full prolateral
row of cheliceral teeth.

Male (holotype) — Size. Carapace length 5.82, width 5.70.
Abdomen length 5.90, width 4.29. — Colour. Basic colour
pattern. — Carapace. Almost as wide as long, cephalic area
strongly raised. Height 2.75. Frontal width 4.47. Fovea wide,
procurved. Mid-dorsal cephalic setae do not reach fovea.
Anterior strial setae absent. Anterolateral carapace angle
with a few weak bristles. — Eyes. Central eye region slightly
raised. Eye group width 1.71. Median ocular quadrangle
length 0.70, anterior width 0.85, posterior width 1.22.
Diameters: AME 0.28, ALE 0.42, PLE 0.32, PME 0.26.

Figure 95. Hadronyche raveni, male (S179): (A) palp, prolateral;
(B) distal patella and tibia I, ventral; (C) metatarsus I, ventral; (D)
distal patella and tibia II, ventral. Scale lines: A, B, 0.3 mm, C 0.1
mm and D 0.2 mm.

Gray: Australian funnel-web spiders

379

Table 28. Male morphological data— Hadronyche raveni (n = 3).

character

range

mean

character

range

mean

character

ratio

SD

CL*

5.82-5.94

5.88

MtlS

19-25

22

CW/CL

0.97

0.015

CW

5.62-5.74

5.69

TalS

10-15

13

CH/CW

0.49

0.012

CH

2.75-2.91

2.81

Ti2S*

2-6

4

CFW/CL

0.77

0.01

CFW

4.47-4.53

4.50

STC2teeth

9

9

CHGW/L

0.15

0.01

ChGL

1.80-1.93

1.87

BulbL

1.94-2.03

1.97

SW/SL

0.78

0.025

ChGW

0.27-0.30

0.28

EmbL

1.22-1.27

1.24

LL/LW

0.83

0.045

ChGCT

3-4

4

BulbW

0.88-0.98

0.92

PLSAPW/L*

0.63

0.038

LL

1.04-1.07

1.05

EmbmidW

0.15-0.16

0.16

Bulb/EmbL*

0.74

0.026

LW

1.22-1.34

1.26

PalpTibL

2.76-2.93

2.85

EmbmidW/L*

0.13

0.006

CUSP*

202-224

213

PalpTibW

1.32-1.38

1.34

PalpTibW/L

0.47

0.012

SL

3.61-3.88

3.75

PalpTibS

0

—

BulbL/TibL*

0.69

0.017

SW

2.92-2.97

2.94

PalpPatS

0

—

PLSAPW

0.40-0.46

0.42

PalpFemS

0

—

PLSAPL

0.64-0.68

0.65

FelS

0

PalS

0

— Chelicerae. Groove narrow, margins diverging distally;
length 1.90, middle width 0.27. Cheliceral teeth: 3 central,
at base of groove; 11 prolateral; 11 retrolateral. — Labium.
Wider than long, not apically indented, shallow groove
behind cuspules. Length 1.04, width 1.24. Labiosternal
sigilla slightly narrowed toward midline. Cuspules c. 202,
number moderate. — Sternum. Ovoid. Length 3.76, width
2.92. Posterior sigilla narrow, elongate. — Palp. Tegular area
wider than long. Middle haematodocha exposed. Embolus
short, shaft of moderate width, weakly curved and tapered.
Distal embolus weakly twisted. Embolus base weakly offset
from tegulum. Bulb length 1.94, width 0.88. Embolus length
1.22, midwidth 0.15. Length of femur 2.76, patella 1.10, tibia
2.76. Width of tibia 1.32. Spination: spines absent. Sinuous
bristles on distal femur. — Legs. 4213 or 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.08

2.13

4.10

3.81

2.38

17.50

2

5.00

2.46

4.26

3.99

2.46

18.17

3

4.25

2.06

3.18

3.52

2.35

15.36

4

5.36

2.22

4.62

4.56

2.55

19.31

Tibia I width 0.80. Femora I—III lacking dorsal spines. Leg
I unmodified, tibial and metatarsal spines scattered. Leg II
without apophysis, tibial spines few, grouped proximally.
Anterior coxal hairs weak, unmodified. Scopulae leg I-IV:
tarsus weak-moderate; metatarsus weak-absent. Tarsi III,
IV weakly swollen, longer than tarsi I, II. Tarsal claw teeth
legs I, II: superior 9, 9; inferior 3, 3. Trichobothria legs I,
II: tarsus 10, 8; metatarsus 7, 6; tibia p4 r6, p5 r6. — Leg
spination. Leg I: femur 0, patella 0, tibia 16, metatarsus 25,
tarsus 14. Leg II: femur 0(1 large bristle), patella 0, tibia 6,
metatarsus 19, tarsus 25. Leg III: femur 0, patella 12 (pll
rl), tibia 8 (p3 r2), metatarsus 20, tarsus 27. Leg IV: femur
6 bristle-like spines, patella 0, tibia 0, metatarsus 16, tarsus
38. — Abdomen. Posterior lateral spinnerets with short apical
segment. Lengths: total 2.21; basal segment 1.00; middle
0.56; apical 0.65. Apical segment width 0.40.

Distribution. Conondale Range, southeastern Queensland
(Fig. 96).

Comments. This species has both the largest labium and
the largest number of labial cuspules in the species group.

Hadronyche lamingtonensis n.sp.

Figures 96-98; Table 29

Etymology. The specific epithet is taken from Lamington
National Park, the type locality.

Types. Holotype male:QMB S394 (QMB). O’Reillys, Lamington National
Park, Queensland, 28°14'S 153°08'E, G.B. and S.R. Monteith, 31 January-
11 April 1976 (in pit fall trap). Paratypes (all QMB). Males: Queensland.
QMB S159, Palm Grove, Tamborine Mtn., 27°55’S 153°10'E, 25 May-13
October 1975, G.B. & S.R. Monteith. New South Wales. QMB S166, Whian
Whian State Forest, via Dunoon, 28°36'S 153°22’E, 26 December 1974-23
March 1975, G.B. & S.R. Monteith.

Diagnosis. CL 6.77-8.04 (male). Small to medium sized
atracine spiders. Differs from H. raveni by having fewer
labial cuspules (44-58) and tibia II more spinose( 15-23)
(Fig. 97F); from H. annachristiae and H. monteithi by the
embolus being relatively short and weakly curved (Fig. 97);
and from H. anzses by male tarsi III, IV unmodified and full
row of promarginal cheliceral teeth.

Male (holotype) — Size. Carapace length 7.86, width 7.75.
Abdomen length 8.38, width 5.96. — Colour. Basic colour
pattern. — Carapace. Slightly longer than wide, cephalic

Figure 96. Collection records for Hadronyche raveni (squares) and
H. lamingtonensis (circles).

380

Records of the Australian Museum (2010) Vol. 62

Figure 97. Hadronyche lamingtonensis, male: (A) spinnerets; (B)
cephalothorax and chelicerae, lateral; (C) palp, prolateral; (D) leg
II, prolateral; (E) cheliceral groove teeth; (F) sternum, labium and
maxilla; (G) cephalothorax and chelicerae, dorsal; ( H ) palpal organ.
Scale lines 1 mm.

area moderately raised. Height 3.49. Frontal width 5.41.
Fovea strongly procurved. Mid-dorsal cephalic setae few,
reaching only halfway to fovea. Anterior strial setae absent.
Anterolateral angle of carapace with a few very weak hairs
only. — Eyes. Sessile. Eye group width 1.97. Median ocular
quadrangle length 0.65, anterior width 0.84, posterior width
1.39. Diameters: AME 0.30, ALE 0.41, PLE 0.37, PME 0.20.
— Chelicerae. Anterodorsalpaturon bristles short. Cheliceral
groove narrow, margins parallel. Groove length 2.55, middle
width 0.40. Cheliceral teeth: 13 central, confined mainly
to basal half of groove (one distal tooth); 12 prolateral;
9 retrolateral. — Labium. Wider than long, not indented
apically. Length 1.28, width 1.41. Labiosternal sigilla entire.
Cuspules c. 58, small in size, number low. — Sternum. Very
broad, subcircular. Length 4.55, width 4.02. Posterior sigilla
large, broad, elongate, anterior margin almost at same level
as anterior margin of middle sigilla. — Palp. Tegular area
wider than long, middle haematodocha exposed. Embolus
gently curved, medium length, shaft hardly tapering distally.
Distal embolus twisted, flange partially reflexedy. Embolus
base weakly offset from tegulum. Bulb length 2.51, width
1.10. Embolus length 1.67, midwidth 0.17. Length of femur
3.32, patella 1.25, tibia 2.99. Width of tibia 1.38. Spination:
femur 3, patella 2, tibia 0. Sinuous bristles on distal femur.
— Legs. 4123.

Figure 98. Hadronyche lamingtonensis, male (S1058): (A) Palp,
prolateral; ( B ) distal patella and tibia I, ventral; (C) metatarsus I,
ventral; (D) distal patella and tibia II, ventral. (E) metatarsus II,
ventral. Scale lines: A,B 0.3 mm and C-E 0.2 mm.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

6.72

2.91

4.77

4.76

2.79

21.95

2

5.82

2.75

4.39

4.22

2.75

19.93

3

5.70

2.48

3.35

4.25

2.88

18.66

4

6.77

2.88

4.98

5.90

3.62

24.15

Gray: Australian funnel-web spiders

381

Table 29. Male morphological data— Hadronyche lamingtonensis (n =

3).

character

range

mean

character

range

mean

character

ratio

SD

CL

6.77-8.04

7.55

MtlS

35-39 (p0-3)

37

CW/CL

0.95

0.036

CW

6.26-7.75

7.20

TalS

29^19

36

CH/CW

0.48

0.03

CH

3.20-3.65

3.44

Ti2S

15-23

19

CFW/CL

0.69

0.01

CFW

4.59-5.62

5.20

STC2teeth

7-8

8

CHGW/L

0.15

0.012

ChGL

2.28-2.68

2.50

BulbL

1.90-2.51

2.28

SW/SL

0.85

0.055

ChGW

0.32-0.40

0.36

EmbL

1.20-1.67

1.47

LL/LW

0.78

0.028

ChGCT

4-14

7

BulbW

0.73-1.10

0.92

PLSAPW/L

0.53

0.027

LL

1.00-1.28

1.16

EmbmidW

0.12-0.17

0.14

BulbW/EmbL* 0.63

0.029

LW

1.32-1.50

1.41

PalpTibL

2.46-2.99

2.75

EmbmidW/L

0.10

0.006

CUSP*

44-58

53

PalpTibW

1.19-1.40

1.32

PalpTibW/L

0.48

0.02

SL

3.94-4.69

4.39

PalpTibS

0

—

BulbL/TibL

0.83

0.051

SW

3.52-4.02

3.74

PalpPatS

0-3

2

PLSAPW

0.33-0.42

0.38

PalpFemS

0-3

2

PLSAPL

0.60-0.78

0.72

FelS

0

PalS

0-5 (pO-2)

3

Tibia I width 1.23. Femora I and II typically without dorsal
spines, but occasionally on femur II. Legs I, II unmodified,
tibial and metatarsal ventral spines scattered. Patellae III, IV
with many prolateral spines. Coxal hairs unmodified. Tarsi
III, IV unmodified. Tarsal scopulae weak to moderately
developed, distal metatarsal scopulae weak to absent. Tarsal
claw teeth legs I, II: superior 10,8; inferior 2,1. Trichobothria
legs I, II: tarsus 7, 6; metatarsus 9, 9; tibia p5 r5, p4 r4. —
Leg spination. Leg I: femur 0, patella 4, tibia 29, metatarsus
36 (p3), tarsus 31. Leg II: femur 0, patella 3 (pi), tibia 23,
metatarsus 51, tarsus 29. Leg III: femur 4, patella 34 (p29
r5), tibia 30 (p8 r5), metatarsus 38, tarsus 34. Leg IV: femur
10, patella 20 (pl4 r6), tibia 22 (p4 r7), metatarsus 29, tarsus
46. Femoral spines mostly bristle-like, a few strong bristles
on legs I, II. — Abdomen. Posterior lateral spinnerets with
short apical segment. Lengths: total 2.84; basal segment
1.40; middle 0.66; apical 0.78. Apical segment width 0.39.

