MEMOIRS

OF THE
QUEENSLAND MUSEUM
BRISBANE VOLUME 38

1 DECEMBER 1995 PART2

REVISION OF PHILIPIS (COLEOPTERA: CARABIDAE: BEMBIDIINAE), A GENUS
OF ARBOREAL TACHYINE BEETLES FROM THE RAINFORESTS OF EASTERN

AUSTRALIA: TAXONOMY, PHYLOGENY AND BIOGEOGRAPHY
MARTIN BAEHR

Baehr, M. 1995 12 01: Revision of Philipis (Coleoptera: Carabidae; Bembidiinae), a genus
of arboreal tachyine beetles from the rainforests of eastern Australia. Taxonomy, phylogeny
and biogeography. Memoirs of the Queensland Museum 38(2): 315-381, Brisbane, ISSN
0079-8835,

Philipis Erwin, 1994 was erected for the unique Tachys trunci Darlington. Extensive fogging
activities in montane rainforests of eastern Queensland produced rich material and the
following new species of Philipis are described: agnicapitis, alticola, alutacea, atra, bicolor,
castanea, cooki, distinguenda, ellioti, frerei, geoffreyi, heatherae, inermis, laevigata, laevis,
lustrans, minor, perstriata, picea, picta, planicola, quadraticollis, reticulata, rufescens,
ruficollis, sinuata, spurgeoni, striata, subtropica, sulcata, thompsoni, tribulationis, unicolor,
and vicina. Philipisis redefined and described, with P. trunci Darlington redescribed. Almost
all species occur in montane rainforest above 900m, generally in moss on tree trunks. The
genus occupies the Great Dividing Range in eastem Queensland from the Queensland/New
South Wales border to Mt. Finnigan south of Cooktown, with most species in northeastern
Queensland. Many species occur on single mountain tops or tablelands, fewer inhabit
adjacent uplands, and extremely few species have a wide range or occur on two distant ranges.
The Carbine Tableland and the Bellenden Ker Range east of Atherton Tableland are centres
of evolution and of species richness. From there the isolated mountain tops of МІ, Finnigan
and Thornton Peak in the north, and the higher tops along the east coast in the south have
been colonized, According to patterns of distribution and phylogenetic evidence the history
of the genus is hypothetized: Philipis may be a part of the so-called "Old Gondwanan
element" in Australia, with its nearest relatives perhaps the South American Xystosomus
lincage. High species diversity, however, may be a recent event: caused by uplift of the Great
Dividing Range during Pliocene and Pleistocene and its subsequent disintegration into
isolated tops and tablelands due to rapid erosion, and also disintegration of former widespread
rainforests as a consequence of the increasingly warmer and dries climate during late Tertiary.
These events divided the ranges of formerly widespread жен and supported allopatric
speciation on single mountain tops or ranges, Superimposed on this gradual climatic change
were other radical changes during Pleistocene that caused the rainforest to repeatedly retreal
up the mountains and then descend again. In that way populations of Philipis were repeatedly
divided then rejoined, eventually becoming split into the many closely related species.
According to phylogenetic evidence, the present 35 species éan te thus referred to only 5
original stocks, representatives of which occur in both centres of evolution. _] Coleoptera,
Carabidae, Bembidiinae, Philipis, phylogeny, biogeography.

Martin Baehr, Zoologische Siaatssanunlung, MünchhausenstraBe 21, D-81247 München,
Germany; 10 August 1995

Australian tachyine beetles (Tachys sensu lato
in the sense of Sloane, 1921) are not very numer-
ous (currently about 75 species are known), but
are very diverse. Hence, some of Sloane's species
groups have been since included in other genera,
e.g. Pericompsus Schaum (Erwin, 1974a),
Taüchyta Kirby (Erwin, 1975; Baehr, 1986),
Elaphropus Motschulsky (including Tachyura
Motschulsky, Sphaerotachys Müller, and thc
politus-group of Andrewes, 1925) (Erwin,
1974b: Baehr, 1987), or have been given even the
rank of a new genus, e.g. Tasmanitachoides
Erwin (Erwin, 1972; Baehr, 19902). Apart from
several species not yet revised but easily referred

to the well known genera Tachys Stephens sensu
stricto, Paratachys Casey (including Eotachys
Jeannel), and Polyderis Motschulsky, there still
remain other isolated species or small groups of
species of unknown relationships that are at pres-
ent best ranked as unspecified species-groups,
e.g. eclromoides-group (Darlington, 1962;
Baehr, 1989; 1991), atridermis-group (Tachys
atridermis Sloane, ? T. carinatulus Sloane),
mitchelli-group (T. mitchelli Sloane), mulwaten-
sis-group (T. mulwalensiy Sloane), blackburni-
group (T. blackburn! Sloane), and irunci-group
(T. trunci Dartington).

316

Although the habits of the Australian species of
Pericompsus, Tasmanitachoides, Elaphropus,
Tachyta, Tachys, Paratachys, and Polyderis are
generally known, little is known of the isolated
species groups or species mentioned above. It is
known that some species of the ectromoides-
group live in leaf litter of temperate rainforest,
others under bark of eucalypts, others on mossy
trunks of rainforest trees (Darlington, 1962;
Baehr, 1989; 1991), and the unique specimen of
Tachys trunci was also found on the trunk of à
tree in montane rainforest (Darlington, 1963).

While looking for additional material for a
review of Elaphropus (Baehr, 1987) in the ANIC,
Canberra, | found three specimens of a species
from the Lamington Plateau in southern Queens-
land that I was unable to identify atthe first glance
and that later appeared not to belong to the genus
Elaphropus. Geoff Monteith, Queensland Mu-
seum, informed me that he had found several
small tachyine species on mountain tops in North
Queensland by means of pyrethrum knockdown
on the mossy trunks of rainforest trees. He asked
me to identify his specimens and sent me a lot of
specimens which turned out to belong to the same
genus as those from Lamington Plateau. Since the
occurrence of another, numerous, group of arbo-
real tachyine beetles in eastern Australia neither
related to Tachyta nor to the Tachys ectromoides-
group was unexpected, I began work on these
specimens, because it would ratse most interest-
ing questions as to origin and relarionships of this
group, the phylogenctic relations of the many
included species, and their biogeographic history,
However, the word was set aside until more ma-
terial was available. Now enough material has
heen accumulated to work on this group, even
though additional new species may come to light
on as yet unworked mountain tops,

Detailed study of the description and later the
type specimen of Tachys trunci Darlington re-
vealed that this species belongs to the mentioned
group and is conspecific with one of the species
collected by Geoff Monteith on Thornton Peak,
the type locality of T. trunci.

Study of all available species revealed several
character states common to all species. Some are
apparently very primitive, the phylogenetic state
of others is still obscure, but they are rather un-
usual in Tachyini, In some phylogenetically rel-
evant characters (e. g. the truncate anterior tibia,
the lateral position of the apical striole, the elon-
gate terminal palpomere of both palpi), as well as
in сепа characteristics of shape and habitus,
this new genus 15 very similar to the South Amer-

MEMOIRS OF THE QUEENSLAND MUSEUM

ican genus Xystosomus Schaum (Erwin, 1973)
and its allies (Erwin, 1994) which include at least
some arboreal or semiarboreal species (Erwin,
1973; 1974b, 1994).

Erwin (1994) redefined the Xystosomus-lin-
eage and during his study, reexamined the type
specimen of Tachys trunci Darlington and
erected Philipis to accomodate it, Due to his very
limited material, Erwin's generic concept is, un-
fortunately, rather limited and the diagnosis given
in his key proves true only for Philipis trunci and
a few other species. Moreover, Erwin's key does
not permit uncquivocal differentiation of Philipis
from the American Mioptachys. 1 had no oppor-
tunity to study Mioptachys in detail, thocefare I
am not prepared to give a well founded differen-
tiation of both genera at present.

*
MEASUREMENTS

Measurements were made with a stereo micro-
scope using an ocular micrometer. Length was
measured from tip of labrum to apex of elytra,
hence these measurements may slightly differ
from those made by other authors, Length of
elytra was measured from humeral tooth to apex,
length of pronotum from anterior angle to middle
of basis, width of apex of pronotum between the
position where the anterior transverse sulcus
meets the anterior border, and width of head
including eves.

ABBREVIATIONS

Institutions: ANIC- Australian National Insect
Collection, Canberra; CBM- Collection M.
Baehr, München: MCZ-Muscum of Comparative
Zoology, Cambridge, Massachusetts; MDPI-
Queensland Departmen! of Primary Industries,
Mareeba; QM- Queensland Museum, Brisbane;
USNM- United States National Museum, Wash-
ington, D. C.; ZSM- Zoologische Staats-
sammlung, München.

Collectors: AG- Alistair Graham; DC- Doug
Cook; DY-David Yeates; ES- Evan Schmidt;
EW- Earthwatch; GM- Geoff Monteith; GT-
Geoff Thompson; HJ- Heather Janetzki; JF- John
Lawrence; LR- Lewis Roberts; QM- Queensland
Museum; RS- Rae Sheridan; SH- Simon Hamlet;
SM- Sybil Monteith; TW- Tom Weir.

RF- rainforest; Pyr.- Pyrethrum knockdown.

COLLECTING METRODS

Almost all material considered in this revision
has been collected by С. Monteith and his co-
workers al the Queensland Museum. Altogether
486 specimens have been examined, Because the

REVISION OF PHILIPIS

method used for the collection of almost all spec-
imens is not generally known, a description of the
method is provided here by Geoff Monteith:

Sheets of light nylon fabric about | Sm square
are laid fiat on the ground beneath any vertical
surfaces which can be found, These are mostly
the trunks of standing trees, both dead and alive,
but other surfaces such as large logs, stumps,
boulders, rock faces and earth banks are also
utilized. The vertical surfaces are then sprayed
with cans of household pyrethrum aerosol insec-
ticide. Small insects on the surfaces, which ate
usually densely mossy, fall on to the fabric sheets.
After about 30 minutes the sheets are carefully
picked up and the insects shaken into a suspended
fabric funnel with a jar of alcohol attached. The
bulk sample thus obtained rs later sorted under the
microscope. Fig. 1 shows the technique being
used in a north Queensland rainforest.

CHARACTERS

The main characters for the distinction of spe-
cies are shape of pronotum and elytra, colour and
pattern, degree of elytral striation, degree of
microreticulation of head, pronotum, and clytra,
and the male genitalia, especially shape of
aedeagus and its apex, structure of the sclerites in
the internal sac, and shape of parameres and
number and position of their apical setae.

TAXONOMIC PRINCIPLES

Many of the species mentioned herein are very
closely related and most of them belong to rather
well defined species groups. Only a few species
bear striking autapomorphic features. Although
many species possess fully developed wings, in
general they seem to be restricted to single moun-
tain formations; with very few widespread spe-
cies. Since most species groups have populations
dispersed over diverse mountain tops or ranges
and hence the species arc allopatric, they could
be described as groups of allopatric subspecies.
However, because the definition of subspecies is
principally more crucial than the definition of
species, | generally prefer to describe species
rather than subspecies, when the differences in
external and/or genitalic characters are constant,
even when they are rather feeble,

PHYLOGENETIC PRINCIPLES

In establishing phylogenetic relationships I
follow principles proposed by Hennig (1966) and
initially adopted by Brundin (1966). I did not
choose the quantitative phyletic approach (per-
haps better called "numerical cladistic") which

317

chiefly proceeds according to the principle of
parsimony because character analysis must be the
prior condition to the construction of a phyloge-
netic tree and the criterion of parsimony to be
used posteriorly to character analysis. Parsimony
may be perhaps not as important when tracing
phylogeny as most proponents believe. Evolution
of species does not necessarily proceed according
to that criterion, but commonly proceeds in a
circutious way in which case application of the
principle of parsimon may easily generate ficti-
tious results,

Although speciation occurs according to the
rules of the biological species concept, in insects
at least, its observation under natural conditions
is rarely possible. Hence characters are needed as
a means for distinguishing species. Although
morphological structures are most widely used,
other characters may be of the same value, e.g.
physiological, etho-ecological, cytological, or
genetical. But even genetics yield only characters
and hence do not solve the problem of applying
the hiological species concept to recognizing spe-
cies,

Reconstruction of the phylogeny and history of
the fauna or of à given superspecific taxon is
primarily based on the acquisition of adaptations
(i.e. characters), therefore analysis of the ances-
tral or derivative status of characters is generally
the only way to reconstruct phylogeny. As Hen-
nig (1966) demonstrated, only derivative
(apomorphic) character states are useful 1n such
phylogenetic classification, because they alone
define monophyletic groups and can be used to
establish sister group relations.

As a first step towards a phylogenetic classifi-
cation and analysis of faunal history the attempt
is made to determine the state of the characters
used. Character analysis, i.c. decision about
plesiomorphy and apomorphy of characters, is
based either on outgroup comparison using char-
acter states found in related taxa of higher cate-
gories as explained by Wiley (1981) and Watrous
& Wheeler (1981), or on group trends as ex-
plained by Ross (1974). The latter method was
used especially in such cases, when strictly syn-
apomorphic states were difficult to perceive be-
cause of parallelisms or reductions of characters.
So it will be noted in the phylogenetic analysis
that only common evolutionary trends can be
traced, rather than well founded sister group re-
lations.

318

HIOGEOGRAPHIC PRINCIPLES

Knowledge of the phylogenetic relations of the
species is prerequisite for any considerations
about historical biogeography. Indeed, 1 do not
trust the unweighed clues to geographical history
used by Darlington (1957, 1971) and repeated by
Erwin (1970). Hence, in general, 1 follow the
considerations of Hennig (1966) and Brundin
(1966) in that plesiomorphy of a taxon and its
geographic origin are commonly correlated, that
the most apomorphic taxa are therefore usually
found at the margins of the range of the supra-
specific taxon, and that this pattern of distribution
is mainly caused by vicariance biogeography and
commonly reflects a continuous distribution on
old land masses or in continuous biota that are
today dismembered. However, in same instances
it may be caused hy dispersal biogeography, e.g.
when new areas are colonized and later separated
by geographical or ecological barriers.

SYTEMATICS

Subfamily BEMBIDIINAE
Subtnbe XYSTOSOMINA

Philipis Erwin, 1994: 567

TYPE SPECIES
Tachys trunci Darlington, 1963, by monotypy.

NOTE

Erwin's description of Philipis was based on
the single species Tachys trunci and actually only
on the male holotype and on two females from G.
Manteith's material sent to him for examination.
Therefore his generic diagnosis and description
are both of limited value and a full redescription
of the genus 15 given below, Unfortunately the
characters for keying out the genus Philipis used
by Erwin (1994) in his key to the genera of the
subtribe Xystosomina are incidental characters of
P. trunci and do not allow the determination of
the genus for most species now known. I am not
familiar enough with the Neotropical taxa, espe-
cially Mioptachys Bates, to allow the proposal of
a generic key to replace Erwin's and thus the
genus Philipis remains recognisable only on
characters of distribution,

DIAGNOSIS

Facial sulci deep, anteriorly more or less dis-
tinctly doubled, reaching clypeus; terminal pal-
pomere of both palpi very elongate; anterior
transverse sulcus of pronotum distinct and usu-

MEMOIRS OF THE QUEENSLAND MUSEUM

ally deep; apex of protibia almost truncate, later-
ally at most slightly oblique; sutural stria deeply
impressed and anteriorly recurved; apical striole
nearer to lateral border than to suture, incurved
anteriorly and meeting 3rd stria or its position;
3rd stria with 2 discal setae and one seta within
the apical striole; aedeagus moderately elongate
with widely rounded apex, a complexly folded
sclerite inside internal sac, apex of internal sac
microtrichiate; parameres triangular, mostly 5-
setose, rarely 4-setose; female stylomere 2 ensi-
form, rather short, with two stout ventral ensiform
setae, a large dorsal ensiform seta situated rather
basally, and a nematiform seta originating in a
groove near apex,

The genus includes Philipis trunci (Darlington)
and 34 new species described below,

DESCRIPTION

Small (1.8-2.8mm long), either fairly elongate
and moderately depressed, or rather short and
convex species.

Colour black, or piceous, or reddish, with or
without a light elytral pattern consisting of a spot
ora transverse or oblique fascia in posterior third,
commonly also with a more or less distinct hu-
meral spot or fascia. In some species pronotum
red. Usually 1st-3rd antennomeres and basal half
of 4th antennomere of antenna yellow, rest dark.
Palpi and mouth parts yellow or partly infuscate,
Lower surface reddish, brown, anteriorly com-
monly slightly darker, Legs yellow or partly in-
fuscate.