Distribution. Macpherson and Tweed Ranges in eastern
Border Ranges of New South Wales and Queensland (Fig.
96).

Hadronyche annachristiae n.sp.

Figures 99-101; Table 30

Etymology. The species is named for Anna-Christie Gray,
my younger daughter.

Types. Holotype male: AMS KS5438, Kerewong State Forest, nr Lome,
31°35'S 152°33'E, D. Milledge, 1980. Paratypes (AMS). New South
Wales. Males: AMS KS1551, Kerewong State Forest, nr. Taree, 31°35'S
152°33'E, 11 May-19 June 1978, D. Milledge; AMS KS5409, Kerewong
State Forest, 31°35'S 152°33’E, 30 May-1 September 1979, D. Milledge;
AMS KS10770, Kerewong State Forest, Wedding Cake Road, nr. Taree,
31°35'S 152°33E, 28 March 1982, H. Pamaby; AMS KS5637, Lome State
Forest, nr. Taree, 31 °33'S 152°37'E, 1 Sept-9 December 1979, D. Milledge;
AMS KS5420, Lome State Forest, nr. Taree, 31°33’S 152°37’E, 30 May-1
September 1979, D. Milledge.

Diagnosis. CL 6.19-6.66 (male). Small atracine spiders
with (“boat shaped”). Differs from H. raveni and H.
lamingtonensis and by embolus long and curved with base
strongly offset from tegulum (Fig. 100B) and presence
of swollen tarsi III, IV (Fig. 3G); from H. monteithi by

tarsus more spinose (24-43);and from H. anzses by long
promarginal cheliceral tooth row.

Male (holotype) — Size. Carapace length 6.52, width 6.41.
Abdomen length 6.97, width 5.56. — Colour. Basic colour
pattern. — Carapace. Slightly longer than wide, moderately
raised. Height 3.12. Frontal width 4.85. Fovea strongly
procurved. Mid-dorsal cephalic setae almost reach fovea.
Anterior strial setae absent. Anterolateral angle of carapace
with a few weak bristles. — Eyes. Sessile. Eye group width

Figure 99. Hadronyche annachristiae Male. (A) cephalothorax
and chelicerae, lateral; ( B ) palp, prolateral; (C) cephalothorax and
chelicerae, dorsal; (D) cheliceral groove teeth; (E) sternum, labium
and maxilla; ( F) spinnerets; (G) leg II, prolateral. Scale lines 1 mm.

382

Records of the Australian Museum (2010) Vol. 62

1.90. Median ocular quadrangle length 0.68, anterior width
0.85, posterior width 1.36. Diameters: AME 0.26, ALE 0.50,
PLE 0.28, PME 0.25. — Chelicerae. Anterodorsal bristles
on paturon rather short. Cheliceral groove narrow, margins
parallel. Groove length 2.22, middle width 0.34. Cheliceral
teeth: 4 central placed at base of groove (occasionally a few
teeth apically); 10 prolateral; 10 retrolateral. — Labium.
Wider than long with long anterior hairs. Length 1.06, width
1.42. Labiosternal sigilla broad, complete. Cuspules c. 83,
number low. — Sternum. Ovoid. Length 4.08, width 3.21.
Posterior sigilla large, elongate. — Palp. Tegular area much

wider than long, middle haematodocha exposed. Embolus
moderately long, shaft hardly tapering distally, strongly
curved. Distal embolus moderately twisted, flange partially
reflexed. Embolus base strongly offset from tegulum. Bulb
length 2.99, width 1.18. Embolus length 2.18, midwidth
0.16. Length of femur 2.84, patella 1.24, tibia 3.39. Width
of tibia 1.54. Spination: none; sinuous bristles on distal
femur. — Legs. 4123.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

5.79

2.67

4.06

4.22

2.34

19.08

2

5.29

2.67

3.87

3.98

2.50

18.31

3

5.02

2.35

3.25

3.69

3.02

17.33

4

5.93

2.68

4.56

4.96

3.52

21.65

Tibia I width 0.92. Femora I and II without dorsal spines.
Legs I and II unmodified. Scattered ventral spines on
metatarsus I and tibia II, with largest spines grouped
proximally. Tarsi III, IV swollen and longer (boat-shaped)
than tarsi I, II. Coxal hairs unmodified. Tarsal scopulae:

Figure 100. Hadronyche annachristiae, male (A,D-G, AMS KS5049;
B,C, AMS KS5420): (A) palp, prolateral; (B) bulb; (C) embolus,
distal; ( D ) distal patella and tibia I, ventral; (E) metatarsus I, ventral;
(F) distal patella and tibia II, ventral. (G) metatarsus II, ventral. Scale
lines: A 0.2 mm, B 1.0 mm, D-F 0.2 mm and G 0.3 mm.

Gray: Australian funnel-web spiders

383

Table 30. Male morphological data— Hadronyche annachristiae (n

= 6).

character

range

mean

character

range

mean

character

ratio

SD

CL*

6.19-6.66

6.44

MtlS

23-32

27

CW/CL

0.94

0.036

CW

5.85-6.41

6.06

TalS

24-43 (1-4 mv) 35

CH/CW

0.48

0.03

CH

2.87-3.12

2.93

Ti2S

9-16

12

CFW/CL

0.71

0.01

CFW

4.42-1.69

4.59

STC2teeth

8-10

8

CHGW/L

0.13

0.012

ChGL

2.11-2.63

2.26

BulbL

2.72-2.99

2.87

SW/SL

0.80

0.055

ChGW

0.27-0.34

0.29

EmbL

1.97-2.48

2.14

LL/LW

0.79

0.028

ChGCT

4-8

6

BulbW

1.01-1.18

1.07

PLSAPW/L

0.56

0.027

LL

0.96-1.16

1.05

EmbmidW

0.15-0.20

0.17

BulbW/EmbL* 0.50

0.029

LW

1.24-1.42

1.33

PalpTibL

3.01-3.39

3.14

EmbmidW/L*

0.08

0.006

CUSP

83-102

89

PalpTibW

1.39-1.54

1.49

PalpTibW/L

0.47

0.02

SL

3.63-4.08

3.84

PalpTibS

0

—

BulbL/TibL

0.91

0.051

SW

2.92-3.28

3.08

PalpPatS

0

—

PLSAPW

0.36-0.41

0.38

PalpFemS

0-3

1

PLSAPL

0.66-0.72

0.67

FelS

0

PalS*

0

leg I weak; legs II-IV moderately to strongly developed.
Distal metatarsal scopulae absent legs I—III, very weak leg
IV. Tarsal spines numerous, a few small mid-ventral spines
somet im es present. Tarsal claw teeth legs I, II: superior 8, 8;
inferior 1,1. Trichobothria legs I, II: tarsus 7,7; metatarsus
7, 8; tibia p5 r5, p5 r5. — Leg spination. Leg I: femur 0,
patella 0, tibia 12, metatarsus 32, tarsus 42. Leg II: femur 0,
patella 0, tibia 16, metatarsus 35, tarsus 42. Leg III: femur
0, patella 31 (p30 rl), tibia 15 (p5 r3), metatarsus 25, tarsus
47. Leg IV: femur 1, patella p8, tibia 6 (pi), metatarsus 27,
tarsus 88. — Abdomen. Posterior lateral spinnerets with
short apical segment. Lengths; total 2.44; basal segment
1.14; middle 0.58; apical 0.72. Apical segment width 0.38.

Distribution. Known only from the Kerewong and Lome
State Forests near Comboyne, New South Wales (Fig. 101).

Hadronyche monteithi n.sp.

Figures 101-102; Table 31

Etymology. The species is named for Dr Geoff Monteith,
a collector of the type specimens and many other spider
species.

Types. Holotype male: QMB SI81 (QMB). The Head, via Killarney,
Queensland, 28°20'S 152°18'E, G.B. and S.R. Monteith, 31 Mar-2 August
1975.

Other material. Queensland. Male: QMB S148, Bald Mountain, via
Emuvale, 28°14’S 152°25E, G.B. and S.R. Monteith, 28 December 1974-30
March 1975.

Diagnosis. CL 5.00-5.19 (male). Small atracine spiders.
Differs from H. raveni and H. lamingtonensis by having
slender, curved embolus (Fig. 102D and tarsi III, IV swollen;
from H. annachristiae by relatively few tarsus I spines (5-7);
and from H. anzses by strongly curved embolus.

Male (holotype) — Size. Carapace length 5.00, width 4.70.
Abdomen length 5.00 width 3.94. — Colour. Basic colour
pattern. Small paler area on anterodorsal abdomen flanked

Figure 101. Collection records for Hadronyche annachristiae
(squares) and H. monteithi (circles).

by a pair of sigilla. — Carapace. Slightly longer than wide,
narrowing rapidly posteriorly, moderately raised. Height
2.54. Frontal width 3.57. Fovea strongly procurved. Mid¬
dorsal cephalic setae do not reach fovea. Anterior strial setae
few or absent. Anterolateral angle of carapace with a few
hairs only. — Eyes. Sessile. Eye group width 1.39. Median
ocular quadrangle length 0.49, anterior width 0.68, posterior
width 0.93. Diameters: AME 0.23, ALE 0.32, PLE 0.27,
PME 0.18. — Chelicerae. Sparse-moderate anterodorsal
cover of short bristles on paturon. Cheliceral groove very
narrow, margins subparallel. Groove length 1.68, middle
width 0.25. Cheliceral teeth: 2 central, in proximal end of
groove; 6 prolateral; 9 retrolateral, absent from distal third
of retromargin. — Labium. Wider than long. Length 0.78,
width 1.05. Cuspules c. 68, number low. Labiosternal sigilla
entire. — Sternum. Broad, ovoid. Length 2.99, width 2.58.
Posterior sigilla large, ovoid. — Palp. Tegular area much
wider than long, middle haematodocha exposed. Embolus
shaft strongly curved, moderately long and slender but hardly
tapered distally. Distal embolus twisted and set at obtuse
angle to shaft. Embolus base weakly offset from tegulum.
Bulb length 1.96, width 0.79. Embolus length 1.44, midwidth
0.10. Length of femur 2.20, patella 0.90, tibia 2.18. Width of
tibia 1.07. Spination: femur 4 (bristle-like). — Legs. 4123.

384

Records of the Australian Museum (2010) Vol. 62

Table 31. Male morphological data— Hadronyche monteithi (n = 2).

character

range

mean

character

range

mean

character

ratio

CL*

5.00-5.19

5.09

MtlS

16-19

18

CW/CL

0.95

CW

4.70-5.02

4.86

TalS*

5-7

6

CH/CW

0.51

CH

2.44-2.54

2.49

Ti2S*

3-7

5

CFW/CL

0.72

CFW

3.57-3.81

3.69

STC2teeth

6-11

9

CHGW/L

0.15

ChGL

1.68-1.86

1.77

BulbL

1.96-1.97

1.97

SW/SL*

0.85

ChGW

0.25-0.30

0.27

EmbL

1.42-1.44

1.43

LL/LW

0.74

ChGCT*

3

3

BulbW

0.79-0.80

0.80

PLSAPW/L

0.60

LL

0.78-0.84

0.81

EmbmidW

0.10-0.11

0.11

BulbW/EmbL 0.55

LW

1.05-1.13

1.09

PalpTibL

2.18-2.40

2.29

EmbmidW/L* 0.07

CUSP

68-81

75

PalpTibW

1.07-1.54

1.30

PalpTibW/L*

0.56

SL

2.99-3.26

3.12

PalpTibS

0

—

BulbL/TibL

0.86

SW

2.58-2.72

2.65

PalpPatS

0

—

PLSAPW

0.32-0.33

0.33

PalpFemS

2-4

3

PLSAPL

0.44-0.68

0.56

FelS

0

PalS

0

A

Figure 102. Hadronyche monteithi, male: (A) spinnerets; ( B)
cephalothorax and chelicerae, lateral; (C) cheliceral groove teeth;
( D ) palpal organ; (E) leg II, prolateral; (F) palp, prolateral; (G)
cephalothorax and chelicerae, dorsal; ( H) sternum, labium and
maxilla. Scale lines 1 mm.

Leg

Femur

Patella

Tibia

Metatarsus Tarsus

Total

1

3.92

1.90

2.87

2.82

1.86

13.37

2

3.48

1.62

2.73

2.64

1.86

12.33

3

3.32

1.60

1.92

2.52

2.12

11.48

4

3.92

1.76

2.76

3.30

2.62

14.36

Tibia I width 0.74. Femora I and II without spines. Legs I and
II unmodified, tibial spines few and metatarsal ventral spines
scattered. Coxal hairs unmodified. Tarsi III, IV swollen,
“boat-shaped”. Scopulae absent on distal metatarsi, weak to
moderately developed on tarsi I-IV. Tarsal claw teeth legs I,
II: superior 7, 6; inferior 0, 0. Trichobothria legs I, II: tarsus
5, 6: metatarsus 8,6; tibia p4 r4, p4 r4. —Leg spination. Leg
I: femur 0, patella 0, tibia 6, metatarsus 16, tarsus 7. Leg II:
femur 0, patella 0, tibia 2-3 (ventral spines ± weak). Leg
III. femur 1, patella 27 (p26 rl), tibia 11 (p6 r2) metatarsus
18 (pd6 rd4), tarsus 8. Leg IV: femur 1, patella p3, tibia 6
(p3 dl), metatarsus 16, tarsus 13. — Abdomen. Posterior
lateral spinnerets with short apical segment. Lengths: total
1.76; basal segment 0.88, middle 0.44, apical 0.44. Apical
segment width 0.32.

Distribution. Border Ranges in Killarney region, south¬
eastern Queensland (Fig. 101).

Hadronyche anzses Raven

Hadronyche anzses Raven, 2000: 225

Types. Holotype male: QMB S18825. Mossman Bluff summit, lOKm west
of Mosman, 16°26'54"S 145°16'59"E, N.E. Queensland, 20 December
1989-15 January 1990, G. Monteith, G. Thompson and ANZSES
Expedition.

Diagnosis. CL 5.31 (male). Small atracine spiders. Differs
from other lamingtonensis group species in having only
a few basal teeth (3) on the cheliceral promargin (Raven,
2000, Fig. 1G) and metatarsus I with fewer spines (c. 10);
and from H. raveni and H. lamingtonensis by tarsi III, IV
swollen, “boat-shaped”.

Distribution. Known only from the type locality.

Gray: Australian funnel-web spiders

385

adelaidensis species group

Description (Figs. 103-104; Tables 32-33). Small atracinae
(CL 5.4-6.9). Burrow entrance without triplines, with side-
chamber closed by a trap-door. Carapace broad, strongly
raised. Labium and sternum wide. Serrula absent. Cheliceral
groove narrow with long central tooth row. STC teeth few
(8-9). Male femora I, II with dorsal spines or bristles. Males
without leg II apophyses; leg I modified (incrassate tibia and
metatarsus) or unmodified. Male palpal tibia bulbous basally;
patella wider than femur.

Included species. Hadronyche adelaidensis (Gray), H.
flindersi (Gray), H. eyrei (Gray).

Distribution. Limited to the Gulf Ranges region of South
Australia.

Comments. Burrows of adult and subadult spiders have a
side chamber with a robust soil door and a simple collapsible,
collar-like silk entrance opening in or under lear litter in open
forest and woodland habitats (Main, 1976; Gray, 1984). The
burrows lack surface triplines. Interestingly, juveniles of
H. adelaidensis make burrows that lack side chambers but
have well-formed, flap-like surface trapdoors (Gray, 1984).

These spiders are separated from the main eastern
distribution of the Atracinae by the dry shrublands/
woodlands of the southern Murray Basin (Fig. 2). They may
originally have been isolated as a western atracine population
during a Tertiary marine incursion into the Murray Basin.

Hadronyche flindersi (Gray)

Figure 105. Collection records for Atraxflindersi (open circles), A.
adelaidensis (black squares) and A. eyrei (closed circles).

Types. Holotype female: AMS KS983, Mt. Remarkable (lower slopes),
3km north of Melrose, Flinders Range, South Australia, 32°47'S 138°04'E,
23 April 1973, M. and G. Gray. Paratype male: AMS KS980, same data as
holotype, 25 April 1973.

Diagnosis. CL 5.4-6.9 (male and female). Small atracine
spiders Males differ from H. adelaidensis by row of strong
dorsal spines (8-9) on femur I; and tibia I, II with 6-9
spines. Male palp with straight, slender embolus, and 12-16
spines on patella. Male tarsi III, IV weakly swollen. Females
with moderately slender spermathecae with a subapical
constriction; few prolateral spines on tibia III (8-9).

Figures 103-105; Table 3 2 Distribution. The southern end of the Flinders Ranges, South

Australia (Fig. 105).

Atrax flindersi Gray, 1984: 446.

Hadronyche flindersi. -Gray, 1988: 114. Comments. Female burrow with juvenile spiders in side

passage closed by trapdoor in April, 1973.

Figure 103 .Hadronyche flindersi, male: (A) cheliceral groove teeth;
( B ) cephalothorax and chelicerae, lateral; (C) leg I, prolateral; (D)
sternum, labium and maxilla; ( E ) cephalothorax and chelicerae,
dorsal; ( F) palp, prolateral; (G) spinnerets. Scale lines 1 mm.

Figure 104. Hadronyche flindersi, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) spinnerets;
( D ) cheliceral groove teeth; (E) cephalothorax and chelicerae,
dorsal; ( F ) leg III, prolateral, patella, tibia, metatarsus; (G)
spermathecae. Scale lines 1 mm.

386

Records of the Australian Museum (2010) Vol. 62

Table 32. Male morphological data —Hadronyche flindersi (n = 2).

character

range

mean

character

range

mean

character

ratio

CL*

5.44-5.86

5.65

MtlS

15-18

17

CW/CL

0.98

CW

5.33-5.72

5.52

TalS

18-21

20

CH/CW

0.45

CH

2.38-2.62

2.50

Ti2S

9

9

CFW/CL

0.70

CFW

3.81-4.10

3.95

STC2teeth

8-9

9

CHGW/L

0.13

ChGL

1.74-1.86

1.80

BulbL

2.20-2.36

2.28

SW/SL

0.90

ChGW

0.24-0.25

0.25

EmbL

1.28-1.42

1.35

LL/LW

0.72

ChGCT

16-18

17

BulbW

1.02-1.04

1.03

PLSAPW/L

0.35

LL

0.90-1.00

0.95

EmbmidW

0.10-0.12

0.11

BulbW/EmbL 0.76

LW

1.31

1.31

PalpTibL

2.81-2.95

2.88

EmbmidW/L* 0.08

CUSP

171-185

178

PalpTibW

1.43-1.54

1.48

PalpTibW/L

0.51

SL

3.16-3.47

3.31

PalpTibS

2-3

3

BulbL/TibL

0.79

SW

2.92-3.10

3.01

PalpPatS*

14-15

15

PLSAPW

0.36-0.38

0.37

PalpFemS

2-7

5

PLSAPL

1.03-1.08

1.05

FelS*

8-9

9

PalS

0

Hadronyche adelaidensis (Gray)

Figures 105-107; Table 33

Atrax adelaidensis. Gray, 1984: 442.

Hadronyche adelaidensis. -Gray, 1988: 114.

Types. Holotype female: N1979146 (SAM). Hackney, Adelaide, South
Australia, 34°54'S 138°37'E, 16 November 1973, J. Batt. Paratype male:
N1979145 (SAM). St. Peters C.G.S. Adelaide, South Australia, 3 June
1971, D. Edwards.

Diagnosis. CL 5.5-6.8 (male and female). Small atracine
spiders. Males differ from H. flindersi by presence of
incrassate tibia I and metatarsus I, reduced spination on

Figure 106. Hadronyche adelaidensis, male: (A) sternum, labium
and maxilla; ( B ) cephalothorax and chelicerae, lateral; (C)
spinnerets; ( D ) cheliceral groove teeth; (E) cephalothorax and
chelicerae, dorsal; (F) leg I, prolateral; (G) palp, prolateral. Scale
lines 1 mm.

tibia I, II (0-2), and absence of dorsal spines (bristles only)
on femur I. Male palp with short, broad embolus. Females
are distinguished by strongly dilated spermathecae; and
numerous prolateral spines on patella III (18-26).

Distribution. Mount Lofty Ranges and the Adelaide region,
South Australia (Fig. 105).

Comments. Williams & Goode (1978) found evidence that
the Miocene Murray River in South Australia may have run
west into the Spencer Gulf south of Port Pirie, before its
flow was diverted southward near Morgan. The progenitor
populations of H. adelaidensis and H. flindersi may have
speciated on either side of a barrier formed by the ancient
Murray River in South Australia.

Figure 107. Hadronyche adelaidensis, female: (A) sternum,
labium and maxilla; ( B ) cephalothorax and chelicerae, lateral;
(C) spinnerets; (D) cheliceral groove teeth; (£) cephalothorax and
chelicerae, dorsal; (F) spermathecae; (G) leg III, prolateral, patella,
tibia, metatarsus. Scale lines 1 mm.