Head short and compact. Eyes large. Neck
wide. Frons and vertex convex. Clypeus sepa-
rated from frons by shallow suture; frons usually
with shallow circular groove in middle, com-
monly also with circular or oblique groove medi-
ally of facial sulci, Labrum short, 6-setose.
Mandibles elongate, at apex incurved. Mentum
with prominent tooth, biselose, with two deep,
though not perforate foveae. Gula bisetose.
Glossa narrow, rather triangular at apex, 3- or
5-setose, with the longest seta in middle;
paraglossa membraneous, elongate, as long as
glossa or nearly so. Lacinia elongate, apex in-
curved, inner margin with a senes of strong,
elongate bristles. Penultimate palpomere of labial
palpus short, globose, pilose, apical palpomere
thin, remarkably elongate, usually almost as long
as penultimate palpomiere, Maxillary palpus
rather elongate, pilose, apical palpomere thin,
very elongate, Antenna rather short, sparsely pi-
lose from 2nd antennomere, densely pilose from
apical half of 4th anteünomere, Median an-

REVISION OF PHILIPIS

319

FIG. 1. Queensland Museum staff using the technique of pyrethrum knockdown in a North Queensland rainforest.
A, Nylon sheet spread on ground at base of tree while trunk is sprayed with aerosol pyrethrum. B, Fallen insects
on sheets are then shaken into a suspended funnel with alcohol jar attached,

tennomeres globose or up to twice as long as
wide.

Pronotum much wider than head. Base usually
much wider than apex; anterior angle rounded
off, not or barely projecting; lateral margins near
base but slightly convex or almost straight; pos-
terior angle usually about rectangular, commonly
posteriorly slightly projecting. Anterior trans-
verse sulcus always distinct, usually deep, v-
shaped, impunctate. Median line distinct, though
ususally fine, not reaching base. Posterior trans-
verse sulcus deep, impunctate, commonly inter-
rupted in middle by a deep fovea. Lateral channel
deep and rather wide, posteriorly more or less
widened; with two lateral setae, the anterior seta
situated at or behind anterior third, the posterior
at basal angle. Basal grooves deep, laterally bor-
dered by a conspicuous, elongate, usually straight
to slightly oblique carina. Prosternum with
sparse, erect pilosity.

Elytra fairly elongate and depressed, or more
or less convex and short. When elytra depressed,

then commonly with a distinct transverse impres-
sion in anterior third, sometimes each elytron
with a large, circular impression behind middle.
Lateral channel deep and wide. Lateral margin
commonly finely serrate and pilose. Sutural stria
always deeply impressed and anteriorly curved
towards scutellar pore. 2nd-7th striae present or
reduced to various degrees, or all or some striae
indicated as rows of punctures, or deeply chan-
nelled, or completely absent. 8th stria present,
usually attaining at least the posterior pore of the
anterior group of marginal pores, rarely ending
shortly behind this pore; usually deeply im-
pressed, commonly punctate-crenulate. Apical
striole situated rather close to lateral border, but
anteriorly more or less incurved to meet the end
of the 3rd stria or its position; striole usually deep,
deepened at the position of incurvation. 3rd stria
with two discal pores and setae at about anterior
third and behind middle, and with a pore and seta
within the apical striole. Marginal pores large,
setae not very elongate, but usually markedly

320

different, anlerior group consisting of 4, posterior
group of 2 + 2 pores, separated by a wide glabrous
space. Wings fully developed or more or less
redüced. Prosternum with few erect hairs.
Metepisternum almost quadrate to c. 1.5 x as ong
as wide at apex.

Abdomen smooth, impilose, each sternite with
a pair of ambulatory setae, last visible sternite in
males with 2, in females with 4 setae near border
in a transverse row.

Legs normal shaped, apex of protibia truncate,
lateral border not at all excised, Two basal tar-
someres of male protarsus slightly asymetrically
widened and clothed with adhesive setae. Claws
large, elongate, not serrate.

Microreticulation on head usually rather
coarse, isodiametric, though commonly reduced
and more or less superficial on frons and vertex,
but still present on labrum, clypeus and near
lateral margin. Pronotum with or without isodia-
metric microreticulation. Elytra with distinct or
superficial microreticulation, or without any
microreticulation. Microreticulation usually
transverse, rarely almost isodiametric. Some-
times intervals with very sparse, extremely fine
puncturation.

Male genital ring tnangular, feebly asymmet-
ric, Aedeagus variously shaped, with asymmetri-
cal basal lobe and usually short, convex apex.
Internal sac in middle with complexly coiled
sclerites. Apex nf internal sac with a densely
irichose or spinose field. Parameres with 4 or 5
apical or subapical setae, left paramere larger
than right.

Female stylomere 2 ensiform, with 2 strong
ventral ensiform setae, 1 dorsal ensiform seta
situated rather basally, and a nematiform seta
originating in a groove. Shape of stylomeres
highly umform throughout the genus.

DISTRIBUTION

Rainforests of eastern Queensland from the
Queensiand/New South Wales border north to
about Cooktown, though most species on the
eastern fringes of the Atherton and Carbine Ta-
blelands in north Queensland.

HABITAT

Almost all specimens collected by pyrethrum
knockdown from mossy tree trunks of rainforest
trees; a few were from Berlese extraction of moss
and litter. They occur with few exceptiuns on
high mountain tops above c. 900m,

MEMOIRS OF THE QUEENSLAND MUSEUM

KEY TO THE SPECIES OF THE GENUS
PHILIPIS ERWIN

Because many species are apparently restricted
toa single range or even to one mountain top, and
perhaps even to a narrow altitude range, the
known range of the species is gencrally recorded
in the key. It should be noted, however, that the
recorded geographic and altitudinal ranges may
change due to additional collecting work. So dis-
tribution should not be used as a prima facie

character for determination,
1. Elytra without distinct colourpattern .... . 2
Elytra with distinct colour pattem, each elytron
unimaculate or bimaculate, orfasciate 2... 5

2.Rather short, dorsally convex species, lateral bor-
ders of pronotum convex throughout, though
sometimes irregularly convex. Aedeagus as in
Figs 4D-F. Bellenden Ker Range, and moun-
tains near Eungella, northwest of Mackay .. 3
Rather elongate and depressed species, lateral
borders of pronotum in posterior half straight or
even faintly concave, though sometimes slightly
oblique, Aedeagus as in Fig, 4C or unknown.
panera near Mossman and near Cape Tribu-
ation

3, Smaller, shorter species, body length <2.25mm,
ratio length/width of elytra <1.3, Colour pice-
ous. Lateral margin of pronotum evenly
rounded, basal angle slightly produced back-
wards, Aedeagus, Fig. 4D, Bellenden Ker and
Malbon Thompson Ranges, between 900-1450m

SPE) ар MEL DD unicolar sp. nov.
Larger, more elongate species, body length
2.40mm, ratio length/width of elytra 1.37. Col-
our black. Lateral margin of pronotum irregu-
larly rounded, compressed in basal half, basal
angle not produced backwards. Aedeagus, Fig.
4E. Mt. Macartney north of Eungella, northwest
of Mackay... sese atra sp. nov.

4, Lateral borders of pronotum faintly concave, pro-
notum at base аз wide as in middle. At least 5
inner striae well visible. Aedeagus unknown,
Mossman Bluff at c.8S0m picea sp. nov.

Lateral borders of pronotum straight, pronotum at
hase narrower than in middle. Only 2 inner
striae well visible, others very faint. Acdeagus,
Fig. 4C. Thornton Peak and mountains west of
Cape Tribulation above ¢.750m ........

——— — — а Ээ ep el a castanea sp. nov,

5. Elongate, depressed species, elytra only with
postmedian fascia . ,
Short, convex species, pattern of elytra variahle

6. Elytra without distinct transverse impression in an-
terior third (doubtful species under both cou-
plets)

REVISION OF PHILIPIS

Elytra with distinct impression in anterior third

eom б kom 9 n lg n on

7, At least 4 inner intervals of elytra distinctly con-

R,

9,

10.

11.

12.

vex in anterior half. Large species, body length
>2.7mm. Pronotum in middle barely wider than
at base. Aedeagus unknown, Mountains north-
west of Mossman, Massey Range west of
Bellenden Ker Range, between 1000-1200m .

Only 2 inner intervals of elytra distinctly convex
in anterior half. Smaller species, body length

«2,35mm. Pronotum variable. Adeagus, Fig. m

or unknown

M c

Pronotum in middle perceptibly wider than at
base. Smaller species, body length «2.25mim. 2
inner intervals of elytra distinctly convex, Lower
surface of aedeagus straight, Fig. 4B. Mt. Spur-
geon above 1 100m ‚ laevigata sp. nov,

Pronotum in middle barely wider than at hase.
Larger species, body length c. 2.35mm. Only su-
tural interval of elytra distinctly convex,
Acdeagus unknown, Isley Hills south of Cairns

heatherae sp. nov.

Pronotum in middle barely wider than at base,
ratio widest part/base «1.04. Postmedian elyiral
fascia not s-shaped, Mountains east of Atherton
Tableland 10

Pronotum in middle distinctly wider than at base,
ratio widest part/base >1.07. Postmedian elytral
fascia morc or less distinctly s-shaped. Moun-
tains west of Mossman

«+ ж ж» o € m ж to t og ot og ot on n9 n n

11

Large species, body length >2.7mm. Colour dark
reddish, Transverse impression in anterior third
of elytra deep. 2nd stria distinctly impressed,
3rd-Sth striae distinct, Aedeagus unknown,
Bellenden Ker Range at 1560m . . i,
alticola sp. nov.
Smaller species, body length c. 2.35mm. Colour
dark piceous. Transverse impression in anterior
third of elytra shallow. 2nd stria barely im-
pressed, 3rd-5th striae very weak. Acdeagus
unknown. Isley Hills south of Cairns , . .
heatherae sp. nov.
Larger species, body length 22.35mm. Pronotum
wider, base wider, ralio base/apex 71.45.
Aedeagus longer with narrower apex, both
parameres 4-setose, Fig. 4A, Mountains im-
mediately west of Mossman above 1000m
cooki sp. nov,
Smaller species, body length «2.25mm. Prono-
tum narrower, base narrower, ratio base/apex
«1.40. Adeagus shorter with wider apex, both
parameres 5-selose, Fig. 4B. Mt. Spurgeon at
TOU: on sae nem roy om vs laevigata sp. nov.
Elytra unimaculate or -fasciate 13
Elytra distinctly bimaculate or -fasciale (doult-
ful species under both couplets)... =. 22

RRT ERLT fu

IX

quadraticollis sp. nov.

321

13, Elytra with circular or slightly transverse macula

14, Lar,

оғ fascia (Figs 12D-E). Pronotum reddish or pro-
motum remarkably narrow, ratio width/length
«1.35 , with natrow base, ratio base/apex «1.35.
Elytra Tather clongate, ratio length/width »1.35.
Aedeagus as in Figs AF, 5А. Mr. Finnigan . . 14
Elytra with conspicuously s-shaped, oblique fas-
cia (Figs I2F, 13B-D). Pronotum not reddish
and pronotum wider, ratio widrh/length 21.40,
with wider base, ratio hase/apex 71.38, Elytra
shorter, ratio length/width «1.35 , Aedeagus as
in Figs SB-F, 6A. Distribution further south. .
ia ida а у elio wis ‚15
Larges species, body length 32: 3mm. Pronotum
ish, wider, ratio width/Iength >1.40, with
wider base, ratio base/apex 21,45, Aedeagus,
Fig. AF. Above 850m ..... ruficollis sp. nov.
Smaller species, body length «2mm. Pronotum
dark piceous ta black, narrower, ratio
width/length «1.35, with narrow base, ratio
base/apex «1.35. Aedeagus, Fig. 5A. Above
110m inermis sp. nov.

+ - * + + + + t << ж

15, Lateral borders of pronotum in posterior half

straight, hase as wide as or even wider than wid-
est diameter in middle. Lower surface of
aedeagus almost straight, apex not perceptibly
curved down (Fig. 5C). Eastern foothills of
Bellenden Ker Range below 100m... ...
planicola sp, тоу.
Lateral borders of pronotum in posterior half
perceptibly convex, base distinctly narrower
than widest diameter in middle. Lower surface
of aedeagus either concave or apical third
curved down (Figs 5B, 50-Е, 6A), Distribution

nesq»... ж у = ро рва

different or upland species ......... 16
16, 5th-7th intervals not perceptibly weaker than
innef intervals „ааа 17

Sth-7th intervals markedly weaker than inner in-
tervals, or абе... 18

17, Striac not impressed, intervals not convex,

Microreliculation of elytra weak, surface glossy.
Base of pronotum wider, ratio base/npex 21.5.
Acdeagus, Fig, 5B, Cape Tribulation below
150m striata sp. nov.
Ѕіліае in basal half slighily impressed, intervals
perceptibly convex, Microreticulation of elytra
distinct, surface rather dull. Base of pronotum
less wide, гапо base/apex 1.45. Aedeagus un-
known. Bellenden Ker Range at 1000m .
reticulata sp. nov.
‚ Ist and 2nd striae equally deeply impressed, 3rd-
Sth striae very inconspicuous or absent,
Aedeagus му (Fig. 5D). Thornton Peak
above Ze tee ‚ trunci (Darlington)
2nd stria distinctly less deeply impressed than
| st, or not impressed, 3rd-Sth strige visible as
rows of punctures. Aedeagus longer (Pigs ae
Г, 6A), Distribution different

wor m o pomo ro tm orm + m oap om n

Ett + t t B Rm

322

19. Smaller species, body length «2.15mm. and base
of pronotum ralher narrow, ratio base/apex
<1.42. Acdeagus, Fig. 5E. Mountains west of
Cape Tribulation and Windsor Tableland above
JOM wie vee е. tribulationis sp. nov.
Commonly larger species, body Jength

>2.20mm, or base of pronotum wider, ratio
base/apex 21.45, Aedcagus as in Figs 5Е, 6A
Distribution different |... ... iss 20

20. Aedeagus shorter, with shorter apex and lower
surface nol perceptibly bisinuate (Fig. SF) and
surface of elytra without distinct microreticula-
tion and outer striae very fine and base of prono-
tum wider, ratio base/apex »1.45, Mountains
from Atherton Tableland south to Cardwell
Range above 74m, -, . , thompsoni sp. nov.
Either aedeagus longer, with longer apex and

lower surface perceptibly bisinuate (Fig. 6A)
or surface of elytra distinctly microreticulate or
outer striac rather distinet or base of pronotum
narrower, ratio base/apex «1.45, Distribution
different -sssaaa othe Pe wl

21. Microreticulation of elytra almost absent, уша
as glossy as pronotum, Aedeagus, Fig, 6A, М!
Misery south of Helenvale above 850m -~

s VMicina sp. nov.

Microreticulation of elytra distinct, clyira për-

ceptibly less glossy than pranotum. Aedcagus

Unknown. Mt. Spurgeon area $t 1330m . . .

spurgenni sp. nov.

22, Only sutüral stria of elytra present. Surface of
pronotum and elytra extremely glossy, without
any trace af microreticulalion. Wide, highly con-
vex species with very short elytra, ratio
length/width «128 , Aedengus, Fig. 6B. Thorn-
ton Peak above 000m... .. . laevis sp. nov.
At least traces of outer striae visible, Sucface

usually less glossy, at least with some traces of
microreticulation. Usually less wide and con-
vex species with longer elytra, ratio
length/width >1.29, rarely less, but then elytra
completely and deeply striate. Aedeagus differ-
ent. Distribution different... 2.2... - 23

23, Elytra completely striate and striae deeply im-
pressed, outer striae not perceptibly weaker than
inner, all intervals remarkably canvex
Acdeagus with elongate apex and deep concav-
ity in [ront of apex (Fig. ӨС), or unknown . . 24
At least outer striae not deeply impressed, dis-

tincily weaker than inner, often elytra nol com-
pletely striate, not all intervals-convex.
Aedeagus different... . ., ‚25

24. Wider and on the average larger st species, body
length 2.05-2.50mm, Pronotum wider, ratio
width/length 71.53, ratio base/apex 71,48, Ely-
tra shorter, ratio length/width «1.31. Aedeagus.
Fig. 6C, Mountains west of Mossman between
480-1300m , » . ЗМ/сата sp, nov.

Foro ro tor eo gr or f tr ng n9

mE VT

MEMOIRS OF THE QUEENSLAND MUSEUM

Narrower and smaller species, body length
2.05mm. Pronotum narrower, ratio
width/length 1.41, ratio base/apex 1.37. Elvira
langer, ratio length/width 21.38. Aedeagus un-
known, Mi, Bartle Frere above [600m , , . .

ae eer oe ae perstriata sp. nov.
, Posterior elytral fascia distinctly s-shaped and
basal elytral macula well delimited, triangular,
meeting lateral border... 50... "+46

Posterior elytral fascia variable, when s-shaped,
then basal elytral macula less well delimited,
often rather vague, and not meeting lateral bor-
ВСга: see es css estes (0420

2n, Pronotum reddish, elytral pattern very distinc
At least 6 inner striae well marked, though only
sulural stria impressed. Aedeagus, Fig, 7E, Mt.
Demi southwest of Mossman above 1100m . .
bu uM sv bicolor sp. nov.
Pronotum piceous, elytra! pattern less disunct.
A1 most 4 inner striae well marked and more or
less impressed, Aedeagus as in Figs 7D, 7F,
8A. Distribution different... 2. isa- 27
27, Elytra barely striate, only sutural stria distinct
and impressed, other striae very indistinct. On
the average larger species, body length 2.2 mm.
Acdeagus very short, with short apex (Fig. 7D).
Mt. Finnigan above 1100m , . , . picta sp. nov.
Elytra more extensively striate, at least 3 miner
striae distinct and somewhat impressed. On the
average smaller species, body length <2.2min-
Aedeagus longer, with longer apex (Figs TER
8A). Distribuuon different
28, Larger and wider species, body length 22, wk
ratio length/width of elytra «1.32, with wider
pronotum, ratio widlh/length 21.55. Lower sur-
face of aedeagus almost straight, apex narrower
(Fig. ТЕ). Mt. Spurgeon area above 1250m . .
geoffreyi sp. nov.
Smaller and narrower species, body length
2mm, ratio length/width of elytra 1.35, with
narrower pronotum, ratio width/length «1.5.
Lower surface of aedeagus distinctly bisinuate,
apex wider (Fig. 8A). Mossman Bluff west of
Mossman at 1180m ...... minor sp. nov,

[I
“л

ка us aws аана э „җә

29, Posterior fascia distinctly s-shaped . ‚М
Posterior fascia not distinctly s-shaped (doubt-

ful species under both couplets). . . . . . . 33

30., Larger species, body length 22.6mm ..... 31

Smaller species, body length <2.5mm , . , . 32

31, Wider species, ratio length/width of elytra

«1,36, with wider pronotum, ratio width/length

51.5, ratio base/apex 21.5, Lower surface of

nedeagus gently concave (Fig. 8B). Bellenden

Ker Range above 1500m . sinuata sp. nov.

Narrower species, ratio length/width of elytra
71.38, with narrowe\ pronotum, ratio
width/length «1.48, ratio base/apex «1.48.
Lower surface of aedeagus gently bisinuate

REVISION OF PHILIPIS

(Fig. 8C). Mossman Bluff west of Mossman
between 480-1200m . . distinguenda sp. nov,
32. More or less reddish species, but at least prono-
tum somewhat reddish. Pronotum and elytra
strongly microreticulate. Posterior elytral fascia
rather weakly s-shaped (Fig. 15A). Aedeagus
еи lower surface gently concave, parameres
4-selose (Fig. 7A). Bellenden Ker Range above
1500m rufescens sp. nav,
Piceous species, pronotum not reddish, Prono-
tum and elytra glossy, almost devoid of
microrcticulation. Posterior elytral fascia mark-
edly s-shaped (Fig. 16E), Acdeagus short, com-
pact, lower surface bisinuate, parameres
5-seinse (Fig. 8D). Bellenden Ker Range and
Lambs Head, between 900-1200m . , 2...
lustrans sp. nov.
. Pronotum remarkably wide at base, ratio
base/apex >1.6, ratio widest diameter/base
«1.06. Microrcticulation of elytra isodiametric,
surface rather dull. Lower surface of aedeagus
markedly bisinuate, apex very wide (Fig. ӨР),
Bellenden Ker above 1500m . alutacea sp, nov.
Pronotum less wide at base, ratio base/apex
«1.5, ratio widest diameter/base 71.10. Micro-
reticulation of elytra transverse, surface less
dull. Aedeagus different (Figs GD-E, 7A-C) .
"m 34
34. Pronotum and elytra strongly microreticulate.
Lateral borders of pronotum posteriorly not con-
vex, though oblique. Posterior el He fascia
oblique, feebly s-shaped (Fig. 15A), Aedeagus,
Fig. 7A. Bellenden Ker Range above 1500m .
+ rufescens sp. nov.
Microreticulation of pronotum and elytra
weaker or absent. Lateral borders of pronotum
posteriorly usually perceptibly convex, when
straight, then posterior elytral fascia transverse.
Posterior fascia transverse. ar oblique, but not
at all s-shaped (Figs 14D-E, 15C-E), Aedeagus
as in Figs 6D-E, 7B-C, Distribution different

"+ mom scm Rom Bom om n om ж m P ot жу n t +

Ow ү „ 2. "u$ come ®

*oy 9 y og og! ® NAR CS M

35

35. Only sutural stria impressed. On average larger
species, body length 2.3-2.8mm. Posterior elyt-
ral fascia transverse. Aedeagus with elongate,
wide apex and à strongly sclerotized peg in inter-
nal sac (Fig. 6E). Lamington Plateau, south
Queensland, above 1000m

subtropica sp. nov.
Additional striae perceptibly impressed. On av-
erage smaller species, body length 1.85-
2.45mm. When larger than 2.3mm, then inner
3-4 striae markedly impressed and posterior
elytral fascia oblique. Aedcagus different (Figs
6D, 7B-C), north Queensland ў 36
36. Posterior elytral fascia oblique (Fig. 140). On
average larger species, body length 2.05-
2.45mm, with wider pronotum, ratio

323

width/length >1.6. 3-4 inner striae conspicu-
ously impressed and feebly punctate. Aedeagus,
Fig. 6D. Mi Bartle Frere above 1500m . . .
frerei sp. nov.
Posterior elytral fascia rather transverse (Figs
17C-D), On average smaller species, body
length 1.85-2.3mm, with narrower pronotum,
ratio width/length «1.57. Inner strjae less con-
spicuously depressed, but distinctly punctate,
Aedeagus as in Figs 7B-C. Distribution differ-
ent
37. Elytra slightly darker, hence pattern more con-
irasting. Lateral border of pronotum usually less
convex in posterior half, base wider, ratio
base/apex «1.38. Aedeagus narrower, with
larger apex, parameres with 4, rarely left
paramere with 5 setae (Fig. 7B). Mt. Elliot
above 1000m ellioti sp. nov.
Elytra slightly lighter, hence pattern less con-
trasting, Lateral border of pronotum usually
rather convex in posterior half, base narrower,
ralio base/apex «1,37. Aedeagus wider, with
smaller apex, both parameres always 5-seiuse
(Fig. 7С), Lambs Head near Edmonton, at
1200m agnicapitis sp. nov,

eblete 9$?4 1254257715

= о = + gy t oi 4+

б, eot o e pot gon

NOTE

In the following descriptions only those char-
acters are mentioned in detail that are distinctive
for the respective species.

Philipis quadraticollis sp. nov.
(Figs LIA, 16F, 17C)

MATERIAL EXAMINED

Ноготуре: QMT 13508; ©, Devils Thumb area 10km
NW Mossman, МЕО, 10 Oct 1982, 1000-1 180т, Mon-
teilh, Yeates & Thompson, Pyrethrum knockdown,
RF,

Paratype; | 9, 17°14'S x 145 "48'E Massey Ra,
6km NW of Bellenden Ker, МЕО, 115013, 11 Oct. 1991,
Pyr., GM & HJ (CBM),

DIAGNOSIS

Large, depressed, With an oblique fascia in
posterior half of elytra. Distinguished by absence
of transverse impression in anterior third of ely-
tra, large size, and conyex four inner intervals of
elytra.

DESCRIPTION

Measurements. Length: 2.7-2.8mm; width:
1,20-1,25mm. Ratios: Width pronotumvhead;
1.48-1.49; width/length of pronotum: 1.55-1.58;
width base/apex of pronotum: 1.57-1.60; width
widest diameter/base of pronolum: 0.99;
length/width of elytra: 1.43.

324

Colour. Chestnut brown, head slightly darker.
Sutural interval lighter posteriarly. Elytra in pos-
terior 2/5 with an oblique, at medial end strongly
recurrent yellow fascia not attaining suture,

Head. Median antennomeres c. 1.2 x as long
as wide. Microreticulation on frons distinct,
though somewhat superficial, isodiametric, on
vertex less distinct. Surface moderately glossy,
on vertex rather glossy.

Pronotum. Fairly depressed, transverse, widest
near base, base much wider than apex. Lateral
margin anteriorly strongly rounded, posteriorly
straight. Posterior angle rectangular, faintly pro-
duced over lateral part of base. Carina at posterior
angle elongate, oblique, slightly incurved. Ante-
rior transverse suleus comparatively shallow, un-
interrupted in middle. Posterior transverse sulcus
slightly interrupted in middle, Anterior lateral
seta situated slightly behind anterior third af mar-
gin. Surface almost devoid of microreticulation,
with scattered, extremely fine punctures, highly
glossy.

Elytra, Rather elongate, moderately convex,
widest in middle; lateral border almost evenly
curved, though in middle almost straight. Disk
not impressed in anterior third. Sutural stria finely
punctulate. 2nd-5th striae impressed, punctate,
6th and 7th striae visible as rows of fine punc-
tures. All innner striae traceable to far down apex.
Most intervals, apart from Tth and 8th, distinctly
convex, at least m anterior half. Sth stria deeply
impressed, punctate, attaining posterior marginal
pore, Recurrent striole deep, punctulate, clearly
meeting 3rd stria. Anterior discal pore in anterior
third, posterior pore in posterior 2/5.
Microreticulation on disk barely visible, at apex
slightly more distinct, composed of transverse
meshes. Surface with scattered, extremely fine
punctures, highly glossy. Wings fully developed,

Lower surface. Metepisternum 1.5 x as long as
wide,

Genitalia, Male unknown, Female stylomere 2
with dorsal ensiform seta situated rather basally,
- Variation. Apart from some variation of rela-
tive shape of pronotum, little variation noted.

DISTRIBUTION

Mountains to the east of Atherton Tableland
and to the northwest of Mossman, north Queens-
tand,

HABITAT

Both known specimens collected by pyrethrum
knockdown nn mossy tree trunks in montane
rainforests above 1000m.

MEMOIRS OF THE QUEENSLAND MUSEUM

ETYMOLOGY
Refers 10 the quadrate shape of pronotum.

REMARKS

Because both specimens are females, it is not
fully settled, whether they are actually conspe-
cific. Discovery of males from both localities
would be very useful,

Philipis alticola sp. nov.
(Figs 11B, 17C)

MATERIAL EXAMINED
HoLorvPE: QMT13509; 9.Bellenden Ker summit,
NEQ, 10 June 1980, б, B. Monteith.

DIAGNOSIS

Large, depressed, with oblique fascia on pos-
terior half of elytra. Distinguished by wide base
of pronotum that is almost as wide as in middle,
by fascta of elytra not s-shaped, and by impressed
2nd interval and fairly distinct 3rd-5th intervals.

DESCRIPTION

Measurements. Length; 2.75mm; width:
1.24mm. Ratios: Width pronotum/head: 1.41;
width/length of pronotum: 1.45; width base/apex
of pronotum: 1,48; width widest diameter/base of
pronotum: 1.04; length/width of elytra: 1.45.

Colour. Chestnut brown, head and lateral parts
of elytra slightly darker. Sutural interval posteri-
orly fecbly lighter. Elytra in posterior 2/5 with
oblique, feebly s-shaped, at medial margin not
recurrent yellow fascia not attaining suture.

Head. Median antennomeres c. 1,5 x as long
as Wide, Microreticulation on frons distinct, iso-
diametric, slightly less distinct on vertex. Surface
moderately dull, more glossy on vertex,

Pronotum. Fairly depressed, transverse, widest
well in front of middle, base much wider than
apex. Lateral margin posteriorly almost straight,
though somewhat oblique. Posterior angles rec-
tangular, faintly produced over lateral part of
base. Carina at posterior angle elongate, oblique,
slightly incurved. Anterior transverse sulcus
rather deep, interrupted in middle. Posterior
transverse sulcus very deep, interrupted by a
fovea. Anterior lateral seta situated at anterior
third of margin, Surface with fairly superficial,
almost isodiametric microreticulation, fairly
glossy,

Elytra. Rather elongate, moderately convex,
widest in middle. Lateral border almost evenly
curved, though in middle almost straight. Disk in
anterior third at position of anterior discal seta

REVISION OF PHILIPIS 325

FIG. 2. Dorsal view of Philipis thompsoni sp. nov.

with conspicuous, rather deep depression. Sutural
stria finely punctulate. 2nd stria almost corn-
pletely impressed, punctate, 3rd-5th striae visible
a8 rows of fine punctures. Outer striae only trace-
able in anterior half as rows of extremely fine
punctures. Intervals, apart from sutural and 2nd
intervals completely depressed. 8th stria deeply
impressed, punctate, anteriorly shallower, not at-
taining posterior marginal pore. Recurrent striole
elongate, oblique, meeting 3rd stria. Anterior dis-
cal pore in anterior third, posterior pore in poste-
rior 2/5. Microreticulation distinct, though fairly
superficial, composed of transverse meshes. Sur-
face rather glossy. Wings fully developed. Vari-
ation unknown.

Lower surface. Metepisternum c. 1.5 x as long
as wide.

Genitalia. Male unknown. Female stylomere 2
with dorsal ensiform seta situated rather basally.

DISTRIBUTION

Bellenden Ker Range, to the east of Atherton
Tableland, north Queensland. Known only from
type locality,

HABITAT

Collecting circumstances unknown. This is a
high mountain species, collected above 1500m in
montane rainforest in June.

ETYMOLOGY
Refers to the occurrence on the summit of
Bellenden Ker.

Philipis cooki sp. nov.
(Figs 4A, 11C, 17C)

MATERIAL EXAMINED

HOLOTYPE: QMT13510; d, 5.5km N. of Mt. Lewis,
via Julatten, NEQ, 1200m, 13 Sept 1981, G. Monteith
& D. Cook.

PARATYPES: 1 d, 1 9, same data (QM); 1 d, 2.5km
N Mt. Lewis via Julatten, NEQ, 3 Nov 1983, 1040m,
DY & GT, Pyr. in RF (QM); 2 d, Devils Thumb area
10km NW Mossman, МЕС, 10 Oct 1982, 1000-1180m,
GM, DY & GT, Pyr., RF (CBM, QM); 1 d, 1 9,
Mossman Bluff Summit 10km W Mossman, NEQ, 18
Dec 1988, 1300m, GM & GT, Pyr. Trees & Rocks
(QM); 1 3,3 9, Mt, Demi, 7km SW of Mossman,
NEQ, 29 Oct 1983, 1100m, DY & GT, Pyr. in RF
(CBM, QM, USNM, ZSM); 4 d, Carbine Tableland,
NEQ, Plane Crash Site, 1330m, 28 Nov 1990, GM &
HJ, Pyr. - Logs & Trees (ANIC, CBM, QM).

DIAGNOSIS

Medium sized to rather large, depressed, with
an oblique, s-shaped fascia in posterior half of
elytra. Distinguished by distinct transverse im-
pression in anterior third of elytra, narrower base
of pronotum, distinctly s-shaped elytral fascia,
and longer aedeagus with longer apex, and 4-se-
tose parameres.

DESCRIPTION
Measurements. Length: 2.35-2.70mm; width:
1.10-1.18mm. Ratios: Width pronotum/head:
1.44-1.48; width/length of pronotum: 1.42-1.47;
width base/apex of pronotum: 1.45-1.51; width
widest diameter/base of pronotum: 1.07-1.09;
length/width of elytra: 1.41-1.45.
Colour. Dark piceous, pronotum, base and
apex of elytra and posterior part of suture faintly

326

lighter, Elytra in posterior 2/5 with an oblique,
strongly s-shaped, at median end recurrent yellow
fascia not attaining suture. Legs yellow, tibiae
sometimes faintly darker.

Head. Median antennomeres c. 1.75 x as long
as wide, Microreticulation very indistinct and
superficial, about isodiametric, even less distinct
on vertex, Surface glossy,

Pronotum. Fairly depressed, transverse, widest
slightly in front of middle, base much wider than
apex. Lateral margin posteriorly feebly rounded
or even almost straight, though somewhat
oblique. Posterior angles rectangular, faintly pro-
duced over lateral part of base. Carina at postenor
angle elongate, oblique, slightly incurved, Ante-
rior transyerse sulcus rather deep, uninterrupted.
Posterior transverse sulcus very deep, interrupted
by a large fovea. Anterior lateral seta situated at
anterior third of margin. Microreticulation ex-
tremely superficial, indistinct, slightly trans-
verse. Surface highly glossy.