Gray: Australian funnel-web spiders

387

Table 33. Male morphological data —Hadronyche adelaidensis (n =

3).

character

range

mean

character

range

mean

character

ratio

SD

CL*

5.55-6.40

6.04

MtlS

19-31

25

CW/CL

0.93

0.064

CW

5.30-6.15

5.63

TalS

12-20

16

CH/CW

0.47

0.046

CH

2.57-2.75

2.65

Ti2S*

1-2

1

CFW/CL

0.74

0.017

CFW

4.21-4.67

4.45

STC2teeth

8

8

CHGW/L

0.10

0.006

ChGL

2.07-2.39

2.22

BulbL

2.10-2.26

2.18

SW/SL

0.89

0.02

ChGW

0.22-0.24

0.22

EmbL

1.24-1.36

1.30

LL/LW

0.76

0.021

ChGCT

10-12

11

BulbW

1.02-1.06

1.04

PLSAPW/L

0.40

0.015

LL

1.00-1.06

1.03

EmbmidW

0.22-0.24

0.23

BulbW/EmbL 0.80

0.05

LW

1.30-1.44

1.37

PalpTibL

2.95-3.14

3.03

EmbmidW/L* 0.18

0.006

CUSP

181-230

207

PalpTibW

1.48-1.60

1.54

PalpTibW/L

0.51

0.006

SL

3.45-3.73

3.59

PalpTibS

0

—

BulbL/TibL

0.72

0.01

SW

3.14-3.26

3.20

PalpPatS

0

—

PLSAPW

0.36-0.38

0.37

PalpFemS

0-1

1

PLSAPL

0.92-0.94

0.93

FelS

0*

PalS

0

Figure 108. Hadronyche eyrei, female: (A) sternum, labium and
maxilla; ( B ) cephalothorax and chelicerae, lateral; (C) cheliceral
groove teeth; ( D ) spinnerets; (E) cephalothorax and chelicerae,
dorsal; (F) spermathecae; (G) leg III, prolateral, patella, tibia,
metatarsus. Scale lines 1 mm.

Hadronyche eyrei (Gray)

Figures 105, 108

Atrax eyrei. Gray, 1984: 444.

Hadronyche eyrei. -Gray, 1988: 114.

Types. Holotype female: AMS KS4509, 6.5km south of Coulta, Eyre
Peninsula, South Australia, 34°23'S 135°28'E, 18 December 1952, B.Y.
Main.

Diagnosis. CL 6.6 (female). Small atracine spiders. Differs
from other adelaidensis group species by its rather uniform
maroon-brown abdominal pigmentation (much lighter
laterally and ventrally in other species). Prolateral patella
with 5-10 spines. Spermathecae unmodified, sac-like.

Distribution. Southern Eyre Peninsula, South Australia
(Fig. 105).

Comments. This species represents the known western
extent of the atracine spiders in Australia.

Acknowledgments. The late Vera Levitt-Gregg was a Sydney
naturalist and Australian Museum Associate, whose knowledge and
enthusiasm initiated my interest in these highly venomous spiders.
My thanks to Dr Robert Raven for his useful comments on the
manuscript. Curatorial staff at the Queensland Museum, Museum
Victoria, the Tasmanian Museum and Art Gallery, the Launceston
Museum and Art Gallery and the South Australian Museum
assisted with the loan of material. Field work was supported by a
grant from the Australian Biological Research Study—thanks to
Harry Parnaby for his field assistance. Funding from the Australian
Museum assisted with mapping and SEM editing work by Vicky
Tzsoumiz. Sue Lindsay took the SEM pictures and Kate Attwood
kindly assisted with re-arrangement of drawings. Thanks to my
wife, Greta Jensen Gray, who assisted in so many ways, including
data compilation, field work and coffee.

Table 34. Female morphological data for Atrax, Illawarra and Hadronyche species; means within parentheses, ratios within square brackets, measurements in mm; n = 6 for
each species except//, emmalizae and H. lynabrae n = 3. Abbreviations: CL , carapace length; CW, carapace width; CH, carapace height; SL, sternum length; SW, sternum width;
LL, labium length; LW, labium width; ChGL, cheliceral groove length; ChGW, cheliceral groove width; PLSAPL, posterior lateral spinneret apical segment length; PLSAPW,
posterior lateral spinneret apical segment width; TilS, tibia I spine number; Pa3proS, patella III prolateral dorsal spine number.

388

Records of the Australian Museum (2010) Vol. 62

cq

t-H

os

'xl-

00

OO

O'

H

Os

p

p

CO

OG

O

os

O'

O'

&

00

7

sd

CN

o4

04

o

H

o

CN

P

CO

04

o

O

04

o

o

o

os

p

CO

p

'xf

04

p

O

p

00

H

os

>n

d

in

04

04

cd

H

—H

o

<44

Cj

1

O'

4

4

1

1

m

1

in

1

O'

44

1

CO

I

SO

so

N-

p

°p

p

p

P

i—H

in

os

•n

sd

sd

cd

cd

i-H

i-H

04

o

o

o

IO4^H00^tS0O4O4^t0S

;qoqqpo4qoqpp

'7 so in 7 7 7 d 7 d

invo^tooooio^tit

loo^M^ooiooinc^

inoointNtNcn
I I I I I I
S0O40S^H00^j-_ .._

oosso^-*oinsoosso

- u -,,- n -,,- H -,

I I I so in Tt O' oo os

\TIT, -tnoiOOOO

54,

O O O O O O

^04 7 7 no in no O
, - omti h o m in

oin-H'xi-in— iO'SOO'Sooo
p p p oo cn p p p cn oo p

oo 6 cn Tt h h d o o o

^■OrnOHinin'OiO'too

p p p p p p p p p p in

os oo’ 7 7 7 7 — 7 o —’ o 7_,,,_, ( ____..._ r .__

I I I I I I I I I I I .^^770 os 7 so

oOfN^ONOoO'OOM^tm o oo ^ rH IO

osr-ooo4i^o4ini^co^tcowp:^;^7X.7

7 sd 7 7 7 7 7 7 o o o o co ££££££

CO

^_

_

,_ N

,_

^_

,_

^_

^_

00

^_

,_

,_ s

_

^_

,_ s

,_

,_

p

o

i—i

7"

in

00

m

O

os

04

•n

^H

so

O'

t-H

o

in

'xf

7

04

CO

o

00

cq

o

p

cn

00

p

CO

P

•n

os

p

p

o

p

p

p

O'

p

so

o^

cd

in

cd

d

d

CN

o

o

00

7

sd

7

qi

cd

d

o

o

'—^

—

■—

in

04

o

04

os

o

oo

so

O'

m

CO

p

m

o

o

so

7—1

co

04

04

os

d

p

CO

p

p

04

04

'xl-

cn

so

SJ

o

CO

p

OS

p

p

of

p

Os

P

p

3

7

04

00

4

7

in

sd

1

in

7

of

7

O'

oi

1

O'

cd

1

O'

2

oo

04

1

os

2

<7

rn

7

CO

00

F^ 1

pj-

7

p

in

p

7

7

p

a

7

O'

os

1

y—H

in

1

oo

sd

4

>n

4

04

1

in

04

1

04

cd

1

os

o

I

04

7

CO

7

04

P

04

p

O

04

p

p

04

p

m

V ’ J

1

7

00

O

p

p

p

p

P

m

in

os

^1-

o :

sd

04

P

cd

o

t-H

o

(N

CO

o,

o

o

o

o

o

d

rd

o^

cd

7

cd

i-H

04

o

o

o

s

cn

7

PP

7

7

7

7

CO

7

&PJ

CO

' .

7

oo

so

7

7

7

in

7

£

p

p

cn

oo

P

p

oo

p

'xf

p

in

w

j

p

t—H

04

p

p

04

00

o

O'

SS

00

7

7

7

7

7

7

7

o

7

o

~5

'xl-

w

7

oo

sd

7

7

7

o

7

d

'q

v — '

v —^

v — y

n '

s —^

«

00

H

s y

s — y

00

04

so

CO

'xf

co

o

00

04

00

so

l>

m

O'

oo

00

m

m

so

m

IS;

i—H

p

oo

p

CO

p

o

so

p

p

XI

CN

’—1

00

o

p

00

o

o-

p

as

4

00

7

4

7

l

os

i

7

00

7

4

7

l

CO

o

I

m

7

C"

7

m

7

cn

1/7

'Os'

vn

pj-

°o

7

p

7

oo

cn

7

in

p

4

sd

1

o

os so

44

7

l

o-

7

I

00

7

^h 04

44

7

o-

p

p

p

p

p

co

o-

u i

1

H i

1

H

p

m

p

o

o

p

so

p

so

7

7

sd

7

7

7

7

7

7

d

o

o

<N

O

o

O

o

o

o

Ed

ON

7

sd

7

7

7

7

O

7

o

in

o

00

P-x

p

rn

,— N

,— s

,— N

,— s

^— s

,— s

<N

00

s — N

,— s

,— N

,— s

,— s

rXl

00

7

m

in

04

7

o

O'

p

in

7

CO

o

7

so

o

V. N
t-H

77

p

os

p

P

p

so

P

00

w

H

qs

p

o

OO

p

p

p

Os

oo

os

7

sd

7

7

7

7

d

7

d

7

7

7

7

7

7

o

7

o

On

On

'—^

s —^

.N

m

o

'— /

s —'

s —^

oo

On

o

o

o

os

CO

m

so

so

so

CN

os

04

04

m

04

m

04

O'

7

p

04

CO

p

o

'xl-

so

04

p

^4

o

so

p

in

•n

SO

so

p

p

H

4

4

7

4

OO

4

sd

7

7

4

T

CO

d

I

04

7

4

cn

rr\

7

'7

00

r7

pj-

7

p

7

oo

NO

H

i-H

cn

4

4

sd

1

O'

7

l

O'

sd

1

CO

04 04 CO

444

7

OO

7

I

O'

4

so

p

p

p

p

oo

^r

so

s’ t

1

v’ i

1

Ed

p

p

p

00

p

p

q

so

so

os

7

7

7

7

7

CN

7

o

7

o

(N

(N

o

O,

o

o

O,

o

oo

7

7

7

7

7

CN

7

o

7

o

> 7 > 7 > c /3 on

p

7R: S zt ; - i o,

Si

a!

J

! PU m w n 5 S
P1> ffi > H O Pj

7

ad

hOinninmonoiHON
cn oo in t"- in cn m o omt;

hOcn^tcn-

< '-H (N O O O

lOrHfn'onrHvocoONoocn

oo 7
I I

(NON .

"nT CN SO

't it, m — (N N O — O -ia _.._ hH _i,_ H _ i

I I I I I I I I I

on oo o on ^ n O', cn r , ^ ^ ^ ^ co no
so- ON-OsosoinpP^X'.'.X X

sd sd 7 7 ri 2 2 d o o o - co £££££ £

24

O' r-

7 q
4°

in

o£

00

ososinoosooor^r-'
p — in in in p p p p
7 7 m] cn 7 7 d 7 ©

r^soooscNsomi^cN
cnoopphinpori
7 7 m] 7 oi 7 d oi — 7 7_,,_ H _ h _ a _,

l l l l l l l l l

co O' cn co os in in os w-, qq in O' o cn in
— (N o (N (N go in | | vj l) • V- X

7 d >n cn cn co o — O co in £££;7£ £

r- co oj r- vo oo oo oo — co co

(N O p - h OO ITi IT, O O

oo 7 7 7 7 — — cn d — d

M CO O co N h- ON (N Cl OO

p p p p oo cq cq p p p p

OS 00 7 7 7 CN cn 7 d i —i d 77

vi-Ud>4tJ.<i<±<4 ' '

os o oo in ^ so <n \i-o m i-

sd sd 7 7 7 7 7 7 o 7 o o cn £££££ £

- - Cl - Cl Ti o O
^ oi mj os in O' os oi Tt

rHNOOOrHO ITj

■ rH os in o os cn ^
i ddddod

o .
o <

O O O -H o o

in O' cn O' oo o
q \ os »n oo os cn in

ddddod

U r N rk OI

7 > 7 > 7 7

U S3^ s^SsdSS

^ U oo

7
Oh

°d

species H. nimoola H. modesta H. meridiana H. infensa H. orana H. valida

CL 5.88-9.24 (8.00) 6.68-8.12 (7.29) 5.90-7.86 (6.84) 9.56-12.97 (11.18) 7.93-10.74 (9.16) 7.96-12.05 (9.89)

CW 5.76-8.65 (7.78) 5.97-8.19 (6.95) 6.29-7.39 (6.77) 9.42-10.73 (10.05) 8.15-9.17 (8.76) 7.19-10.40 (8.69)

CH 3.08-5.44(4.76) 3.16-4.43 (3.72) 3.55-4.43 (3.97) 4.76-6.42 (5.49) 3.85-5.37 (4.82) 3.84-6.18 (4.80)

SL 3.80-5.78 (4.99) 4.22-5.65 (4.95) 4.21-5.21 (4.77) 5.44-7.58 (6.69) 5.48-6.71 (6.31) 5.21-7.68 (6.42)

Gray: Australian funnel-web spiders

389

o

p

N-

m

Qi

3

e2

VO On ^ ^ IT)

OS 00 oo IN SO SO

rd O-£o

so oo os in os
N cn in cn oo cn so

3333333 !

Tf oi o o o) i 222.SS 2,

^oo in so in so o

cn cn oo so so n in

HOOOOOOlOh

7 £ £22 £ 3 £

Tt cn cn so cn <N
in cn (N O O oo oo
in cn cn cn 4
I I I i I . .

oo — r'-cNcNsoN-
so oo oo in N cn so

<(NO'

O (N in

£7

O O co

so in oo o oo
os in oo os cn Tj-

d> <5 d> d> d> d>

cn cn ■t os os ■t cn
(NcNppr^r^r-;
n rd rd rn p-£o

m os os o O (N os
os in oo os rn os

in cn <N cn © <N © ,

l £ J. J _L J, ww O in oo in cn (N

.• • • • • I o o o o o o

Tj- cn (N cn o '

OO — N sO — (N — 1
r- in p in in p in
cn 4 4 <p o o o

so c-~ ^ cn tj- tj- oo
p p — p in — n

4 — CN CN d —- O p \d_____ _

I I I I I I I £^££573o n
rnnHcidddol ££££££

cn

IN

in

<N

so

CN

o

>3

cn

os

os

IN

cn

•n

os

s—H

p

p

p

SO

»n

os

in

K

l—J

ps

p

CN

o

CN

p

IN

p

00

p

4

t-H

CN

o

o

o

p

sd

>n

c4

CN

cn

d

s-H

O

s —'

Q

00

»n

,s —^

■^t

o

o

H

oo

so

SO

p—\

s

p

cn

os

oo

so

•n

os

o

o

so

CN

so

N

o

so

p

»n

H

N"

o

p

p

so

p

00

cn

O

7

»n

r—H

1

N-

7

os

cn

1

O

3

00

7

OS

3

^os

In

p

cn

in

cn

p

57

IN

i-H

CN

4 1

in

U

a

S

7

so

7

00

7

N

3

7

cn

7

IN

CN

1

os

cn

1

ST

3

so

7

N"

s-H

1

p

CN

•n

i-H

cn

so

cn

CO

1 |

p

CN

p

i—H

CN

•n

N

N

so

p

m

cn

CN

o

o

o

o "T

o

o

o

o

o

o

ad

N

IN

cn

in

CN

o

o

so so cn os so
^ so so ^ so o so

2L33££3£
oo o so cn os os cn ^

O — N" OO t"-- CN Os ^CN

in (N (N CN I

L F7 4 1 57 57 oo cn
I os so oo os (N so

. , ,'3 ,

'T os so IN (N

cn^H(Nin^i-so^i-'|'_i.T. / ^isT.\--
^dHHoddoH£££££ £

!>4>;4>;4>G0G04>;4>;4 4

P y < 7 H Ohn: p > 7 l> >

o S2

4

C

GO

J

4

5J

so

■T

,_ N

,_ N

^_ N

,_ N

,_ s

^_ N

n

cn

•n

00

so

CN

00

00

O

00

N

os

£

*G

CN

so

p

p

N

IN

(N

so

Oo

p

sd

'T

—

—

CN

o

s-H

o

so

00

s —^

'— /

N —'

S- /

s — /

^^

v —^

CN

cn

CN

'T

os

so

so

•n

m

so

p

d

00

o

IN

CN

cn

p

00

IN

7

00

7

7

7

cn

o

s-H

9

p

£

!_,

,_,

,_,

,_,

,_,

1

CN

o

p

i

CN

p

o

cn

o

p

p

in

p

SO

so

■T

1

s—H

os

in

7

00

1

N

00

in

in

'T

N

N

os

in

r i

CN

in

tti

IN

•n

cn

'T

cn

s-H

i-H

CN

o

s—H

o

s—H

o

o

o

o

o

o

oocNoo^tNCNOooso

p p — — p p p p p

4 so in cn r i cn © 4 ©

o '
oo ■

o 1 ^ 1

-h in i

4 A

(N p l
oo cn 1

IN

•n

o

so

CN

cn

O

p

•n

os

p

os

»n

os

in

CN

CN

'T

o

CN

9 CN

;£

!_!

!_!

_!

_!

_!

!_!

1

s-H

46

SO N

CN

N

i

cn

so

os

'T

os 7
in P

N

in

00

CN

in

IN

N

■T

os

'T

CN

00

cn

'T

CN

o

o o

4 "

o

o

o

o

o

o

r-> ,
os -

^ i-H inoorHrncN cn
;cn^noO'xl-oooscN

i odd odd.

in so cn cn

cn

cn

o os os

■T

■T

os

»n os

p

00

so

p

SO

oo oo T in

p

p

cn cn o

d

x '

^— /

^^ ^^

s — /

>- o

s —^

^^

in

_

'T O

so

m

cn

00

cn

•n

00

p

p

in cn

m

p

ri

p

IN

00

so

7

00

1

333

7

7

cn

1

3

7

3

(N

o

IN 00

CN

cn

N"

'T

SO

cn

p

p

p p

cn

N

P

O

SO

p

so

00

IN

■T in

4

4

4

cn

o

4

o

(rnininhcn cn
os in in oo cn "T
! o o o o o d

4£:ffi4>:4>:4>:4>;cnp4>:4>;44

uGu I/ ’«- | 3on^&-?syB52Sss

U S3«

&3

GO

4

4

390

Records of the Australian Museum (2010) Vol. 62

References

Ausserer, A., 1871. Beitrage zur Kenntniss der Arachniden-Familie
der Territelariae Thorell—(Mygalidae). Verhhandlungen
zoologisch-botanischen Gesellschaft in Wien 21: 117-224.

Ausserer, A., 1875. Zweiter Beitrag zur Kenntniss der Arachniden-
Familie der Territelariae Thorell (Mygalidae). Verhhandlungen
zoologisch-botanischen Gesellschaft in Wien 25: 125-206.

Beavis, A.S., & D.M. Rowell, 2006. Phylogeography of two
Australian funnel-web spiders (Araneae: Mygalomorphae:
Hexathelidae) in Tallaganda State Forest, New South Wales.
In Insect Biodiversity and Dead Wood: Proceedings of a
Symposium for the 22nd International Congress of Entomology.
General Technical Report SRS-93, ed. Simon J. Grove and
James L. Hanula. Asheville, NC: US Department of Agriculture
Forest Service, Southern Research Station. 1 lOp.

Bonnet, P., 1955. Bibliographia araneorum. Toulouse, 2(1): 1-918.

Bonnet, P., 1957. Bibliographia araneorum. Toulouse, 2(3):
1927-3026.

Bonnet, P., 1958. Bibliographia araneorum. Toulouse, 2(4):
3027^4230.

Bradley, R.A., 1993. Seasonal activity patterns in Sydney funnel-
web spiders, Atrax spp. (Araneae: Hexathelidae). Bulletin of the
British Arachnological Society 9(6): 189-197.

Brignoli, P.M., 1983. A Catalogue of the Araneae Described
Between 1940 and 1981. Manchester University Press, 755 pp.

Biicherl, W., 1971. Spiders. In Venomous Animals and their Venoms.
Academic Press, New York, 3: 197-277.

Cambridge, O.P-, 1877. On some new genera and species of
Araneidea. Annals of the Magazine of natural History (4)19:
26-39.

Cooley, A., 1989. An apparent case of character displacement in
sympatric populations of atracine spiders of the genera Atrax
and Hadronyche. Honours thesis, Australian National University.

Coyle, F., 1986. Courtship, mating and the function of male-
specific leg structures in the mygalomorph spider genus Euagrus
(Araneae, Dipluridae). Proceedings of the IX International
Congress of Arachnology, Panama. Pp. 33-38.

Goloboff, P.A., 1993. A reanalysis of mygalomorph spider families
(Araneae). American Museum Novitates 3056: 1-32.

Gray, M.R., 1978. Venoms of Dipluridae, A. Systematics and
Distribution. In Arthropod Venoms, ed. S. Bettini, pp. 121-126.
Berlin: Springer-Verlag.

Gray, M.R., 1981. Getting to know funnel-web spiders. Australian
Natural History 20: 265-270.

Gray, M.R., 1984. The taxonomy of the Atrax adelaidensis
species-group (Macrothelinae: Mygalomorphae) with notes on
burrowing behaviour. Records of the South Ausralian Museum
18; 441-452.

Gray, M.R., 1986. A systematic study of the funnel-web spiders
(Mygalomorphae: Hexathelidae: Atracinae). PhD thesis.
Macquarie University.

Gray, M.R., 1987. Distribution of the funnel-web spiders. In Toxic
Plants and Animals: A Guide for Australia, ed. J. Covacevich, P.
Davie & J. Peam, pp. 312-321. Brisbane: Queensland Museum.

Gray, M.R., 1988. Aspects of the systematics of the Australian
funnel-web spiders (Araneae: Hexathelidae: Atracinae) based
upon morphological and electrophoretic data. Australian
entomological Society miscellaneous Publication 5: 113-125.

Hedin, M., & J.E. Bond, 2006. Molecular phylogenetics of the
spider infraorder Mygalomorphae using nuclear rRNA genes
(18S and 28S): Conflict and agreement with the current system
of classification. Molecular Phylogenetics and Evolution 41:
454-471.

https://dx.doi.org/10.1016/j.ympev.2006.05.017

Hickman, V.V., 1927. Studies in Tasmanian spiders. Part I. Papers
and Proceedings of the Royal Society of Tasmania 1926: 52-86.

Hickman, V.V., 1964. On Atrax infensus sp. n. (Araneida:
Dipluridae) its habits and a method of trapping the males. Papers
and Proceedings of the Royal Society of Tasmania 98: 107-112.
Hickman, V.V., 1967. Some Common Spiders of Tasmania.