Elytra. Fairly elongate, moderately convex, re-
versely egg-shaped, widest about in middle. Lat-
eral border evenly curved. Disk in anterior third
at position of anterior discal seta with conspicu-
ous, rather deep depression. Sutural stria finely
punctulate. 2nd stria visible as a row of smail
punctures, in anterior half even faintly impressed,
3rd-Sth striae anteriorly more or less well visible
as indistinct rows of extremely fine punctures.
Outer striae almost invisible. Intervals absolutely
depressed, apart from sutural and 2nd in anterior
half. 8th stria deeply impressed, punctate, poste-
riorly punctate-crenulate, attaming postenor
marginal pore. Recurrent striole meeting position
of 3rd stria, Anterior discal pore in anterior third,
posterior pore posterior 2/5. Microreticulation
distinct, though fairly superficial, composed of
very transverse meshes, surface rather glossy.
Wings fully developed.

Lower surface. Metepisternum c. 1.5 x as long
as wide.

Male genitalia, Genital nng slightly asymmet-
rical, apex rather acute. Aedeagus elongate,
rather depressed, lower surface absolutely
straight. Apex short, rounded off. Internal sac in
middle with two areas of microtrichia, Both
parameres 4-setose seta on upper border absent.

Female genitalia, Stylomere 2 with dorsal
ensiform seta situated rather basally.

Variation. Apart from minor differences in rel-
alive shape of pronotum and elytra and degree of
microreticulatron, little variation noted.

MEMOIRS OF THE QUEENSLAND MUSEUM

DISTRIBUTION
Widespread on the Carbine Tableland west of
Mossman, north Queensland.

HABITAT

Rainforest on mountains tops above 1000m by
pyrethrum knockdown on trees, “on trees and
rocks", or "logs and trees", So far collected from
September lo December.

ETYMOLOGY
For Doug Cook, collector of many specimens
of Philipis,

Philipis laevigata sp. nov.
(Figs 4B, 11D, 17C)

MATERIAL EXAMINED

Houoryre: QMT 13520; d, 2km SE Mt. Spurgeon via
Mt. Carbine, МЕО, 20 Dec 1988, 1100m, Monteith &
Thompson, Pyrethrum/Trees & Logs.

PARATYPE: | d, Акт NNE МІ. Spurgeon, NEQ, 15
Oct 1991, GM & HJ, QM Berlesate Nr, 855, 1624'S,
145? 13'Е, RF, 1250m, Sieved litter (CBM).

DIAGNOSIS

Depressed, with an oblique fascia in posterior
half of elytra. Very similar to P. cooki, but
smaller, pronotum narrower, 2nd stria on elytra
deeply impressed and 2nd interval convex,
microreticulation of elytra almost absent,
aedeagus shorter with larger apex, and both
parameres 5-setose.

DESCRIPTION

Measurements. Length: 2.2mm; width: 1.00-
1.02mm. Ratios: Width pronotunvhead; 1.41-
1.44; width/length of pronotum: 1.41-1.45;
width base/apex of pronotum: 1.39; width widest
diameter/base of pronotum: 1.09-1.11;
length/width of elytra: 1.41-1.44.

Colour. Dark piceous, pronotum, base and apex
of elytra and posterior part of suture faintly ligh-
ter. Elytra in posterior 2/5 with a rather inconspic-
uous, moderately well delimited, oblique,
s-shaped, at median end recurrent yellow fascia
attaining about 2nd stria.

Head. Median antennomeres c. 1.5 x as long
as wide. Microreticulation rather distinct, though
somewhat superficial, less distinct on vertex,
about isodiametric. Surface rather glossy,

Pronotum. Fairly depressed, transverse, widest
slightly in front of middle, base much wider than
apex. Lateral rnargin posteriorly feebly rounded,
though somewhat oblique. Posterior angles rec-
tangular, faintly produced over lateral part of

REVISION OF PHILIPIS 327

FIG. 3. Dorsal view of Philipis bicolor зр. nov.

base. Carina at posterior angle elongate, oblique,
slightly incurved. Anterior transverse sulcus
rather deep, uninterrupted. Posterior transverse
sulcus very deep, interrupted by a large fovea.
Anterior lateral seta situated at anterior third of
margin. Microreticulation indistinct, highly su-
perficial, slightly transverse. Surface glossy.
Elytra. Fairly elongate, moderately convex, re-
versely egg-shaped, widest about in middle. Lat-
eral border evenly curved. Disk in anterior third
without transverse depression. Sutural stria finely
punctulate. 2nd stria apparently also rather deeply
impressed (though elytra of type somewhat
creased), finely punctate. 3rd-Sth striae anteriorly
just visible as indistinct rows of extremely fine
punctures. Outer striae almost invisible. Sutural
and 2nd intervals convex, outer intervals abso-
lutely depressed. 8th stria deeply impressed
throughout, almost impunctate, attaining poste-
rior marginal pore. Recurrent striole elongate,

oblique, meeting position of 3rd stria. Anlerior
discal pore in anterior third, posterior pore
slightly in front of posterior third. Microreticula-
tion indistinct, highly superficial, composed of
irregular, very transverse meshes, surface highly
glossy. Wings fully developed,

Lower surface, Metepistemum с. 1.5 x as long
as wide.

Male genitalia. Genital ring asymmetrically
triangular, apex rather obtuse. Aedeagus short
and compact, lower surface straight. Apex short,
wide, widely rounded off. Internal sac in middle
with two areas of microtrichia. Both parameres
5-sctose. Female genitalia unknown.

Variation. Very little variation noted.

DISTRIBUTION
Mt. Spurgeon and vicinity, Carbine Tableland,
north Queensland.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks and logs and by sieving litter in mon-
tane rainforest above 1100m. Collected October
and December.

ETYMOLOGY
Refers to the smooth, glossy elytra.

Philipis heatherae sp. nov.
(Figs 11E, 18A)

MATERIAL EXAMINED
HOLOTYPE: QMT13521; €, 17°03'S x 145"42" E Isley
Hills, МЕО, 1050m 30 Nov 1993 Monteith & Janetzki,
Pyrethrum/irees & rocks,

DIAGNOSIS

Medium-sized, depressed, with an oblique fas-
cia in posterior half of elytra. Distinguished bv
wide base of pronotum that is almost as wide as
diameter of pronotum in middle, by fascia of
elytra not s-shaped, and by 2nd interval not im-
pressed and 3rd-5th intervals indistinct.

DESCRIPTION

Measurements. Length: c. 2.35mm:; width: c.
1.08mm. Ratios: Width pronotum/head: 1.41;
width/length of pronotum: 1.46; width base/apex
of pronoturn: 1,50; width widest diameter/base of
pronotum: 1.03; length/width of elytra: 1.42.

Colour. Dark piceous, suture and lateral mar-
gins of elytra faintly lighter. Elytra in posterior
2/5 with a moderately conspicuous, fairly well
delimited, oblique, faintly s-shaped, at median

end recurrent yellow fascia attaining about 2nd
stria.

Head. Median àntennomeres c, 1.5 x as long as
wide. Microreticulation rather distinct, though
somewhat superficial, less distinct on vertex,
about jsodiametric. Surface rather glossy.

Pronotum. Fairly depressed, transverse, quad-
rate, widest slightly in front of middle, though
barely narrowed to base, base much wider than
apex. Lateral margin posteriorly almost straight
oreven feebly concave, faintly oblique. Posterior
angles rectangular, faintly produced over lateral
part of base. Carina at posterior angle elongate,
oblique, slightly incurved. Anterior transverse
sulcus rather deep, uninterrupted. Posterior trans-
verse sulcus very decp, interrupted by a large
fovea, Anterior lateral seta situated at anterior
third of margin. Disk with some shallow, trans-
verse wrinkles in middle, in anterior two fifth on
either side with a shallow, punctiform impres-
sion. Microrcticulation rather indistinct, superfi-
cial, slightly transverse. Surface glossy.

Elytra. Fairly elongate, moderately convex, re-
versely egg-shaped, widest about in middle, Lat-
eral border evenly curved. Disk in anterior third
with very slight transverse depression, Sutural
stria finely punctulatc. 2nd stria barely im-
pressed, finely punctate. 3rd-5th striae just. visi-
ble as indistinct rows of very spaced and fine
punctures. Outer striae almost invisible, Sutural
interval convex, 2nd interval barely convex, outer
intervals absolutely depressed. Sth stria deeply
impressed throughout, finely punctate, attaining
posterior marginal pore. Recurrent striole elon-
gate, oblique, meeting position of 3rd stria, Ante-
rior discal pore in anterior third, posterior pore
slightly in front of posterior third. Microreticula-
tion indistinct, highly superficial, composed of
irregular, transverse meshes, surface highly
glossy. Wings fully developed. Variation un-
known.

Lower surface. Metepisternum c. 1.5 x as long
as wide.

Genitalia, Male unknown. Female stylomere 2
with dorsal ensiform seta situated rather basally.

DISTRIBUTION
Isley Hills, south of Cairns, north Queensland.
Known only from type locality.

HABITAT

Collected hy pyrethrum knockdown on mossy
trees and rocks in montane rainforest at 1050m.
Captured end of November,

MEMOIRS OF THE QUEENSLAND MUSEUM

ETYMOLOGY
For Heather Janetzki, Queensland Museum,
collector of many species of Philipis.

Philipis picea sp. nov.
(Figs HF, 18A)

MATERIAL EXAMINED

HOLOTYPE: QMT13522; 9, Mossman Bluff Track,
9km W Mossman, NEQ, 20 Dec 1989, 860m, Monteith
& Thompson, Pyrethrum (Site 6).

DIAGNOSIS

Medium sized, rather elongate and fairly de-
pressed, winged. Easily distinguished by de-
pressed shape and uniform, piceous colour
without any elytral pattern. Distinguished from Р.
castanea by wide base of pronotum and absence
of transverse depression in anterior third of elytra.

DESCRIPTION

Measurements. Length: 2.50mm; width:
l.16mm, Ratios: Width pronotum/head: 1.40;
width/length of pronotum: 1.51; width base/apex
of pronotum: 1.57; width widest diameter/base of
pronotum: 1,01; length/width of elytra: 1.37.

Colour. Very dark piceous. lateral borders of
pronotum and elytra and elytral suture posteriorly
faintly lighter. Legs dark yellow, femora slightly
infuscate.

Head, Median antennomeres c, 1.6 x as long
as wide. Microreticulation on labrum, clypeus,
and frons fairly distinct, though somewhat super-
ficial, on vertex reduced, about isodiametric, Sur-
face moderately glossy.

Pronotum. Fairly depressed, transverse, widest
in front of middle, base much wider than apex,
Lateral margin posteriorly very faintly concave,
barely oblique, hence pronotum not perceptibly
narrowed to posterior angles. Posterior angles
rectangular. Carina at posterior angle elongate,
oblique, slightly incurved, Anterior transverse
sulcus comparatively shallow, uninterrupted.
Posterior transverse sulcus very deep, interrupted
by a rather large fovea. Anterior lateral seta situ-
ated at anterior third of margin. Microreticulation
absent, surface highly glossy.

Elytra, Fairly elongate, moderately convex, re-
versely egg-shaped, widest slightly in front of
middle. Lateral border evenly curved. Disk with-
Our transverse depression, Sutural stria finely
punctulate. All other striae visible as rows of
rather fine punctures, though becoming finer lat-
erally and towards apex. 2nd stria in anterior half
faintly impressed. Only sutural interval convex,
others depressed. 8th stria deeply impressed,

REVISION OF PHILIPIS 329

FIG. 4. d genitalia. A, Philipis cooki sp. nov. B, P. laevigata sp. nov. C, P. castanea sp. nov. D, P. unicolor sp.
nov. E, P. atra sp. nov. F, P. ruficollis sp. nov.

330 MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1. Summary of measures and ratios of Philipis. 1: length (mm); w: width (mm); w p/h: ratio of width
pronotum/head; w/l p: ratio of width/length of pronotum; b/a p: ratio of width of base/apex of pronotum; d/b p:
ratio of widest diameter/base of pronotum; I/w e: ratio of length/width of elytra.
ee eee eee
| 099 |
lea — | 275 | am | im | з | її | иш | 145 |
CERE че ми аш es
шшш | is ue ы с лш ы ү
picea — | 250 | 11 | 14 | 1s | їз | ım | 137 |
teen ГУП auae | xus | labial | Ims т | CUM |
PIPEMELICNEIIURNUTCHECTCEETICENIIEENETE
eris — | 195299 | 389090 | пазам anas [iaar | 10940 | 136138 |
1.03-1.07
0.97-0.98
[155 | 107 | 130 |
iss Lo | 151-156 | 151-153 | 13&144
092-098 | 148-L54 | 150-153 | 140-142
thompsoni Tisis | ооо | aas | зва лвла [bein | зын)
vicina Lim [om [oam oam [oie oam | 133
erei | 20s2as | 100115 | 1o16 | iasisi iata onas | iaia |
jsubrropica | 230-280 | 100125 | 154164 | 142-147 | 143-47 | 1.132115 | 135-136 |
bars. | 225245 | 1л? | restos | ва | 1816). 10106 | 2135 |
16-63 | 1.36-1.45 [rastar | азаа [ raras |
bus | nm | we | ы ie p iar [ы ан
135-14] |
жн Зама “eed ase мые | шше ныч
Wem [миё [ав [| ыз [мз мб ы [ 4 ]
sinuata | 260275 | 120-130 | 165-173 | 151156 | 15159 | 109-101 | 1341.36 |
[lustrans — | 220-250 | 100-116 | 156162 | 151157 | 1д6151 | 109113 | 1334137 |
finely punctate, becoming shallower ànteriorly, Lower surface. Metepisternum elongate, c. 1.5
attaining posterior marginal pore. Recurrent * Cs dem pi n ныт и
, , i à enitalia. Male unknown. Female stylomere
striole elongate, oblique, meeting 3rd stria. Ante- with dorsal ensiform seta situated ик» basally.
rior discal pore in anterior third, posterior pore
slightly in front of posterior 2/5. Microrenicula- DISTRIBUTION
tion absent, surface highly glossy. Wings fully Mountains west of Mossman, north Queens-
developed. Variation unknown. land. Known so far from type locality only.

REVISION OF PHILIPIS

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks in montane rainforest at 850m. Col-
lected December only,

ETYMOLOGY
Refers to the piceous colour.

Philipis castanea sp. nov.
(Figs 4C, 12A, 18A)

MATERIAL EXAMINED

HOLOTYPE: QMT13523; 4, Thornton Peak summit,
via Daintree, NEQ, 24-27 Sept 1984, G. B. & S. R.
Monteith/QM Berleseate No. 662, Rainforest, 1100-
1300m, sieved litter & moss.

PARATYPES: 2 d, 4 9, Thornton Peak, 11km NE
Daintree, NEQ, 1000-1200m, 30 Oct-1 Nov 1983, GM,
DY & GT (ANIC, CBM, QM, ZSM); 2 d, 4 9,
Thornton Peak via Daintree, МЕО, 1000-1300m, 20-22
Sept 1981, GM & DC (CBM, QM, USNM), 1 8, ,
4.5-5.0km W of Cape Tribulation, Top Camp, МЕО,
760-780m, 27 Sept-7 Oct 1982, GM. DY & GT (QM);
2 d, | 9, 4.5-5.0km W of Cape Tribulation, Top
Camp, NEQ, 1-6 Oct 1982, 760-780m, GM, DY & GT,
Pyr., RF (QM); І 9,, 5.0km W of Cape Tribulation,
NEQ, (Site 10), 28 Sept 1982, 780m, GM, DY & GT,
Pyr., RF (QM).

DIAGNOSIS

Medium sized, rather elongate and fairly de-
pressed, winged. Easily distinguished by de-
pressed shape and uniform, chestnut brown
colour without any elytral pattern. Distinguished
from P. picea by narrower base of pronotum and
presence of a transverse depression in anterior
third of elytra.

DESCRIPTION

Measurements. Length: 2.15-2.50mm; width:
0,95-1.15mm. Ratios: Width pronotum/head-
1.40-1.45; width/length of pronotum: 1.48-1.51;
width base/apex of pronotum: 1,50-1.53; width
widest diameter/base of pronotum: 1.05-1.07;
length/width of elytra: 1.36-1.38.

Colour. Chestnut brown, head piceous, elytral
suture posteriorly faintly lighter. Legs yellow,
tibiae sometimes faintly darker.

Head. Median antennomeres c. 1.75 x as long
as wide. Microreticulation present on labrum,
clypeus, and frons, but very indistinct and super-
ficial, about isodiametric, visible only under high
magnification. Surface glossy.

Pronotum. Fairly depressed, transverse, widest
in front of middle, base much wider than apex.
Lateral margin posteriorly feebly rounded or
even almost straight, though somewhat oblique.

331

Posterior angles rectangular. Carina at posterior
angle elongate, oblique, slightly incurved. Ante-
rior transverse sulcus comparatively shallow, un-
inlerrupted. Posterior transverse sulcus very
deep, interrupted by arather large fovea. Anterior
lateral seta situated at anterior third of margin.
Microreticulation absent, surface highly glossy.