Tasmanian Museum and Art Gallery, 112 pp.

Hogg, H.R., 1901. On Australian and New Zealand spiders of the
suborder Mygalomorphae. Proceedings of the zoological Society
of London 1901: 218-279.

Hogg, H.R., 1902. On some additions to the Australian spiders of
the suborder Mygalomorphae. Proceedings of the zoological
Society of London 1902: 121-142.

Karsch, F., 1878. Exotisch-araneologisches. Zeitschrift fur die
gesamten Naturwissenschaften 51: 332-333.

Koch, L., 1873. DieArachnidenAustraliens. Niirnberg 1: 369-472.
Levitt, V., 1961. The funnel web spider in captivity. Proceedings of
the Royal Zoological Society (NSW) 1958: 80-84.

Main, B.Y., 1967. Spiders of Australia. Jacaranda Press, 125 pp.
Main, B.Y., 1976. Spiders. Collins, Sydney, 296 pp.

Main, B.Y., 1982. Some zoogeographic considerations of f ami lies of
spiders occurring in New Guinea. In Biogeography and Ecology
in New Guinea, ed. J.L. Gressitt, pp. 583-602. The Hague: Junk.
Main, B.Y., 1985. Arachnida : Mygalomorphae. In Zoological
Catalogue of Australia 3: 1-183.

Mascord, R.E., 1980. Spiders of Australia: A Field Guide. Sydney:
A. H. and A. W. Reid.

McKeown, K.C., 1963. Australian Spiders. Sydney: Sirius Books:
Angus and Robertson.

Musgrave, A., 1927. Some poisonous Australian Spiders. Records
of the Australian Museum 16: 33^46.

https://dx.doi. org/10.3853/j.0067-1975.16.1927.777

Musgrave, A., 1948. A catalogue of the spiders of Tasmania.

Records of the Queen Victoria Museum 2: 75-91.

Nishikawa, Y., 1976. Notes on the venomous spiders (funnel-web
spider and red-back spider) in Australia. Otemon Bulletin for
Australian Studies 2: 176-193.

Platnick, N.I., 2010. The world spider catalogue, version 9.5.
American Museum of Natural History, online at:

http://research.amnh.org/entomology/spiders/catalogue/index.html

Rainbow, W.J., 1911. A census of Australian Araneidae. Records
of the Australian Museum 9: 107-319.

https://dx.doi.org/10.3853/j.0067-1975.9.1911.928

Rainbow, W. J., 1913. Arachnida from the Solomon Islands. Records
of the Australian Museum 10: 1-16.

https://dx.doi.org/10.3853/j.0067-1975.10.1913.893

Rainbow, W.J., 1914. Studies in the Australian Araneidae. No.
6. The Terretelariae. Records of the Australian Museum 10:
187-270.

https://dx.doi.org/10.3853/j.0067-1975.10.1914.901

Rainbow, W.J., 1920. Trap-door spiders of the “Chevert” Ex¬
pedition. Records of the Australian Museum 13: 77-85.

https://dx.doi.org/10.3853/j.0067-1975.13.1920.857

Rainbow, W.J., & R.H. Pulleine, 1918. Australian trap-door spiders.
Records of the Australian Museum 12: 81-169.

https://dx.doi.org/10.3853/j.0067-1975.12.1918.882

Raven, R.J., 1980. The evolution and biogeography of the mygalo¬
morph spider family Hexathelidae (Araneae, Chelicerata).
Journal of Arachnology 8: 251-266.

Raven, R.J., 1985. The spider infraorder Mygalomorphae
(Araneae): Cladistics and systematics. Bulletin of the American
Museum of natural History 182: 1-180.

Raven, R.J., 2000. A new species of funnel-web spider (Hadron¬
yche, Hexathelidae, Mygalomorphae) from north Queensland.
Memoirs of the Queensland Museum 46: 225-230.

Roewer, C.F., 1942. Katalog der Araneae 1: 1-1040. Bremen.
Simon, E., 1891. Etudes arachnologiques. 23eMemoire. XXXVIII.
Descriptions d’especes et de genres nouveaux de la famille des
Aviculariidae. Annales Societe entomologique de France 60:
300-312.

Gray: Australian funnel-web spiders

391

Simon, E., 1892. Histoire naturelle des araignees. 1: 1-256. Paris.
Simon, E., 1903. Histoire naturelle des araignees. 2: 669-1080.
Paris.

Snazel, R., & R. Allison, 1989. The genus Macrothele Ausserer
(Araneae, Hexathelidae) in Europe. Bulletin of the British
arachnological Society 8(3): 65-72.

Sutherland, S.K., & J. Tibballs, 2001. Australian Animal Toxins.

Oxford University Press, 856 pp.

Walker, R, 1982. The tree funnel-web spider, Atrax formidabilis.
Australian Arachnology 7: 4-5

Wilson, D.T.R., 2001. The identification and characterisation of
Australian funnel-web spider venom. PhD thesis. University
of Queensland.

Wishart, G., 1993. The biology of spiders and phenology of
wandering males in a forest remnant. Memoirs of the Queensland
Museum 33(2): 675-680.

Woodman, J.D., J.E. Ashe & D.M. Rowell, 2006. Population
structure in a funnelweb spider (Hexathelidae: Hadronyche)
along a forested rainfall gradient. Journal of Zoology 268 (4):
325-333.

https://dx.doi.org/10.1111/j. 1469-7998.2005.00012.x

Manuscript submitted 23 September 2009, revised 12 August 2010, and
accepted 25 August 2010.

Appendix 1

Morphological characters and measurement points

A Palpal organ. (1) Bulb length—tegulum+embolus. (2) Bulb width—across tegulum and
subtegulum. (3) Embolus length. (4) Embolus mid-width. (5) Offset of basal embolus from
tegulum. (6) Twisted and “flanged” distal part of embolus.

B Cheliceral tooth row: length and mid-width.

C Carapace, dorsal: length, width and clypeal frontal width.

D Labium: length, width. Sternum: length, width. (Note: the presence of setae on the posterior
labium helps differentiate it from the smooth hairless cuticle of the labiosternal sigillum

E Leg segment lengths: femur, patella, tibia, metatarsus, tarsus.

F Posterior lateral spinnerets, apical segment: length and mid-width.

G Male palpal tibia: length, width.

H Carapace, lateral: height.

392

Records of the Australian Museum (2010) Vol. 62

Appendix 2

List of atracine spider species, with numbers previously allocated to new species. Undescribed species dealt with in Gray
(1986,1987, 1988) were identified by species numbers. These numbers are given below with the corresponding new species
names allocated here. Note that number 5 in the Hadronyche species number list is intentionally omitted.

species names species number

Atrax robustus O.P.-Cambridge, 1877

Atrax sutherlandi n.sp. Atrax sp.l

Atrax yorkmainorum n.sp. Atrax sp.2

Hadronyche adelaidensis (Gray, 1984)
Hadronyche anzses Raven, 2000
Hadronyche cerberea L. Koch, 1873
Hadronyche eyrei (Gray, 1984)

Hadronyche formidabilis (Rainbow, 1914)
Hadronyche flindersi (Gray, 1984)

Hadronyche infens a (Hickman, 1964)
Hadronyche meridiana Hogg, 1902
Hadronyche modesta (Simon, 1891)
Hadronyche pulvinator (Hickman, 1927)
Hadronyche valida (Rainbow & Pulleine, 1918)
Hadronyche venenata (Hickman, 1927)
Hadronyche versuta (Rainbow, 1914)

Hadronyche emmalizae n.sp. Hadronyche sp. 1

Hadronyche alpina n.sp. Hadronyche sp. 2

Hadronyche marracoonda n.sp. Hadronyche sp. 3

Hadronyche monaro n.sp. Hadronyche sp. 4

Hadronyche tambo n.sp. Hadronyche sp. 6

Hadronyche nimoola n.sp. Hadronyche sp. 7

Hadronyche mascordi n.sp. Hadronyche sp. 8

Hadronyche jensenae n.sp. Hadronyche sp. 9

Hadronyche orana n.sp. Hadronyche sp. 10

Hadronyche lynabrae n.sp. Hadronyche sp. 11

Hadronyche kaputarensis n.sp. Hadronyche sp. 12

Hadronyche levittgreggae n.sp. Hadronyche sp. 13

Hadronyche macquariensis n.sp. Hadronyche sp. 14

Hadronyche walkeri n.sp. Hadronyche sp. 15

Hadronyche raveni n.sp. Hadronyche sp. 16

Hadronyche lamingtonensis n.sp. Hadronyche sp. 17

Hadronyche annachristiae n.sp. Hadronyche sp. 18

Hadronyche monteithi n.sp. Hadronyche sp. 19

Illawarra xvisharti n.sp. Hadronyche sp. 20

© The Authors, 2010. Journal compilation © Australian Museum, Sydney, 2010
Records of the Australian Museum (2010) Vol. 62: 393-394. ISSN 0067-1975
doi: 10.3853/j.0067-1975.62.2010.1564

Note on Hydroides malleolaspinus from the Kimberleys of
Western Australia (Polychaeta: Serpulidae)

Anna Murray 1 *, Pat Hutchings 1 & T. Gottfried Pillai 2

1 Australian Museum, 6 College Street, Sydney NSW 2010, Australia
anna.murray@austmus.gov.au

2 Zoology Department, Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom

Abstract. Pillai (2009) described a series of specimens from the Kimberley area, Western Australia,
considered them to belong to an unknown species and proposed the name “ Hydroides trihamulatus” for
them. In error, no type material was designated, so the name is unavailable. After reexamination of the
same specimens, as well as the holotype of Hydroides malleolaspinus Straughan, 1967, we found that
the specimens correspond to this latter species.

Murray, Anna, Pat Hutchings & T. Gottfried Pillai, 2010. Note on Hydroides malleolaspinus from the
Kimberleys of Western Australia (Polychaeta: Serpulidae). Records of the Australian Museum 62(3): 393-394.

Pillai (2009: 131-134) described a new species for which
no types were designated (“ Hydroides trihamulatus ,, ).
Because of this error the name is unavailable. Initially we
thought to correct this by reexamining the same specimens,
nominating type material, and describing them as a new
species. However, as Hydroides malleolaspinus Straughan,
1967, was considered to be most similar to “ H. trihamulatus ”
(see Pillai’s remarks on p.132), we also examined that
holotype and other specimens identified by Straughan as H.
malleolaspinus. We then discovered features on the holotype
of H. malleolaspinus that were not included in Straughan’s
original description, and which Pillai (2009) used as
distinguishing characters separating the two species. We
herein expand the description of Hydroides malleolaspinus
Straughan, 1967, to include these characters.

Genus Hydroides Gunnerus, 1768

Type species. Hydroides norvegica Gunnerus, 1768.

Diagnosis. See Pillai (2009), and ten Hove & Kupriyanova
(2009).

Hydroides malleolaspinus Straughan, 1967
Fig. 1A-B

Hydroides malleolaspina Straughan, 1967:222-224, figs. 7a-f.
“Hydroides trihamulatus” Pillai, 2009:131-134, figs. 24A-G,
25A-C, 26A-L (name unavailable, types not designated).