Elyrra. Fairly elongate, moderately convex, re-
versely egg-shaped, widest slightly in front of
middle. Lateral border evenly curved. Disk in
anterior third at position of anterior discal seta
with shallow transverse depression. Sutural stria
punctulate. 2nd siria visible as a row of small
punctures, in anterior half even faintly impresed,
3rd-5th striae anteriorly more or less well visible
as indistinct rows of extremely fine punctures.
Outer striae absent. Intervals, apart from sutural
and 2nd in anterior half, absolutely depressed. 8th
stria deeply impressed, punctate, posteriorly
punctate-crenulale, attaining posterior marginal
pore. Recurrent striole clongate, oblique, meet-
ing position of 3rd stria. Anterior discal pore in
anterior third, posterior pore behind middle.
Microreticulation absent, except for the very
apex, surface highly glossy. Wings fully devel-
oped.

Lower surface. Metepisternum c. 1.5 x as long
as wide.

Male genitalia, Genital ring regularly triangu-
lar, apex rather acute, Aedeagus large, lower sur-
face slightly bisinuate, apex stout, elongate,
slightly bent down, at tip rounded off. Both
parameres 5-setose,

Female genitalia. Stylomere 2 with dorsal
ensiform seta situated rather basally.

Variation, Apart from some variation of shape
of pronotum and distinctness of elytral striae,
little variation noted.

DISTRIBUTION

Thornton Peak and adjacent uplands above
Cape Tribulation, north of Daintree, north
Queensland,

HABITAT

A montane species, occurring in rainforest
above 750m. Collected by pyrethrum knockdown
on mossy tree trunks and by Berlese extraction of
litter and moss. Collected only in the period of
September-November.

ETYMOLOGY
Refers to the castaneous colour of upper siir-
face.

332
Philipis unicolor sp. nov.
(Figs 4D, 12B, 18A)
MATERIAL EXAMINED

HOLOTYPE: QMT13536; d , Bellenden Ker, Cableway
Tower No 3, МЕО, 1000m. Pyrethrum/trees, 25 Sept
1981, G. Monteith.

PARATYPES: 2 3, 10 9, same data (ANIC, CBM, OM,
USNM, ZSM); 14,5 9, Bellenden Ker Range, Cable
Tower 3, NEQ, 1054m, 17 Oct-5 Nov 1981, EW/OM,
Pyr. (CBM, QM); 1 9, Bellenden Ker Range, Cable
Tower 3, NEQ, 1054m, 17-24 Oct 1981, EW/OM, Pyr.
on logs, stones and tree trunks (QM); 1 9, Bartle-
Frere, NW Peak, NEQ, 1440m, Pyr. on mossy rocks 24
Sept 1981, GM (QM); 2 5, Mt. Bartle Frere, МЕО,
West Side, 1050m, 8 Dec 1990, GM, GT & RS, Pyr.-
trees & rocks (QM); 1 9, Bell Peak North 10km E
Gordonvale, NEQ, В a 1982, 900-1000m, GM, DY
& GT, Pyr., RF (QM

DIAGNOSIS

Small, piceous, unpatterned, short and highly
convex, distinguished from Р, atra sp. nov. by
smaller size (length), lighter colour, evenly con-
vex posterior part of lateral margin of pronotum,
shorter elytra, and shorter and wider apex and
evenly rounded lower surface of aedeagus.

DESCRIPTION

Measurements. Length: 1.85-2.25mm; width:
0.9-1.1mm. Ratios: Width pronotum/head: 1.54-
1.58; width/length of pronotum: 1.56-1.60; width
base/apex of pronotum: 1.45-1.50; width widest
diameter/base of pronotum: 1.10-1.13;
length/width of elytra: 1.28-1.30.

Colour. Piceous, sutural interval and marginal
channel of elytra reddish, Elytra without pattern.
Antenna yellow. Legs yellow, femora reddish to
reddish-brown.

Head. Median antennomeres c. 1.35 x as long
as wide, Microreticulation on frons barely visi-
ble, on labrum and anterior border of clypeus
distinct, isodiametric. Surface glossy.

Pronotum. Moderately convex, transverse,
widest in middle, base much wider than apex,
Lateral margin evenly curved, though slightly
more curved to apex than to base, Posterior angles
rectangular, faintly produced over lateral part of
base. Carina at posterior angle elongate, oblique,
slightly incurved, Anterior transverse sulcus
comparatively shallow, uninterrupted. Posterior
transverse sulcus decp, interrupted by a large
fovea. Anterior lateral seta situated slightly be-
hind anterior third of margin. Microreticulation
absent, surface highly glossy.

Elytra. Short and highly convex, lateral border
evenly curved, widest in middle. Lateral margin

MEMOIRS OF THE QUEENSLAND MUSEUM

*

extremely finely serrrate and pilose. Sutural stria
in anterior half coarsely punctate, posteriorly im-
punctate, 2nd stria in basal half indicated as a row
of rather coarse punctures, but stria beginnin ng
only at some distance from base, posteriorly a
most reduced, Outer striae composed of ex-
tremely fine rows of delicate punctures,
posteriorly almost reduced, difficult to recognize.
Only sutural interval and 2nd interval in basal
third, slightly convex. 8th stria deeply impressed,
loosely punctate, not attaining postenor marginal
pore. Recurrent striole rather short, markedly
oblique, only anteriorly shortly incurved and
hardly meeting position of 3rd stria. Anterior
discal pore in anterior third, posterior pore
slightly behind middle. Microreticulation absent,
surface highly glossy. Wings absent.

Lower surface. Metepisternum about as long as
wide,

Male genitalia. Genital ring almost symmetri-
cal, apex rather wide, Aedeagus short and com-
pact, lower surface slightly and evenly curved.
Apex short, thick, rounded off. Both parameres
5-sctose.

Female genitalia. Stylomere 2 with dorsal en-
siform seta situated rather basally.

Variation. Apart from some, partly sexual, dif-
ferences of size, and some differences of relative
shape of pronotum, little variation noted.

DISTRIBUTION

Bellenden Ker Range and nearby Malbon
Thompson Range, east of Atherton Tableland,
north Queensland,

HABITAT

Collected by pyrethrum knockdown of mossy
trees and rocks in montane rainforest above
900m, in the period of September to November.

ETYMOLOGY
Refers to the unpatterned dorsal surface.

Philipis atra sp, nov.
(Figs 4E, 12C, 17A)

MATERIAL EXAMINED

HOLOTYPE: QMT13553; 8, 20750'S, 148"34'E Mt.
Macartney, CEQ, 19 Nov 1992, 900m Monteith,
Thompson & Janetzki, Pyrethrum.

PARATYPES: 2 9, same data (СВМ, QM).

DIAGNOSIS

Medium sized, black, unpatterned, short and
highly convex, distinguished from P. unicolor sp.
nov. by larger size (length 2.45mm), darker col-

REVISION OF PHILIPIS 333

FIG. 5. d genitalia. A, Philipis inermis sp. nov. B, P. striata sp. nov. C, P. planicola sp. nov. D, P. trunci
(Darlington). E, P. tribulationis sp. nov. F, P. thompsoni sp. nov.

334

our, irregularly convex posterior part of lateral
margin of pronotum, longer elytra, and longer
and narrower apex and basally straight lower
surface of aedeagus.

DESCRIPTION

Measurements. Length: 2.45-2.60mm; width:
1.12-1.20mm. Ratios: Width pronotum/head:
1,55-1,58; width/length of pronotum: 1.49-1.53;
width base/apex of pronotum: 1.39-1.43; width
widest diameter/base of pronotum: 1,11-1,13;
length/width of elytra: 1.37-1.38.

Colour. Completely black or very dark pice-
ous-black, Elytra without pattern. Legs dirty yel-
low, femora barcly darker.

Head. Median antennomeres slightly «1.5 x as
long as wide. Microreticulation on frons rather
superficial, on labrum and anterior border of
clypeus distinct, isodiametric, Surface glossy.

Pronotum. Moderately convex, transverse,
widest in middle, base much wider than apex.
Lateral margin anteriorly evenly curved, though
posteriorly with a faint compression behind mid-
dly, and near basal angles curved in, Posterior
angles rectangular, at apex obtuse, not produced
over lateral part of base. Carina at posterior angle
elongate, oblique, slightly incurved. Anterior
transverse sulcus comparatively shallow, unin-
terrupted. Posterior transverse sulcus deep, inter-
rupted by à large fovea. Anterior lateral seta
situated slightly behind anterior third of margin.
Microreticulation absent, surface highly glossy.

Elytra. Rather short and highly convex, egg-
shaped, widest in middle, Lateral border evenly
curved, extremely finely serrate and pilose. Su-
tural stria in anterior half finely punctate, poste-
попу impunctate, 2nd stria in basal half slightly
impressed, finely punciate, but stria beginning
only at some distance from base, becoming
weaker towards apex. Outer striae gradually finer
towards lateral margin, striae composed of rows
of fine punctures, the ouler ones posteriorly al-
most reduced, difficult to recognize. Only sutural
interval and 2nd interval in basal half, slightly
convex. Sth stria deeply impressed, loosely punc-
tate, just attaining posterior marginal porc. Re-
current striole rather short, markedly oblique,
anteriorly barely incurved, not attaining position
of 3rd stria. Anterior discal pore situated in ante-
rior third, posterior pore in posterior third,
Microreticulation absent, except for near apex,
surface highly glossy. Wings present.

Lawer surface, Metepistemum slightly longer
than wie.

MEMOIRS OF THE QUEENSLAND MUSEUM

Male genitalia. Genital ring almost symmetri-
cal, apex rather wide. Aedeagus short and com-
pact, lower surface basally straight, then evenly
curved. Apex short, rather compact, rounded off.
Both parameres 5-setose.

Female genitalia. Stylomere 2 with very stout
dorsal ensiform seta situated about medially.

Variation. Apart from some minor differences
in size and relative shape of pronotum, little vart-
ation noted.

DISTRIBUTION

Mt. Macartney north of Eungella National
Park, central eastern Queensland. Known only
from this mountain top,

HABITAT

Collected in montane rainforest above 900m
by pyrethrum knockdown, on mossy tree trunks,
Collected November only.

ETYMOLOGY
Refers to the unicolorous black surface.

Philipis ruficollis sp. nov.
(Figs 4F, 12D, 18B)

MATERIAL EXAMINED
HOLOTYPE; QMT13555; d, Mt. Finnigan, 850-
1100m, 37km S Cooktown, NEQ, 19-22 Apr 1982, RF
Monteith, Yeates & Cook.
PARATYPES: | 5, same data (CBM); 1 9 Mt, Finnigan,
1100m, 37km S Cooktown, NEQ, 20 Dec 1982, LR,
n (QM); 1 3, Mt, Finnigan Summit via Helenvale,
EQ, 3-5 Dec 1990, 1050m, GM, GT, DC, RS & LR
(QM); 1 3, Mt. Finnigan Summit via Helenvale, NEQ,
28-30 Nov 1985, 1100m, GM, DC & LR (QM), | d,
Mt. Finnigan Summit via Helenvale, NEQ, 28 Nov
1985, 1100m, GM & DC, Pyr/RF (QM); 2 3, Mt.
Finnigan Summit via Helenvale, NEQ, 3-5 Dec 1990,
1050m, GM, RS, LR & GT, Рут. (ANIC, ОМ),

DIAGNOSIS

Medium sized, moderately convex, distin-
guished by rufous pronotum, almost transverse
elytral fascia not s-shaped, and strongly bisinuate
lower surface of acdeagus.

DESCRIPTION

Measurements. Length: 2.35-2.50mm; width:
1.12-1.15mm. Ratios; Width pronotum/head:
1.43-1.44; width/length of pronotum: 1.42-1.44;
width base/apex of pronotum: 1.47-1,49; width
widest diameter/base of pronotum: 1.06-1.08;
Jength/width of elytra; 1,37-1,41.

Colour, Head and elytra dark piceous, prono-
tum contrastingly reddish. Elytra with an ill-de-

REVISION OF PHILIPIS 335

FIG. 6. ó genitalia. A, Philipis vicina sp. nov. B, P. laevis sp. nov. C, P. sulcata sp. nov. D, P. frerei sp. nov. E,
P. subtropica sp. nov. F, P. alutacea sp. nov,

336

fined, somewhat transverse spot in posterior 2/5
not reaching suture nor lateral border, and suture.
posteriorly reddish. Legs yellow.

Head. Median antennomeres c. 1.3 x as long as
wide. Microreticulation visible on frons, though
rather superficial, almost reduced on vertex,
about isodiametric. Surface rather glossy.

Pronotum. Rather depressed, transverse, wid-
est slightly in front of middle, base much wider
than apex. Lateral margin posteriorly almost
straight, though somewhat oblique. Posterior an-
gles rectangular, feebly produced over lateral part
of base. Marginal Lateral channel posteriorly
slightly punctate, Carina at posterior angle elon-
gate, oblique, slightly incurved. Anterior trans-
verse sulcus rather shallow, slightly interrupied
or at least becoming shallower. Posterior trans-
verse suleus very deep, interrupted by a rather
small, inconspicuous fovea. Anterior lateral seta
situated slightly behind anterior third nf margin.
Microreticulation completely absent, surface
highly glossy.

Elytra. Moderately elongate, fairly convex, re-
versely egg-shaped, widest well in front of mid-
dle. Lateral border evenly curved. Sutural stria
crenulate. Other striae in anterior half well devel-
oped as rows of fairly coarse punctures, though
6th and 7th striae very delicate. All striae poste-
riorly reduced, Only sutural interval convex. 8th
stria deeply impressed, punctate, attaining poste-
rior marginal pore. Recurrent striole clongate,
meeting position of 3rd stria. Anterior discal pore
in anterior third, posterior pore slightly behind
posterior 2/5. Microreticulation on disk com-
pletely absent, but present ах superficial rans-
verse meshes at apex, Surface of disk highly
glossy. Wings fully developed. Little variation
noted.

Lower surface, Metepisternum с, 1.3 x as long
as wide.

Male genitalia. Genital ring triangular, slightly
asymmetrical, apex rather acute. Aedeagus rather
short and compact, lower surface remarkably
bisinuate, apex slightly bent down, short, widely
rounded off. Both parameres 5-setose,

Female genitalia, Stylomere 2 with dorsal
ensiform seta situated rather medially.

DISTRIBUTION

Mt. Finnigan south of Cooktown, north
Queensland, Known only from that mountain top.

MEMOIRS OF THE QUEENSLAND MUSEUM

HARITAT

Collected by pyrethrum knockdown on mossy
tree trunks in montane rainforest above 850m.
Collected April and November-December.

ETYMOLOGY
Refers to the red colour of pronotum.

Philipis inermis sp. nov.
(Figs 5A, L2bE, IBB)

MATERIAL EXAMINED

HOLOTYPE: QMT13561; d, Mt. Finnigan Surmmit,
NEQ, 29 Nov 1985, G. Monteith, QM Berlesate No.
699, 15"48S 145°17E, moss on trees.

PARATYPES: | d, Mt. Finnigan, 1 100m, 37km 5 Cook-
town, NEQ, 20 Dec 1982, LR, Pyr. (CBM),

DIAGNOSIS

Small, narrow. distinguished by small size.
narrow pronotum, and apical elytral fascia not
s-shaped,

DESCRIPTION

Measurements. Length: 1,95-2.0mm; width:
0.89-0,90mm. Ratios: Width pronotum/head:
1.43-1.44; width/length of pronotum; 1.32-1.34;
width base/apex of pronotum: 1.31-1.33; width
widest diameter/base of pronotum: 1.09-1.10;
length/width of elytra: 1.36-1.38.

Colour, Very dark piceous to almost black,
pronotum faintly lighter, posterior half of elytral
suture reddish. Elytra with a moderately well-de-
fined, slightly transverse, dark yellow or light
reddish spot in posterior 2/5 medially reaching
almost to 2nd stria. Legs reddish-piceous, apex of
femora and tibiae, and tarsi yellow. Lower sur-
face piceous, anteriorly even slightly darker.

Head. Median antennomeres с, 1,15 x as long
as Wide, Microreticulation distinct on labrum and
clypeus, becoming more superficial on frons, al-
most absent on vertex, anteriorly about isodia-
metric, posteriorly slightly transverse. Surface
fairly glossy.

Pronotum. Comparatively narrow, convex,
widest about in middle, base not much wider than
apex. Lateral margin evenly curved, but slightly
less so posteriorly. Posterior angles subrectangu-
lar, slightly obtuse, not produced over lateral part
of base. Carina at posterior angle comparatively
short, oblique, slightly incurved, Anterior trans-
verse sulcus moderately deep, barely interrupted.
Posterior transverse sulcus deep, slightly inter-
rupted, Anterior lateral seta situated well behind
anterior third of margin. Microreticulation super-

REVISION OF PHILIPIS 337

Fig. 7. д genitalia. A, Philipis rufescens sp. nov. B, P. ellioti sp. nov. C, P. agnicapitis sp. nov. D, P. picta sp.
nov. E, P. bicolor sp. nov. F, P. geoffreyi sp. nov.

338

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 8. & genitalia. A, Philipis minor sp. nov. B, P. sinuata sp. nov. C, P. distinguenda sp. nov. D, P. lustrans

sp. nov.

ficial, though visible, slightly transverse, surface
fairly glossy.

Elytra. Short, convex, egg-shaped, widest
about in middle. Lateral border evenly curved,
finely serrate and pilose throughout. Sutural stria
in basal half punctate, anteriorly barely recurved
and not deepened. 2nd and 3rd striae near base
indicated as rows of moderately fine punctures,
reduced posteriorly. Outer striae almost invisible,
at most 4th and 5th vaguely indicated near base.
Only sutural stria well marked to apex. Only
sutural interval and 2nd interval in basal half
somewhat convex. 8th stria posteriorly deeply
impressed, punctate, becoming weaker anteri-
orly, not attaining posterior marginal pore. Re-

current striole markedly oblique, meeting posi-
tion of 3rd stria. Ridge laterally bordering recur-
rent striole markedly carinate. Anterior discal
pore in anterior third, posterior pore slightly in
front of posterior third. Microreticulation dis-
tinct, rather coarse, though slightly superficial,
composed of moderately transverse meshes. Sur-
face but moderately glossy. Wings slightly short-
ened. No perceptible variation noted.

Lower surface. Metepisternum barely longer
than wide.

Genitalia. Male genital ring regularly triangu-
lar, narrow, slightly asymmetrical, apex rather
wide. Aedeagus short and compact, lower surface
straight, apex very short, widely rounded off.

REVISION OF PHILIPIS

Parameres moderately elongate, both 5-setose.
Female genitalia unknown.

DISTRIBUTION
Known only from mountain top, Mt. Finnigan
south of Cooktown, north Queensland.

HABITAT

Collected by pyrethrum knockdown and Berl-
ese extraction of mossy tree trunks iñ montane
rainforest above 1100m. Collected November
and December,

ETYMOLOGY !
Refers to the small size and narrow shape.

Philipis reticulata sp. nov,
(Figs 12F, 18B)

MATERIAL EXAMINED

HOLOTYPE: QMTI3562; 9, Bellenden Ker Range,
NEQ, Cable Tower 3, 1054m, 17-24 Oct 1981, Earth-
watch/Old, Museum.

DIAGNOSIS

Small, convex, with s-shaped fascia in poste-
rior half of elytra. Distinguished by well devel-
oped striae and presence of distinct
microreticulation on elytra.

DESCRIPTION

Measurements. Length: 2.20mm; width:
1.02mm. Ratios: Width pronotum/head: 1.45;
width/length of pronotum: 1.52; width base/apex
of pronotum: 1.45; width widest diameter/base of
pronotum: 1.07; length/width of elytra; 1,32.

Colour. Piceous, base and apex of elytra faintly
lighter. Elytra with a well delimited, conspicuous,
oblique, rather s-shaped, yellow fascia in poste-
rior 2/5 medially slightly surpassing 2nd stria.
Legs yellow.

Head, Frons medially of facial sulcus on either
side with two distinct, rather deep, circular im-
pressions. Facial sulci on clypeal suture with a
conspicuous circular groove. Median an-
tennomeres c. 1.2 x as long as wide.
Microreticulation distinct on labrum and whole
clypeus, more superficial, though still distinct on
frons, reduced on vertex, about isodiametric to
slightly transverse. Surface moderately glossy.

Pronotum. Transverse, convex, widest about in
middle, base much wider than apex. Lateral mar-
gin posteriorly almost straight, even very faintly
concave in front of posterior angles, slightly
oblique. Posterior angles rectangular, feebly pro-
duced over lateral part of base, Carina at posterior

339

angle elongate, markedly oblique, slightly in-
curyed. Anterior transverse sulcus rather deep,
barely interrupted. Posterior transverse sulcus
very deep, interrupted by a large fovea. Anterior
lateral seta situated well behind anterior third of
margin. Microreticulation absent on disk, though
still present on base and apex, surface glossy,

Elytra. Short, highly convex, egg-shaped, wid-
estabout in middle. Lateral border evenly curved,
extremely finely serrate and pilose. Sutural stria
crenulate. All other striae well impressed and
coarsely punctate at least in basal half, visible
right to apex. Almost all intervals convex, though
lateral intervals less so than in middle. 8th stria
deeply impressed throughout, strongly punctate-
crenulate, attaining posterior marginal pore with-
out becoming shallower. Recurrent striole rather
oblique, meeting end of 3rd stria. Anterior discal
pore in anterior third, posterior pore slightly in
front of posterior third. Microreticulation distinct
and conspicuous, composed of rather irregular,
moderately transverse meshes. Surface rather
duli. Wings almost fully developed. Variation
unknown,

Lower surface. Metepisternum slightly longer
than wide.

Genitalia, Male unknown. Female stylomere 2
with very stout dorsal ensiform seta situated
rather basally,

DISTRIBUTION
Bellenden Ker Range, north Queensland
Known only from type locality,

HABITAT

Collected by pyrethrum küockdown on mossy
tree trunks in montane rainforest at 1050m, Col-
lected October,

ETYMOLOGY
Refers to the markedly microreticulate surface
of elytra.

Philipis striata sp. nov.
(Figs 5B, 13A, 18B)

MATERIAL EXAMINED
HoLotyPe: QMT 13563; g, 1,5km W of Cape Tribu:
lation {She 3), NEQ, 7 Oct 1982, 150m, Monteith,

Yeates & Thompson, Pyrethrum knockdown, RF,
PARATYPES: 3 ©, same data (CBM, QM); 1 d, Moss-
man Bluff Track, 8km W Mossman, NEQ, 20 Dec
1989, 600m, GM & GT, Pyr. (Site 4) (QM).

quadraticallis
Y alticola
19 3 cooki
laevigata
= heatheras
? picea

IET
7

ue castanales
unicolor
ага
ruficollis
inermis
J reticulata
striata
spurgeani
planicola
thompsoni
vicina
frunai
г tribulationis
laevis
2? sulcata
? тегә
таг subtropica

LE dau
1 нё"
33s 13b p!

108 12^ 27

1 We т

alutacea
rufescens
berstriata
ellioti
L3 - agnicaptius
рїс{г

"n bicalor
4 gaotirayi
miner
sinuata
7 distinguenda

19 lustrans

18^ 6 19125

1s f"

FIG. 9, Cladogram of the supposed relationships of the
species of genus Philipis. Nurnbers of synapomorph-
ies referto Tables 1 and 2, Different apomorphic states
distinguished by lower case letters. States of a
morphocline indicated by; ', '', '"". Comparable or
similar apomorphic states that are likely to have been
convergently evolved marked by numbers: !, 2 etc.

DIAGNOSIS

Medium sized, convex, with s-shaped fascia in
posterior half of elytra. Distinguished by well
developed outer striae but depressed intervals,
and by weak microreticulation of elytra.

DESCRIPTION

Measurements, Length; 2,15-2.45mm; width:
1,0-1.15mm. Ratios: Width pronotum/head:
1.50-1.53; width/length of pronotum: 1.56-1.58;
width base/apex of pronotum: 1.50-1.54; width

MEMOIRS OF THE QUEENSLAND MUSEUM

widest diameter/base of pronotum: 1.03-1.07;
length/width of elytra: 1.32-1.34.

Colour. Dark piceous, pronotum, base to ante-
rior third or half, and apex of elytra slightly
lighter, posterior half of suture even lighter, red-
dish-piceous. Elytra with a well-defined, con-
trasting, oblique, s-shaped, dark yellow or light
reddish fascia in posterior 2/5 medially reaching
2nd stria. Legs yellow.

Head. Median antennomeres c. 1.3 x as long as
wide. Microreticulation distinct on labrum and
anterior border of clypeus, superficial and rather
difficult to see on frons, absent on vertex, about
tsodiametric. Surface fairly glossy.

Pronotum. Transverse, convex, widest about
in middle, base much wider than apex. Lateral
Margin posteriorly almost straight, slightly
oblique. Posterior angles rectangular, feebly pro-
duced over lateral part of base. Carina at posterior
angle elongate, markedly oblique, slightly in-
curved. Anterior transverse sulcus rather deep,
barely interrupted. Posterior transverse sulcus
very deep, interrupted by a large, deep fovea.
Anterior lateral seta situated slightly behind an-
terior third of margin. Microreticulation com-
pletely absent, surface highly glossy.

Elvtra. Short, highly convex, egg-shaped, wid-
est about in middle, Lateral border evenly curved,
extremely finely serrrate and pilose. Sutural stria
crenulate. Other striae well developed as rows of
coarse punctures, laterally and posteriorly not
perceptibly finer, 2nd-5th striae in anterior half
even slightly impressed. All striae well marked to
apex. Sutural interval and basal part of 2nd and
even 3rd intervals somewhat convex. Sth stria
deeply impressed, punctate-crenulate, barely at-
taining posterior marginal pore. Recurrent striale
rather evenly curved, meeting end of 3rd stria.
Anterior discal pore in anterior third, posterior
pore slightly in front of posterior third.
Microreticulation on disk very superficial,
though well visible when seen from laterally or
posteriorly, composed of transverse meshes. Sur-
face rather glossy. Wings slightly shortened.

Lower surface. Metepisternum slightly longer
than wide.

Male genitalia. Genital ring regularly triangu-
lar, slightly asymmetrical, apex rather elongate.
Aedeagus rather short and compact, lower sur-
face fcebly curved, apex perceptibly bent down
short, widely rounded off. Parameres remarkably
short, both 5-setose.

Female genitalia. Stylomere 2 with dorsal en-
siform seta situated rather medially,

REVISION OF PHILIPIS

Variation. Apart from some differences of size
and of shape of pronotum, little variation noted.

DISTRIBUTION

Lowlands at Cape Tribulation and foothills of
Carbine Tableland west of Mossman, north
Queensland.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks in rainforest of low and middle eleva-
tion below 600m. Collected October and Decem-
ber.

ETYMOLOGY
Refers to the complete striation of elytra.

Philipis planicola sp. nov.
(Figs 5C, 13B, 18C)

MATERIAL EXAMINED

HOLOTYPE: QMT13567; d, Bellenden Ker Range,
NEQ Cableway Base Stn, 100m, 17 Oct-9 Nov 1981,
Earthwatch/Old. Museum, pyrethrum knockdown.
PARATYPES: ! d, 1 9, Russell R. at Bellenden Ker
Landing, NEQ, 5m, 1 Nov 1981, EW/QM, QM Berles-
ale No. 361, 17^ 16'S, 145°57°E, palm swamp, moss on
tree tranks (CBM, QM),

DIAGNOSIS

Small, convex, with s-shaped fascia in poste-
rior half of elytra. Distinguished species by wide
base and straight lateral borders of pronotum,

DESCRIPTION

Measurements. Length: 2.1-2.2mm; width:
1,0-1.05mm. Ratios: Width pronotum/head:
1.48-1.49; width/length of pronotum: 1.50-1.54;
width base/apex of pronotum: 1.54-1.55; width
widest diameter/base of pronotum: 0.97-0.98;
length/width of elytra: 1.30- 1.31,

Colour. Head and and posterior 2/3 of elytra
dark piccous, pronotum, anterior third of elytra,
suture, and apex slightly lighter, piceous to red-
dish-piceous. Elytra with a well-defined, con-
trasting, oblique, s-shaped, dark yellow or light
reddish fascia in posterior 2/5 medially reaching
2nd stria. Sometimes the median, slightly wid-
ened part of the fascia isolated. Legs yellowish.

Head. Median antennomeres c. 1.3 x as long
as wide. Microreticulation absent except from
labrum and anterior border of clypeus, there
about isodiametric, Surface glossy

Pronotum. Moderately transverse, markedly
convex, widest immediately at base, base much
wider than apex. Lateral margin strongly and

evenly curved, posteriorly straight, not oblique.
Posterior angles rectangular, feebly produced
over lateral part of base. Carina at posterior angle
elongate, oblique, slightly incurved, markedly
raised. Anterior transverse sulcus deep, slightly
interrupted by a fovea. Posterior transverse sulcus
very deep, interrupted by a very large, deep fovea.
Anterior lateral seta situated slightly behind an-
terior third of margin. Microreticulation com-
pletely absent, surface highly glossy.

Elytra. Short, highly convex, egg-shaped, wid-
est about in middle. Lateral border evenly curved,
extremely finely serrrate and pilose. Sutural stria
crenulate. Other striae well developed as rows of
fairly coarse punctures, though laterally becom-
ing finer. 2nd and 3rd striae in anterior half even
faintly impressed. All striae traceable to apex.
Sutural interval and basal part of 2nd interval
somewhat convex. 8th stria deeply impressed
throughout, punctate, surpassing posterior mar-
ginal pore, Recurrent striole rather evenly curved,
meeting end of 3rd stria. Anterior discal pore in
anterior third, posterior pore slightly in front of
posterior third. Microreticulation completely ab-
sent, surface highly glossy. Wings slightly short-
ened, Little variation noted.

Lower surface. Metepisternum slightly longer
than wide.

Male genitalia. Genital ring triangular, slightly
asymmetrical, apex rather elongate, Aedeagus
rather short and compact, lower surface feebly
curved, apex short, widely rounded off. Internal
sac near apex with a sclerotized piece either at
bottom or at roof, according to the degree of
eversion of intemal sac. Left paramere with 5
setae, right paramere apparently 4-setose.

Female genitalia. Stylomere 2 with dorsal en-
siform seta situated rather medially,

DISTRIBUTION

Lowland at eastern foot of Bellenden Ker
Range, north Queensland.

HABITAT

A lowland species, found in "palm swamp'" and
lowland rainforest on mossy tree trunks by pyre-
thrum knockdown and Berlese extraction. Col-
lected October-November.

ETYMOLOGY
Refers to the occurrence at low elevation only,

342

Rainforest Zones

1.Finnigan
2. Thornton
3.Windsor Tbld

MEMOIRS OF THE QUEENSLAND MUSEUM

4.Carbine ТЫа
5.Hann Tableland
6.Black Mtn
7.Lamb Range-
8.Walsh Range
9.Hugh Nelson
10.Atherton
11.Bellenden Ker
12.Malbon Thomps.
13.Graham Range
14.Walter Hill 4
15.Kirrama/Cardw. 73 1 -

16.Seaview Ва - 0

17.Hinchinbrook 4 0
18.Paluma/Bluewat. - 0
19.Mt Elliot-

0 2

[ T

4 6 8 10

a

12 14

No. of spp. of genus Philipis

FIG. 10. Species numbers of genus Philipis in the north Queensland mountain/rainforest blocks. For explanation

see text.

Philipis spurgeoni sp. nov.
(Figs 13C, 18C)

MATERIAL EXAMINED

HOLOTYPE: QMT13569; $, 16°24’S, 145?13 E,
3.5km NNE Mt. Spurgeon, NEQ, 16 Oct 1991, 1330m,
Monteith & Janetzki, Pyrethrum, trees & rocks.

DIAGNOSIS

Medium sized, convex, with s-shaped fascia in
posterior half of elytra. Distinguished by larger
size, narrower base of elytra, and presence of
microreticulation on elytra.

DESCRIPTION

Measurements. Length: 2.28mm; width:
1.08тт. Ratios: Width pronotum/head: 1.55;
width/length of pronotum: 1.54; width base/apex
of pronotum: 1.44; width widest diameter/base of
pronotum: 1.07; length/width of elytra: 1.30.

Colour. Dark piceous, apex of elytra and pos-
terior part of suture faintly lighter. Elytra with a
fairly well delimited, moderately conspicuous,
oblique, rather s-shaped, light reddish fascia in
posterior 2/5 medially reaching 2nd stria. Legs
light reddish.

Head. Median antennomeres c. 1.2 x as long as
wide. Microreticulation distinct on labrum and
anterior border of clypeus, posteriorly moder-
ately superficial, about isodiametric. Surface
glossy.

Pronotum. Transverse, convex, widest about
in middle, base much wider than apex, lateral
margin posteriorly almost straight, slightly
oblique. Base slightly narrower than widest diam-
eter. Posterior angles rectangular, feebly pro-
duced over lateral part of base. Carina at posterior
angle elongate, rather oblique, slightly incurved.
Anterior transverse sulcus rather deep, not inter-
rupted. Posterior transverse sulcus very deep, in-
terrupted by a fovea. Anterior lateral seta situated

REVISION OF PHILIPIS

well behind anterior third of margin,
Microreticulation absent, surface highly glossy.