Material examined. Holotype of Hydroides malleolaspinus Straughan,
1967, AM W3999, coll. fromPialba, Queensland, 25°16’44"S 152°50’32"E,
June 1962 by G. McKeon. AM W3996 (1) identified by D. Straughan, from
Shoal Point, Mackay, Queensland, coll. 7 Aug 1964, by I. Straughan. AM
W4109 (1) identified by D. Straughan, from Heron Island, Queensland,
coll. 1955 by B. Dew.

“Hydroides trihamulatus”'. AM W202944 (5 specimens, all incomplete and
only one with operculum present, removed from tubes, one posterior end
also in vial), from Kimberley area, Western Australia, from sand cay on
Port George IV, 15°20'S 124°39’E, St. 26, coll. 12 July 1988, by dredge.
AM W21412 (5 specimens, of which two are anterior ends only and lack
tubes, one is juvenile, incomplete posteriorly but possesses a tube, and one
consists of radiolar crown of one side only), from Kimberley area, Western
Australia, from sand cay on Port George IV, 15°20'S 124°39'E, St. 26, coll.
12 July 1988, by dredge. AM W202939 (1 radiolar crown from one side
only, plus operculum), from sand cay on Port George IV, 15°20'S 124°39'E,
12 July 1988, St. 26. AM W202943 (1 adult specimen lacking tube), from
reef north west of Buffon Island, 14°55'S 124°48E, 12 July 1988, St. 85.

* author for correspondence

394

Records of the Australian Museum (2010) Vol. 62

AM W21469, (1 juvenile), from south west corner of Lucas Island,
Kimberleys, Western Australia, 15°13'S 124°31'E, 30 m, 24 July
1988, St. 101. All specimens coll, by P.A. Hutchings.

Description. Holotype incomplete posteriorly,
tube missing, with 7 thoracic and c. 80 abdominal
chaetigers for length of 22 mm, maximum thoracic
width 2.0 mm, 17 radioles plus operculum on right
side, 21 radioles plus rudimentary operculum
on left side, opercular length 2.1 mm, length of
operculum with peduncle 5.5 mm, operculum with
9 unmodified coronal spines with T-shaped tips, plus
1 enlarged modified coronal spine with distal lateral
“points” and large bulbous process directed medially,

21 infundibular spines with T-shaped tips (Fig. 1A); all
spines lack a basal process.

Variation. Specimen from AM W3996 complete but
damaged posteriorly, 17.0 mm in length, maximum thoracic
width 2.0 mm, tube present for most of abdomen, dirty-
white with 3 longitudinal ridges, radioles regrowing, 19
radioles on each side plus rudimentary operculum on left
and operculum on right, opercular length 2.0 mm, 10 un¬
modified coronal spines with T-shaped tips, plus 1 enlarged
modified coronal spine with small distal lateral “points”
and large medial beak-like process directed perpendicular
to axis of spine, 28 infundibular spines with T-shaped
tips (Fig.IB). Specimen from AM W4109 incomplete,
damaged, juvenile, length of 1.5 mm with 3 abdominal
chaetigers, maximum thoracic width 0.4 mm, tube missing,
6 radioles on each side plus rudimentary operculum on right
and operculum on left, opercular length 0.7 mm, 7 unmodi¬
fied coronal spines with T-shaped tips, plus one modified,
enlarged coronal spine with slight medial bulge, 20 infun¬
dibular spines with blunt tips.

All specimens cited by Pillai (2009) under “ Hydroides
trihamulatus ” are incomplete posteriorly, and are smaller
than the type specimen of H. malleolaspinus and specimen
AM W3996, but most are larger than specimen AM W4109.
Maximum thoracic width ranges from 0.5-1 mm, opercular
length ranges from 0.6-1.3 mm, number of radioles
varies from 6-10 per side (plus operculum or rudimentary
operculum), coronal spine number varies from 10-15,
shape of the modified spine ranges from a small bulge to
a beak-like process perpendicular to the axis of the spine.
Number of infundibular lobes/spines ranges from 17-26.
Coronal and infundibular spines show variations associated
with ontogenesis. For a description of the size-related
variability of spine shape/form, refer to Pillai (2009: 131,
figs. 24C-G, 25A-C, 26E-F). Tube is square to trapezoidal
in cross-section; 2-3 longitudinal ridges present along tube:
smaller specimens possess two longitudinal ridges (Pillai,
2009: figs. 24A,B, 26A-C), which may only be weakly
developed (Pillai, 2009: fig. 24E); the larger specimen AM
W3996 possesses three longitudinal ridges along the tube.
For detailed description of chaetae refer to Pillai (2009:
131-132, fig. 26G-L).

Figure 1. Hydroides malleolaspinus. (A) operculum of AM W3999,
holotype; ( B ) operculum of AM W3996. Scale bar is 1 mm.

Remarks. Straughan (1967) described the enlarged modified
coronal spine on the operculum of H. malleolaspinus as
being “somewhat hammer-shaped”, she made no mention
of a medial process, and her illustration (1967: fig. 7a) is
ambiguous. Having confirmed that the holotype and another
specimen identified by Straughan, possess a “bifid” tip and a
large beak-like medial process on the modified coronal spine
of the operculum, we have expanded the description of H.
malleolaspinus to include these characters. We thus conclude
that the specimens cited by Pillai (2009) under “ Hydroides
trihamulatus ” represent this species. The variability in
opercular spine form and count lies within the range that
we regard as normal for conspecific individuals of varying
age and size. The name “ Hydroides trihamulatus ,, should be
treated as unavailable. The geographical distribution of H.
malleolaspinus is thus expanded to include NW Australia.

References

Hove, H.A. ten, & E.K. Kupriyanova, 2009. Taxonomy of
Serpulidae (Annelida, Polychaeta): the state of affairs. Zootaxa
2036: 1-126.

http://www.mapress.com/zootaxa/list/2009/zt02036.html

Pillai, T. G., 2009. Descriptions of new serpulid polychaetes from
the Kimberleys of Australia and discussion of Australian and
Indo-West Pacific species of Spirobranchus and superficially
similar taxa. Records of the Australian Museum 61(2): 93-199.

http://dx.doi.org/10.3853/j.0067-1975.61.2009.1489

Straughan, D., 1967. Marine Serpulidae (Annelida: Polychaeta) of
Eastern Queensland and New South Wales. Australian Journal
of Zoology 15: 201-261.

http://dx.doi.org/10.1071/Z09670201

Records of the Australian Museum (2010) Volume 62. ISSN 0067-1975

CONTENTS

Volume 62 • Numbers 1-3 • 2010

Archer, Michael (see under Worthy).77

Bochenski, Zbigniew M. (see under Campbell).123

Boles, Walter E., 2010. A revision of C.W. De Vis’ fossil cormorants (Aves: Phalacrocoracidae).

In Proceedings of the VII International Meeting of the Society of Avian Paleontology and

Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 145-155.145

doi: 10.3853/j.0067-1975.62.2010.1533

Boles, Walter E., and Trevor H. Worthy, 2010. Foreword to Proceedings of the VII International
Meeting of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H.

Worthy. Pp. 1-6.1

doi: 10.3853/j.0067-1975.62.2010.1532

Boles, Walter E. (see also under Nguyen).45

Boles, Walter E. (see also under Gaff).71

Campbell, Kenneth E., and Zbigniew M. Bochenski, 2010. A new genus for the extinct Late
Pleistocene owl Strix brea Howard (Aves: Strigiformes) from Rancho La Brea, California.

In Proceedings of the VII International Meeting of the Society of Avian Paleontology and

Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 123-144.123

doi: 10.3853/j.0067-1975.62.2010.1534

Dyke, Gareth J., and Gary W. Kaiser, 2010. Cracking a developmental constraint: egg size and
bird evolution. In Proceedings of the VII International Meeting of the Society of Avian

Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 207-216.207

doi: 10.3853/j.0067-1975.62.2010.1547

Dyke, Gareth J. (see also under O’Connor).7

Gaff, Priscilla, and Walter E. Boles, 2010. A new eagle (Aves: Accipitridae) from the Mid Miocene
Bullock Creek Fauna of northern Australia. In Proceedings of the VII International Meeting
of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy.

Pp. 71-76.71

doi: 10.3853/j.0067-1975.62.2010.1557

Geraads, Denis (see under Mourer-Chauvire).157

Gill, B. J., 2010. Regional comparisons of the thickness of moa eggshell fragments (Aves:
Dinornithiformes). In Proceedings of the VII International Meeting of the Society of Avian

Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 115-122.115

doi: 10.3853/j.0067-1975.62.2010.1535

Gohlich, Ursula B., and Cecile Mourer-Chauvire, 2010. A new cormorant-like bird (Aves,
Phalacrocoracoidea) from the Early Miocene of Rauscherod (Southern Germany). In
Proceedings of the VII International Meeting of the Society of Avian Paleontology and

Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 61-70.61

doi: 10.3853/j.0067-1975.62.2010.1536

Gray, Michael R., 2010. A revision of the Australian funnel-web spiders (Hexathelidae: Atracinae).

Pp. 285-392.285

doi: 10.3853/j.0067-1975.62.2010.1556

Hand, Suzanne J. (see under Nguyen).45

Hand, Suzanne J. (see also under Tennyson).105

Hutchings, Pat (see under Murray).393

Jones, Craig M. (see under Tennyson).105

Kaiser, Gary W. (see under Dyke).207

Kohler, Frank, 2010. Uncovering local endemism in the Kimberley, Western Australia: description
of new species of the genus Amplirhagada Iredale, 1933 (Pulmonata: Camaenidae).

Pp. 217-284.

doi: 10.3853/j.0067-1975.62.2010.1554

217

Mayr, Gerald, 2010. A new avian species with tubercle-bearing cervical vertebrae from the Middle
Eocene of Messel (Germany). In Proceedings of the VII International Meeting of the Society

of Avian Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 21-28.21

doi:10.3853/j.0067-1975.62.2010.1537

Mourer-Chauvire, Cecile, and Denis Geraads, 2010. The Upper Pliocene avifauna of Ahl al
Oughlam, Morocco. Systematics and biogeography. In Proceedings of the VII International
Meeting of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H.

Worthy. Pp. 157-184.157

doi:10.3853/j.0067-1975.62.2010.1538

Mourer-Chauvire, Cecile (see also under Gohlich).61

Murray, Anna, Pat Hutchings & T. Gottfried Pillai, 2010. Note on Hydroides malleolaspinus from

the Kimberleys of Western Australia (Polychaeta: Serpulidae). Pp. 393-394.393

doi:10.3853/j.0067-1975.62.2010.1564

Nguyen, Jacqueline M. T., Walter E. Boles and Suzanne J. Hand, 2010. New material of
Barawertornis tedfordi, a dromornithid bird from the Oligo-Miocene of Australia, and its
phylogenetic implications. In Proceedings of the VII International Meeting of the Society

of Avian Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 45-60.45

doi:10.3853/j.0067-1975.62.2010.1539

O’Connor, Jingmai, and Gareth J. Dyke, 2010. A reassessment of Sinornis santensis and
Cathayornis yandica (Aves: Enantiornithes). In Proceedings of the VII International
Meeting of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H.

Worthy. Pp. 7-20.7

doi:10.3853/j.0067-1975.62.2010.1540

Pillai, T. Gottfried (see under Murray).393

Prassack, Kari A., 2010. Late Pliocene avifauna from the hominid-bearing Zinjanthropus land
surface at Olduvai Gorge, Tanzania. In Proceedings of the VII International Meeting
of the Society of Avian Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy.