Elytra. Short, convex, egg-shaped, widest
ahout in middle. Lateral border evenly curved,
extremely finely serrate and pilose. Sutural stria
crenulate. 2nd stria gently impressed, even 3rd-
Sth striae very faintly impressed in anterior half.
Striae moderately punctate, 2nd slightly crenu-
late. Outer striae traceable in anterior half as rows
of fine punctures. Inner four striae just visible at
apex. Sutural interval convex throughout, 2nd
interval in basal half faintly convex. 8th stria
deeply impressed, punctate-crenulate, attaining
posterior marginal pore. Recurrent striole rather
oblique, meeting position of 3rd stria. Anterior
discal pore in anterior third, posterior pore
slightly in front of posterior third. Microreticula-
tion distinct, though somewhat superficial, com-
posed of slightly transverse meshes. Surface
rather glossy, but markedly less glossy than pro-
thorax. Wings slightly shortened. Variation un-
known,

Lower surface, Metepisternum slightly longer
than wide.

Genitalia. Male unknown. Female stylomere 2
with very stout dorsal ensiform seta situated
rather basally.

DISTRIBUTION
Mt. Spurgeon area, Carbine Tableland, north
Queensland. Known only from type locality.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks and rocks in montane rainforest above
1300m. Collected only October.

ETYMOLOGY
Refers to the type locality, Mt, Spurgeon,

Philipis trunci (Darlington)
(Figs 5D, 13D, 18C)

Tachys trunci Darlington, 1963; 31,
Philipis trunci, Erwin 1994: 568.

MATERIAL EXAMINED

HoLoTvPE: M.C.Z. Type No.30335; d, Mt. Alexan-
dra, Thornton Peak Natl, Park, OLD, Dee,” 197 (MCZ).
ADDITIONAL MATERIAL: 10 d, 6 9, Thornton Peak via
Daintree, МЕО, 1000-1300m, 20-22 Sept 1981, GM &
DC(ANIC, CBM, OM, USNM, ZSM); 1 d, Thomton
Peak via Daintree, NEQ, 20-22 Sept 1981, GM & DC,
QM Berlesate No. 301, RF, 1000-1300m, sieved litter
& moss (QM); 2 Z, 1 ?, Thomion Peak, 11 km NE
Daintree, NEQ, 1 Nov 1983, GM, DY & GT, QM

343

Berlesate No. 605, 16° LOS, 145?22E, RF, 1100m, moss
ûn rocks & trees (OM),

DIAGNOSIS

Small, convex, with s-shaped fascia in posle-
rior half of elyira, Distinguished by deeply im-
pressed 1st and 2nd striac and by outer striac
almost absent.

DESCRIPTION

Measurements, Length: 2.0-2.25mm:; width:
0.95-1.03mm. Ratios: Width pronotum/head;
1.51-1.56; width/length of pronotum: 1.51-1.53;
width base/apex of pronotum: 1.38-1.44; width
widest diameter/base of pronotum: 1.09-1.13;
length/width of elytra: 1.28-1.31.

Colour. Dark piceous, apex of clytra and pos-
terior part of suture faintly lighter. Elytra with a
rather ill delimited, inconspicuous, oblique, more
or less s-shaped, light reddish fascia in posterior
2/5 medially reaching almost to 2nd stria. Legs
reddish,

Head. Median antennomeres c. 1.3 x as long as
wide. Microreticulation distinct on labrum and
anterior border nf clypeus, extremely superficial
and very difficult to see on frons, absent on ver-
tex, about isodiametric. Surface highly glossy.

Pronotum. Transverse, convex, widest about
in middle, base much wider than apex. Lateral
margin posteriorly almost straight, slightly
oblique, even extremely faintly concave in front
of posterior angles. Posterior angles rectangular,
feebly produced over lateral part of base. Carina
at posterior angle elongate, markedly oblique,
slightly incurved. Anterior transverse sulcus
rather deep, barely interrupted. Posterior trans-
verse sulcus very deep, interrupted by a very
large, deep fovea. Anterior lateral seta situated
slightly behind anterior third of margin,
Microreticulation absent, surface highly glossy.

Elytra. Short, highly convex, egg-shaped, wid-
estabout in middle. Lateral border evenly curved,
extremely finely serrate and pilose. Sutural stria
crenulate. 2nd stria beginning well behind base,
punctate, almost as deeply impressed as sutural
stria. 3rd-5th striae absent or indicated in anterior
half only as extremely inconspicuous rows of
very fine punctures, outer striae completely ab-
sent. Only sutural stria visible at apex. Sutural
interval and basal half or 2/3 of 2nd interval
convex, 8th stria deeply impressed, punctate-
crenulate, barely attaining posterior marginal
pore, Recurrent striole rather oblique, meeting
end of 3rd stria. Anterior discal pore in anterior
third, posterior pore slightly in front of posterior

344 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 11. Habitus. A, Philipis quadraticollis sp. nov. B, P. alticola sp. nov. C, P. cooki sp. nov. D, P. laevigata
sp. nov. E, Philipis heatherae sp. nov. F, P. picea sp. nov. Lengths: 2.7mm; 2.75mm; 2.6mm; 2.2mm; 2.35mm;
2.5mm,

REVISION OF PHILIPIS

third. Microreticulation on disk extremely super-
ficial, hardly visible, composed of transverse
meshes; or absent. Surface highly glossy. Wings
shortened.

Lower surface. Metepisternum slightly longer
than wide.

Male genitalia, Genital ring triangular, narrow,
rather asymmetric, apex narrow. Aedcagus mod-
erately elongate, lower surface straight or faintly
bisinvate in basal 2/3, in apical third slightly
curved down, apex moderately elongate, widely
rounded off. Internal sac near apex with moder-
ately distinct trianglar fold. Parameres rather
elongate, both 5-setose.

Female genitalia. Stylomere 2 with very stout
dorsal ensiform seta situated rather basally.

Variation. Generally little variation noted,
apart from some differences in distinctness of the
elytral fascia which is in one specimen even
almost invisible. One specimen, however, differs
in following respects: base of pronotum narrower
than usual (ratio width base/apex 1.38 compared
with 1,43-1.44 as usual); 2nd stria far less im-
pressed than usual; oblique fascia on elytra morc
contrasting than usual.

DISTRIBUTION

Thornton Peak north of Daintree, north
Queensland. Known only from this monuntain
top.

HABITAT

Collected by pyrethrum knockdown and Berl-
ese extraction of mossy tree trunks and rocks in
montane rainforest above 1000m. The unique
type was collected "on the trunk of a small tree in
rainforest at probably about 3,000 ft, altitude",
Collected September, November, and December.

Philipis tribulationis sp. nov.
(Figs 5E. 13E, 18C)

MATERIAL EXAMINED

HoLotyPe: QMTI13570; d, 4.0km W of Cape Tribu-
lation (Site R), МЕО, 28 Sept 1982, 720m, Monteith,
Yeates & Thompson, Pyrethrum knockdown, RF.
PARATYPES: 2 p 3.5km W of Cape Tribulation (Site
7), МЕО, 2 Oct 1982, 680m, GM, DY & GT, Pyr, RF
(QM); 1 d, 1.9, 4.5-5.0km W. of Cape Tribulation
(Top Сатр), МЕО, 1-6 Oct 1982, 760-780m, GM, D'Y
& GT, Pyr., RF (CBM, QM); 1 9, 16"04'S, 145724 E,
Mt Pieter Botte, МЕО, 950m, 21 Nov 1993, GM & HI,
Pyr./trees, logs, rocks (QM); 4 d, 16703'S, 145"25' E,
Mt Haleyon, NEQ, 870m, 23 Nov 1993, GM & HJ.
Pyr trees & logs (ANIC, CBM, OM), 1 2, Windsor
Tableland, МЕО, 9 Jan 1989, 1225m, ES & ANZSES
Site 3, Руг, (QM),

DIAGNOSIS

Small, convex, with s-shaped fascia on poste-
rior half of elytra. Distinguished by 2nd stria less
impressed than Ist, smaller size, and rather nar-
row base of pronotum:

DESCRIPTION

Measurements. Length: 1,95-2.14mm; width:
0.92-0.98mm. Ratios; Width pronotum/head:
1.48-1,54; width/length of pronotum: 1.50-1.53;
width base/apex of pronotum; 1.40-1.42; width
widest diameter/base of pronotum: 1.08-1.11;
length/width of elytra: 1.31-1,33.

Colour. Dark piceous, pronotum, basal third
and apex of elytra slightly lighter, posterior half
of sulure even lighter, reddish-piceous. Elytra
with a well-defined, contrasüng, oblique, s-
shaped, dark yellow or light reddish fascia in
posterior 2/5 medially reaching 2nd stria. Legs
yellow,

Head. Frons anteriorly somewhat uneven. Me-
dian anennomeres c, 1,3 x as long as wide.
Microreticulation distinct on labrum and anterior
border of clypeus, very superficial and difficult
to see on frons, absent on vertex, about jsodiamet-
ric. Surface glossy.

Pronorum. Rather transverse, convex, widest
about in middle, base much wider than apex.
Lateral margin évenly curved, though posteriorly
less so than towards apex. Postenor angles rec-
tangular or even slightly acute, feebly produced
over lateral part of base. Carina at posterior angle
elongate, oblique, slightly incurved, Anterior
transverse sulcus rather deep, slightly interrupted
by a fovea. Posterior transverse sulcus very deep,
interrupted by a large, deep fovea. Anterior Lat-
eral seta situated slightly behind anterior third of
margin. Microreticulation absent, surface highly
glossy.

Elytra. Short, highly convex, egg-shaped, wid-
est aboul in middle. Lateral border evenly curved,
extremely finely serrrate and pilose. Sutural stria
crenulate. Other striae well developed as rows of
fairly coarse punctures, though becoming finer
laterally and posteriorly, 2nd and 3rd striae in
anterior half even faintly impressed. At least
inner striae traceable to apex, Sutural interval and
basal part of 2nd interval somewhat convex. 8th
stria deeply impressed, punctate-crenulate,
barely attaining posterior marginal pore. Recur-
rent striole rather evenly curved, meeting end of
3rd stria. Anterior discal pore in anterior third,
posterior pore slightly in front of posterior third.
Microreticulation transverse, on disk strongly re-
duced and extremely superficial, even under high

magnification amost invisible, slightly better
seen near apex. Surface rather glossy. Wings
shortened.

Lower surface. Metepisternum barely longer
than wide.

Male genitalia. Genital ring triangular, rather
asymmetric, especially at base, apex rather elon-
gate. Aedeagus moderately elongate, lower sur-
face basally straight, feebly curved to apex, apex
fairly short, widely rounded off. Both parameres
5-setose,

Female genitalia, Stylomere 2 with dorsal
ensiform seta situated rather medially.

Variation, Apart from some differences of size
and relative width of pronotum, very little varia-
tion noted.

DISTRIBUTION
Mountains some kilometers west of Cape Trib-
ulation to Windsor Tableland, north Queensland.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks m montane rainforest between 680
and 1200m. Collected September-October and
January.

ETYMOLOGY
Refers to the type locality, the vicinity of Cape
Tribulation.

Philipis thompsoni sp. nov,
(Figs 2, 5F, 13F, !7B)

MATERIAL EXAMINED

HOLOTYPE: QMT13578; $, Hugh Nelson Ra., 2.5km
5. of Crater N.P., МО), 5 Dec 1988, 1100m, Monteith
& Thompson, pyrethrum/logs & trees.

PARATYPES: 3 d, 2 9, ] (?sex), same data (CBM,
QM); 1 d, 1 9, Mt. Fisher, 7km SW Millaa Millaa,
NO (Whiteing Rd), 5 May 1983, 1200m, GM, DY, RF,
Pyr. (QM); 2 5, Mt. Edith, Lamb Range, МЕО, 12 Oct
1982, 1000-1100m, GM, DY & GT, Pyr., RF (QM); 1
Ф, Emerald Ck, Lamb Range, МЕО, 11 Oct 1982,
950m, GM, DY & GT, Pyr., RF (CBM); | 3, 1 9,
21кт5 Atherton, МЕО, 1040-1 100m, 5 Nov 1983, DY
& GT, Pyr., RF(CBM, QM); 1 9, Baldy Min Rd, 7 km
SW Atherton, МЕО, 9 Dec 1988, 1 150m, GM & GT,
Pyc/logs & trees (QM); | 3, 1 9, Mt. Father Clancy,
9 km S Millaa Millaa, NEQ, 6 Dec 1988, 1000m, GM
& GT, Pyr./loes & trees (QM): 3 d. Mt. Formartine
South, 10km № Kuranda, NEQ, 23 Nov 1990, 700m.
GM & GT, Pyr./irees & logs (QM); 1 d, E 9,167558,
145?49' E, Mt. Williams, МЕО, 900- 1000m, 2-3 Dec
1993, DC, GM & HJ (QM); 1 4,1 9, 17°03'S,
145?42"E, Isley Hills, МЕО, 1050m, 30 Nov 1993, GM
& HJ, Pyr./trees & logs (QM); 1 &, 17^16S, 145?49" E,
Massey Range, 4km W of Centre Bellenden Ker, NEQ,

MEMOIRS OF THE QUEENSLAND MUSEUM

9-11 Oct 1991, 1250m, GM, HJ & DC (QM); 1 3, Mt.
Bartle Frere, МЕО, West Side, 1050m, 8 Dec 1990.
GM, GT & RS, Pyr./trees & rocks (QM); 1 9,1 (sex?),
Upper Boulder Ck. via Tully, NEQ, 900m, 26 Oct
1983, GM, DY & GT. Pyr., RF(QM); 6 d, 3 2, Upper
Boulder Creek. 11km N Tully, МЕО, 5 Dec 1989,
1000m, GM, СТ, HJ, Pyr/logs & trees (ANIC, СВМ,
QM, USNM, ZSM); 2 5,1 9, Tully Falls, МЕО, 8 Dec
1990, 750m, GM, GT, HJ, Pyr., logs & trees (CBM,
QM); 2 2, Cardwell Range, МЕО, Mt Macalister area,
1000m, 19 Dec 1986, GM, GT & SH, Pyr. (QM),

DIAGNOSIS

Medium sized, convex, with s-shaped fascia in
posterior half of elytra, Distinguished by larger
size, wide base of elytra, absence of
microrcticulation on elytra and shorter aedeagus
with almost straight lower surface.

DESCRIPTION

Measurements. Length: 2.15-2. 30mm; width:
1.0-1.1mm. Ratios; Width pronotum/head: 1.51-
1.55; width/length of pronotum: 1.46-1.52; width
base/apex of pronotum: 1.45-1.49; width widest
diameter/base of pronotum: 1.08-1.11;
lengih/ width of elytra; 1,33-1.34.

Colour. Dark piceous, apex of elytra and pos-
terior part of suture, commonly also base of elytra
faintly lighter. Elytra with a fairly well delimited,
moderately conspicuous, oblique, more or less
s-shaped, light reddish fascia in posterior 2/5
medially reaching 2nd stria, Legs reddish.

Head. Median antennomeres c. 1.3 x as long as
wide, Microreticulation distinct on labrum and
anterior border of clypeus, highly superficial, but
discernible at high magnification on frons, absent
on vertex, about isodiametric. Surface glossy.

Pronotum. Transverse, convex, widest about
in middle, base much wider than apex. Lateral
margin evenly curved, though posteriorly some-
limes almost straight, slightly oblique. Posterior
angles rectangular, fcebly produced over lateral
part of base. Carina at posterior angle elongate,
markedly oblique, slightly incurved. Anterior
transverse sulcus rather deep, nol interrupted,
Posterior transverse sulcus very deep, barely in-
terrupted, Anterior lateral seta situated well be-
hind anterior third of margin. Microreticulation
absent, surface highly glossy.

Elytra. Shon, highly convex, egg-shaped, wid-
est about in middle. Lateral margin evenly
curved, extremely finely serrate and pilose. Su-
tural stria crenulate. 2nd stria well visible as a row
of fairly distinct punctures, sometimes even
faintly impressed in anterior half. Outer striae
traceable at least in anterior half as fine rows of

REVISION OF PHILIPIS

punctures. Only sutural stria visible at apex. Su-
tural interval convex, sometimes also 2nd interval
in basal half faintly convex. 8th stria deeply im-
pressed, punctate-crenulate, clearly attaining
posterior marginal pore. Recurrent striole rather
oblique, meeting end of 3rd зіла, Anterior discal
pore in anterior third, posterior pore slightly in
front of posterior third. In males microreticula-
tion on disk extremely superficial and hardly
visible, or absent, in most females microreticula-
non slightly more distinct, composed of trans-
verse meshes. Surface in males highly glossy, in
females usually slightly less glossy. Wings
slightly shortened.

Lower surface. Metepisternum slightly longer
than wide.

Male genitalia. Genital ring triangular, narrow,
rather asymmetric, apex narrow. Aedeagus mod-
erately elongate, lower surface straight in basal
2/3, in apical third feebly curved down, apex
moderately elongate, widely rounded off. Inter-
nal sac near apex with very dislinct triangular
fold. Parameres rather short, both 5-setose,

Female genitalia. Stylomere 2 with very stout
dorsal ensiform seta situated rather basally,

Variation, Due to the wide range of this species,
some variation of colour, distinctness of pattern,
shape of pronotum, and degree of microreticula-
tion and striation of elytra noted.