Pp. 185-192.185

doi:10.3853/j.0067-1975.62.2010.1541

Scofield, R. Paul, Trevor H. Worthy and Alan J. D. Tennyson, 2010. A heron (Aves: Ardeidae)
from the Early Miocene St Bathans fauna of southern New Zealand. In Proceedings of the
VII International Meeting of the Society of Avian Paleontology and Evolution, ed. W.E.

Boles and T.H. Worthy. Pp. 89-104.89

doi:10.3853/j.0067-1975.62.2010.1542

Scofield, R. Paul (see also under Worthy).77

Scofield, R. Paul (see also under Tennyson).105

Tennyson, Alan J. D., Trevor H. Worthy, Craig M. Jones, R. Paul Scofield & Suzanne Hand, 2010.

Moa’s Ark: Miocene fossils reveal the great antiquity of moa (Aves: Dinornithiformes)
in Zealandia. In Proceedings of the VII International Meeting of the Society of Avian

Paleontology and Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 105-114.105

doi:10.3853/j.0067-1975.62.2010.1546

Tennyson, Alan J. D. (see also under Worthy).77

Tennyson, Alan J. D. (see also under Scofield).89

Tyrberg, Tommy, 2010. Avifaunal responses to warm climate: the message from Last Interglacial
faunas. In Proceedings of the VII International Meeting of the Society of Avian Paleontology

and Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 193-205.193

doi:10.3853/j.0067-1975.62.2010.1543

Weidig, Ilka, 2010. New birds from the Lower Eocene Green River Formation, North America.

In Proceedings of the VII International Meeting of the Society of Avian Paleontology and

Evolution, ed. W.E. Boles and T.H. Worthy. Pp. 29-44.29

doi:10.3853/j.0067-1975.62.2010.1544

Worthy, Trevor H., Alan J. D. Tennyson, Michael Archer, and R. Paul Scofield, 2010. First record
of Palaelodus (Aves: Phoenicopteriformes) from New Zealand. In Proceedings of the VII
International Meeting of the Society of Avian Paleontology and Evolution, ed. W.E. Boles

and T.H. Worthy. Pp. 77-88.77

doi:10.3853/j.0067-1975.62.2010.1545

Worthy, Trevor H. (see also under Boles).1

Worthy, Trevor H. (see also under Scofield).89

Worthy, Trevor H. (see also under Tennyson).105

INSTRUCTIONS TO AUTHORS

Manuscripts must be submitted to The Editor. All manuscripts are
refereed externally.

Only those manuscripts that meet the following requirements
will be considered for publication. Large monographic works are
considered for publication only when the subject matter aligns closely
with the Museum's strategic objectives.

Submit manuscripts electronically and as one printed copy; images
should be high resolution TIFFs (see below). Attach one summary file
or cover sheet giving: the title; the name, address and contact details
of each author; the author responsible for checking proofs; a suggested
running-head of less than 40 character-spaces; and the number of
figures, tables and appendices. Manuscripts must be complete when
submitted.

Text files, tables and charts should be in Rich Text Format (RTF).
Tables and figures should be numbered and referred to in numerical
order in the text. Electronic copy is stripped and reconstructed
during production, so authors should avoid excessive layout or textual
embellishments; a single font should be used throughout; avoid using
uncommon fonts.

All copy is manipulated within a Windows (not Mac) environ¬
ment using Microsoft and Adobe software. The submitted printed
copy of the manuscript should be derived directly from the electronic
file that accompanies it.

Manuscripts should be prepared using recent issues as a guide. There
should be a title (series titles should not be used), author(s) with their
institutional addresses, an abstract (should be intelligible by itself,
informative not indicative), introduction (should open with a few lines
for general, non-specialist readers), materials and methods, results
(usually subdivided with primary, secondary and rarely tertiary-level
headings), discussion, acknowledgments and references. If appropriate,
an appendix may be added after references.

In the titles of zoological works the higher classification of
the group dealt with should be indicated. Except for common
abbreviations, definitions should be given in the materials and
methods section. Sentences should not begin with abbreviations or
numerals. Metric units must be used except when citing original
specimen data. It is desirable to include geo-spatial coordinates;
when reference is made to them, authors must ensure that their
format precludes ambiguity, in particular, avoid formats that confuse
arcminutes and arcseconds.

Label and specimen data should, as a minimum requirement,
indicate where specimens are deposited. Original specimen data—
especially that of type material—is preferred over interpreted data.
If open to interpretation, cite original data between quotation marks
or use “[sic]”.

Rules of the International Code of Zoological Nomenclature must
be followed; authors must put a very strong case if a Recommend¬
ation is not followed. When new taxa are proposed in works having
multiple authors, the identity of the author(s) responsible for the
new name(s) and for satisfying the criteria of availability, should be
made clear in accordance with Recommendations in Chapter XI of
the Code. A scientific name with more than two authors is unwieldy
and should be avoided. Keys are desirable; they must be dichotomous
and not serially indented. Synonymies should be of the short form:
taxon author, year, pages and figures. A period and en-dash must
separate taxon and author except in the case of reference to the original

description. Proposed type material should be explicitly designated
and, unless institutional procedure prohibits it, registered by number
in an institutional collection.

Previously published illustrations will generally not be accepted.
Extra costs resulting from colour production are charged to the
author (AU$1000 for 1-8 pp, AU$2000 for 9-16 pp, etc.; these
charges can be shared by authors of different papers that are printed
consecutively). All images must (a) be rectangular or square and
scalable to a width of 83 mm (one text column) or 172 mm (both
text columns including gutter) and any depth up to 229 mm (the
number of lines in a caption limits depth); (b) have lettering similar
to 14 point, upper case, normal, Helvetica, in final print; (c) have no
unnecessary white or black space; and (d) have vertical or horizontal
scale bars, with the lengths given in the caption and with the
thickness approximately equal to an upper case 14 point letter “I”.

Digital images must be presented as TIFF, or as multilayered
PSD files suitable for Adobe Photoshop version 5.0 or later. Halftone
and colour images must be at a minimum resolution of 300 dpi at
final size (at this resolution 2040 pixels = printed-page width) and
all labelling must be sharp (with anti-aliased active). Black and white
line images (bitmaps) must be at a minimum resolution of 1200 dpi
at final size (at this resolution, 8160 pixels = page width = 172 mm).

When reference is made to figures in the present work use Fig. or
Figs, when in another work use fig. or figs; the same case-rule applies
to the words tables and plates. Figures and tables should be numbered
and referred to in numerical order in the text.

Authors should refer to recent issues of the Records of the Australian
Museum to determine the correct format for listing references and
to The Chicago Manual of Style to resolve other matters of style.
Insert hyperlinks in the Reference section if they are known—use
digital object identifiers (doi) if available (see www.doi.org and www.
crossref.org).

Certain anthropological manuscripts (both text and images)
may deal with culturally sensitive material. Responsibility rests
with authors to ensure that approvals from the appropriate person or
persons have been obtained prior to submission of the manuscript.

Stratigraphic practice should follow the International Stratigraphic
Guide (second edition) and Field Geologist's Guide to Lithostratigraphic
Nomenclature in Australia.

The Editor and Publisher reserve the right to modify manuscripts to
improve communication between author and reader. Essential corrections
only may be made to final proofs. No corrections can be accepted less than
four weeks prior to publication without cost to the author(s). All proofs
should be returned as soon as possible. Fifty reprints may be ordered, the
fee is AU$10.00 times the number of printed pages.

All authors, or the Corresponding Author on their behalf, must
sign a Licence to Publish when a manuscript is submitted, and certify
that the research described has adhered to the Australian Museum’s
Guidelines for Research Practice —or those of their home institution
providing they cover the same issues, especially with respect to
authorship and acknowledgment. While under consideration, a
manuscript may not be submitted elsewhere.

More information and examples are available at our website:

http://publications.australianmuseum.net.au

24 November 2010

Stock no. 010R62B

YOUR SUPPORT MAKES A DIFFERENCE

The Australian Museum strives to inspire the exploration of nature and cultures. We would like to acknowledge
the benefactors and corporate partners who support us in achieving this vision.

These generous individuals contribute to scientific research, education programs, public programs and assist
in the acquisition of items that enrich our collections. We would especially like to acknowledge those who
generously leave a gift to the Australian Museum in their will—a lasting way to benefit generations to come.

If you would like to find out how your support can make a difference to the important work of the Australian
Museum, please contact the Development Branch on +612 9320 6216 or development@austmus.gov.au.
Donations to the Australian Museum and its Foundation are tax deductible.

NSW Government through
Communities NSW

PRINCIPAL PARTNERS

Australian Museum Members
City of Sydney

Department of Environment and Climate
Change NSW
Geddes Group
IBM

JCDecaux

Lizard Island Reef Research Foundation
National Geographic Channel
Panasonic Australia

Raleigh and CPI + Edwards Dunlop Paper

Rio Tinto

Scitech

Sydney Grammar School

Sydney's Child

The Sydney Morning Herald

AUSTRALIAN MUSEUM FOUNDATION
President's Circle

Anita and Luca Belgiorno-Nettis
Tim and Kathryn Berry
Susan Conde
Zeny Edwards
Christopher Grubb
Ian and Stephanie Hardy
Mrs Judy Lee
Diccon Loxton

Richard and Jane Malcolmson
Helen Molesworth
Rob and Helen Rich
David and Daniela Shannon
The Sherman Foundation

Supporters

Antoinette Albert
James and Belinda Allen
Mr and Mrs K R Bell
Jane Beniac
Sir Ron Brierley
Dan Brush
Elizabeth Cameron
Estate of the late Clarence E
Chadwick
John Dickinson
Suellen and Ron Enestrom
Belinda Gibson
Leon Gorr

Owen Griffiths & Biodiversity
Conservation Madagascar Assn
Ken Handley AO and Diana Handley
Bill and Alison Hayward
Ann Hoban
Dan Howard SC and
Dr Rosemary Howard
Frank Howarth
Jonathan Kilborn
A J Loewenthal
David Maloney
Margaret Mashford
Roger Massy-Greene
Robert McDougall
Mabs Melville

Stephanie Miller and Martin Pool
Dame Elisabeth Murdoch AC DBE
John Neuhaus
Justice Henric Nicholas

The Noonan Family
David Norman
Campbell Norwood
Andrew Pardoe
John Pearson
Loris Peggie
Patricia Reid
Kate Richardson
David Robb

Jane and Neville Rowden
Hazel Sproule
Alastair Stevenson
Rosemary Swift
Senta Taft-Hendry
Davod Templeton
Estate of the late

Gwendoline A West
Jill Wran
Jennifer Wright

GIFTS TO COLLECTIONS

Ursula Burgoyne
Louise and Sam Dawson
Rod and Robyn Dent in honour of
Pat Dent and the
Wanindilyaugwa tribe
Mark Hanlon
The Late John Hume
James McColl
Dr William Rieger
Paul Scully-Power
Trevor Shearston
David Twine

nature culture discover

Australian Museum science is freely accessible online at
http://publications.australianmuseum.net.au

ISSN 0067-1975

RECORDS OF THE AUSTRALIAN MUSEUM 2010 Vol. 62 Nos 2 & 3