DISTRIBUTION

A widely distributed species from the Kuranda
area, the Lamb Range, mountains surrounding
the Atherton Tableland, and south to the Cardwell
Range, north Queensland.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks and logs in montane rainforest above
700m. Collected October-December and May.

ETYMOLOGY
Named in honour of Geoff Thompson, collec-
tor of many specimens of Philipis.

Philipis vicina sp. nov.
(Figs 6A, 14A, 17В)

MATERIAL EXAMINED

HoLoTYvPE: QMTI3612; d, Mt. Misery Summit via
Helenyale, NEQ, 6 Dec 1990, 850m, Monteith, Sheri-
dan & Roberts, Pyrethrum-trees & logs.

DIAGNOSIS
Medium sized, convex, with s-shaped fascia in
posterior half of elytra. Distinguished by larger

347

size, narrower base of elytra, absence of
microreticulation on elytra, and longer aedeagus
with bisinuate Jower surface.

DESCRIPTION

Measurements. Length: 2.20mm; width:
1.00mm. Ratios: Width pronotum/head: 1.52;
width/length of pronotum: 1.55; width base/apex
of pronotum: 1.40; width widest diameter/base of
pronotum: 1,10; length/width of elytra: 1.33.

Colour. Dark piceous, apex of elytra and pos-
terior part of suture faintly lighter. Elytra with a
fairly well delimited, moderately conspicuous,
oblique, s-shaped, light reddish fascia in posterior
2/5 medially reaching 2nd stria. Legs light red-
dish.

Head, Median antennomeres c. 1.2 x as long as
wide. Microreticulation distinct on labrum and
anterior border of clypeus, postenorly superfi-
cial, about jsodiametric. Surface glossy.

Pronotum. Transverse, convex, widest about
in middle, base much wider than apex. Lateral
margin posteriorly almost straight, slightly
oblique. Base little narrower than widest diame-
ter. Posterior angles rectangular, feebly produced
over lateral part of base. Carina at posterior an-
gles elongate, rather oblique, slightly incurved.
Anterior transverse sulcus rather deep, not inter-
rupted. Posterior transverse sulcus very deep, in-
terrupted by a fovea. Anterior lateral seta situated
well behind anterior third of margin,
Microreticulation absent, surface highly glossy.

Elytra, Short, convex, egg-shaped, widest
about in middle. Lateral border evenly curved,
extremely finely serrate and pilose. Sutural stria
crenulate. 2nd stria faintly impressed in anterior
half, 3rd-5th striae well visible as rows of mod-
erately fine punctures, outer striae traceable in
anterior half as rows of fine punctures. Inner four
striae just visible at apex. Sutural interval convex
throughout, 2nd interval in basal half faintly con-
vex. 8th stria deeply impressed, punctate-crenu-
late, altaining posterior marginal pore. Recurrent
striole elongate, rather oblique, meeting position
of 3rd stria, Anterior discal pore in anterior third,
posterior pore slightly in front of posterior third.
Microreliculation almost absent. Surface highly
glossy. Wings slightly shortened. Variation ün-
known

Lower surface. Metepisternum slightly longer
than wide.

Genitalia, Male genital ring triangular, narrow,
rather asymmetric, apex narrow. Aedeagus fairly
elongate, lower surface perceptibly bisnuate in
basal 2/3, in apical third feebly curved down,

348 MEMOIRS OF THE QUEENSLAND MUSEUM

ve

i.
20
iv.
IN
hA
p

Dl

FIG. 12. Habitus. A, P. castanea sp. nov. B, P. unicolor sp. nov. C, Philipis atra sp. nov. D, P. ruficollis sp. nov.
E, P. inermis sp. nov. Е, P. reticulata sp. nov. Lengths: 2.35mm; 2.05mm; 2.45mm; 2.5mm; 1.95mm; 2.2mm.

REVISION OF PHILIPIS

apex rather elongate, widely rounded off. Internal
sac near apex with very distinct triangular fold.
Parameres rather elongate, both 5-setose. Female
genitalia unknown.

DISTRIBUTION
Mt. Misery south of Helenvale, north Queens-
land. Known only from type locality.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks and logs in montane rainforest above
850m. Collected December only,

ETYMOLOGY
Refers to the very close relationships of this
species with P. thompsoni and P. spurgeoni,

Philipis laevis sp. nov.
(Figs 6B, 14B, 17B)

MATERIAL EXAMINED

HOLOTYPE: QMT13613; d, Thornton Peak via
Daintree, NEQ, 1000-1300m, 20-22 Sept 1981, G.
Monteith & D. Cook,

PARATYPES: 2 9, same data (QM); 7 d, 6 9, Thom-
ton Peak summit, via Daintree, NEQ, 24-27 Sept 1984,
GM & SM, QM Berlesate NO. 662, RF, 1100-1300m,
sieved litter & moss (ANIC, CBM, QM, USNM,
ZSM).

DIAGNOSIS

Small, short, convex, easily identified by
quadrimaculate pattern, absence of all elytral
striae apart from Ist, and highly glossy surface.

DESCRIPTION

Measurements. Length: 2.05-2.30mm; width:
1.04-1.12mm. Ratios: Width pronotum/head:
1.58-1.62; width/length of pronotum: 1.43-1.49;
width base/apex of pronotum: 1.41-1.45; width
widest diameter/base of pronotum: 1.12-1.15;
length/width of elytra: 1.24-1.28.

Colour. Head and pronotum dark piceous,
elytra slightly lighter, piceous orreddish-piceous.
Elytra with two rather ill-defined spots, a larger,
about triangular one at outer angle of shoulder,
and a smaller, narrower, fascia-like one behind
middle, extending frorn border to position of 3rd
stria. Legs reddish to reddish-brown, apex of
femora and tibiae, and tarsi yellow.

Head. Median antennomeres c. 1.75 x as long
as wide. Microreticulation well visible only on
labrum and anterior half of clypeus, extremely
superficial and hardly recognizable on anterior

349

part of frons, absent on rest of head, about isodi-
ametric. Surface highly glossy.

Pronotum. Rather convex, transverse, widest
about in middle, base much wider than apex.
Lateral margin strongly and evenly curved,
though more to apex than to base. Posterior an-
gles rectangular, distinctly produced over lateral
part of base, Carina at posterior angle compara-
tively short, oblique, slightly incurved. Anterior
transverse sulcus deep, uninterrupted. Posterior
transverse sulcus very deep, interrupted by a large
fovea, Anterior lateral seta situated slightly be-
hind anterior third of margin, Microreticulation
absent, surface highly glossy.

Elytra, Short and wide, highly convex, widest
in middle, lateral border evenly curved. Sutural
stria in anterior half slightly punctulate, posteri-
orly smooth, Other striae absent, but sometimes
2nd stria in basal half faintly indicated as a row
of delicate punctures. Only sutural interval con-
vex. 8th stria deeply impressed, barely punctate,
not attaining posterior marginal pore. Recurrent
striole rather short, close to lateral border and
markedly oblique, anteriorly suddenly incurved,
hardly meeting position of 3rd stra. Anterior
discal pore in anterior third, posterior pore
slightly behind posterior 2/5, Microreticulation
absent, surface highly glossy. Wings extremely
shortened.

Lower surface. Metepisternum about as long as
wide.

Male genitalia. Genital nng slightly asymmet-
rical, apex rather acute, Aedeagus short and com-
pact, lower surface to apex slightly curved. Apex
short, rounded off. Both parameres 5-setose.

Female genitalia. Stylomere 2 with very stout
dorsal ensiform seta situated about medially,

Variation, Apart from some minor differences
insize, relative shape of pronotum, and degrec of
2nd stria of elytra, little variation noted.

DISTRIBUTION
Thornton Peak north of Daintree, north
Queensland. Known only from this mountain top.

HABITAT

Collected in montane rainforest above 1000m
by Berlese extraction frorn litter and moss. Col-
lected September.

ETYMOLOGY
Refers to the remarkably smooth and glossy
surface of elytra.

Philipis sulcata sp. nov.
(Figs 6C, 14C, 18D)

MATERIAL EXAMINED

HorLorvre: QMT13624; 3, nr. Plane Crash Site,
llkm NW Mossman, МЕО, 28 Dec 1989, 1240m,
ANZSES. Pyrethrum.

PARATYPES: 2 d, 1 $, same data (CBM, QM); 2 9,
Pauls Luck, Platypus Ck, 13km W Mossman, МЕО, 1-2
Jan 1990, 1100m, ANZSES, Pyr. (QM); 2 d, 1 2,
Mossman Bluff Summit 10km W Mossman, МЕО, 18
Dec 1988, 1300m, GM & GT, Pyr. trees & rocks (QM,
USNM); 1 d, Mossman Bluff Track 6km W Mossman,
NEQ, 16 Dec 1988, 480m, GM & GT, Pyr. trees &
rocks (QM); 5 9, Devil's Thumb 12km NW
Mossmann, МЕО, 27 Dec 1989, 1000m, ANZSES,
Pyr. (ANIC, CBM, QM, ZSM).

DIAGNOSIS

Small to medium sized, highly convex, easily
identified by the fully and deeply striate elytra,
the elongate, on lower surface near apex strongly
sinuate aedeagus with elongale apex, and the
elongate, parallel parameres; further distin-
guished from P. perstriafa sp. nov. by wider
elytra, absence of microrcticulation on the elytra,
and shorter apical elytral spot,

DESCRIPTION

Measurements. Length: 2.05-2.5mm; width:
0.95-1,2mm. Ratios: Width pronotum/head:
1.58-1.64; width/length of pronotum: 1.53-1.57;
width base/apex of pronotum: 1.48-1.57; width
widest diameter/base of pronotum: 1.11-1.15;
length/width of elytra: 1.27-1.31.

Colour. Dark piceous, pronotum and posterior
half of suture slightly lighter, in light specimens
pronotum even dark reddish. Behind shoulder
and in posterior 2/5 of elytra with a rather incon-
spicuous, slightly transverse, yellowish spot
each, both attaining about 4th stria. Both spots
moderately conspicuous and not well delimited,
Pores on 3rd stria encircled by a small yellowish
spot. Legs dark yellow,

Head. Very short and wide, neck very wide,
separated from frons by a shallow impression.
Median antennomeres barely longer than wide.
Microreticulation distinct on whole surface, even
on vertex, though somewhat superficial, about
isodiametric to slightly transverse. Surface rather
dull,

Pronotum, Transverse, convex, widest about in
middle or slightly in front of it, base much wider
than apex, Lateral margin posteriorly but feebly
convex, fairly oblique. Posterior angles rectangu-
lar, barely ed over lateral part of base.
Carina at posterior angle very elongate, mark-

MEMOIRS OF THE QUEENSLAND MUSEUM

edly oblique, slightly incurved. Anterior trans-
verse sulcus deep, not interrupted. Median line
ralher deeply impressed. Posterior transverse sul-
cus very deep, interrupted by a rather large fovea,
Anterior lateral seta situated well behind anterior
third of margin, Microreticulation very superfi-
cial or almost absent, surface glossy.

Elytra. Very short and convex, egg-shaped,
widest about jn middle, Lateral border evenly
curved, not perceptibly serrate and pilose. All
striae deeply impressed, sulcate, and crenulate till
apex. All intervals markedly convex along their
whole course. 8th stria deeply impressed, anteri-
orly barely shallower, crenulate, altaining poste-
nor marginal pore. Recurrent striole elongate,
markedly oblique, anteriorly suddenly incurved,
meeting 3rd stria. Anterior discal pore well be-
hind anterior third, posterior pore very close [0
anterior pore, shortly behind middle. Both pores
very large, foveiform, interrupting 3rd interval.
Microreticulation highly superficial to almost ab-
sent, moderately transverse. Surface rather
glossy. Wings extremely shortened,

Lower surface. Metepisternum c. 1.2 x as long
as wide.

Male genitalia. Genital ning triangular, rather
asymmetric, moderately wide, apex narrow.
Aedcagus fairly elongate, lower surface conspic-
uously bisinuate near apex, apex clongate, far
protruding, curved, fairly wide and widely
rounded off. Parameres elongate, right narrow,
both with 5 unusually short setae of which only
the most apical seta 15 rather elongate, at or near
apex.

Female genitalia, Stylomere 2 with stout dorsal
ensiform seta situated rather medially.

Variation. Apart from some minor differences
in colour, relative shape of pronotum, and degree
of microreticulation on pronotum and elytra, little
variation noted. However, there 1s one perhaps
not fully coloured and somewhat deformed spec-
imen that shows more distinct aberrations in
shape and microsculpture: it has a remarkably
rough surface of the elytral intervals, and strongly
crenulate striae. However, this may be due to
some creasing of the elytra shortly after emerg-

ing.

DISTRIBUTION
Mountains of the Carbine Tableland west of
Mossman, north Queensland.

HABITAT
Collected by pyrethrum knockdown on mossy
trees and rocks in montane rainforest, usually

REVISION OF PHILIPIS

above 1100m, but one record just below 500m.
Collected December and January.

ETYMOLOGY
Refers to the deeply sulcate elytral striation.

Philipis frerei sp. nov.
(Figs 6D, 14D, 18D)

MATERIAL EXAMINED

HOLOTYPE: QMT13634; d, Mt. Bartle Frere, МЕО,
0.5km N of Sth. Peak, 6-8 Nov 1981, 1500m, Earth-
watch/Old. Museum pyrethrum knockdown.
PARATYPES: 1 d, 6 ©, Mt. Bartle Frere, МЕО, Sth.
Peak Summit, 1620m, 6-8 Nov 1981, EW/QM Pyr.
(ANIC, СВМ, QM, ZSM), 1 2, Mt. Bartle Frere,
summit creek, NEQ, 24 Sept 1981, GM & DC, QM
Berlesate No. 304, RF, 1500m, sieved litter (QM); |
d,1 9, Mt. Bartle Frere, МЕО, Central Ridge, 1500m,
27 Dec 1989, GM, Pyr., logs (CBM, QM).

DIAGNOSIS

Medium sized, quadrimaculate, with reddish
pronotum and transverse apical elytral spot, dis-
tinguished from similarly patterned species by
almost fully striate elytra, very weak
microreticulation on the elytra, small aedeagus
with elongate apex, and 4-setose parameres.

DESCRIPTION

Measurements. Length: 2.05-2.45mm; width:
1.0-1.15mm. Ratios: Width pronotum/head;
1.60-1.64; width/length of pronotum: 1.44-1.51;
width base/apex of pronotum: 1.42-1.48; width
widest diameter/base of pronotum: 1.11-1.19;
length/width of elytra: 1.31-1.34.

Colour. Reddish-piceous, pronotum, apex and
suture of elytra faintly lighter. Elvtra at shoulder
with a very inconspicuous, extremely il] delim-
ited, about triangular, light reddish spot attaining
about position of 4th stria. In posterior 2/5 of
elytra with a moderately conspicuous, fairly well
delimited, narrowly triangular, slightly oblique,
light reddish spot or stripe, attaining about 3rd
stria. Legs light reddish, tibiae sometimes faintly
darker.

Head. Frons without median groove. Median
antennomeres c. 1.2 x as long as wide.
Microreticulation distinct and coarse on most of
head, weaker only on vertex, about isodiametnc
to slightly transverse, Surface somewhat dull.

Pronotum. Transverse, moderately convex,
widest about in middle, base much wider than
apex. Lateral margin posteriorly moderately
curved, rather oblique, at posterior angles faintly
incurved. Base usually distinctly narrower than

351

widest part. Posterior angles fairly obtuse,
slightly produced over lateral part of base. Carina
at posterior angle moderately elongate, rather
oblique, slightly incurved. Anterior transverse
sulcus fairly deep, feebly interrupted. Posterior
transverse sulcus deep, interrupted by large
fovea. Anterior lateral seta situated slightly be-
hind anterior third of margin. Microreticulation
distinct, though superficial, almost isodiametric,
surface moderately glossy.

Elytra, Rather short and convex, egg-shaped,
widest slightly behind anterior third. Lateral bor-
der evenly curved, extremely finely, almost not
perceptibly, serrate and pilose, Ist-3rd striae
deeply impressed, punctate-crenulate, All other
striae al least in basal half perceptibly impressed,
though becoming shallower laterally and api-
cally. AII striae moderately finely punctate. 1st-
4th intervals convex throughout, outer intervals
convex only in basal half, becoming posteriorly
rather depressed. All striae well visible at apex,
though outer striae finer. 8th stria deeply im-
pressed throughout, indistinctly punctate, attain-
ing posterior marginal pore. Recurrent striole
elongate, rather oblique, meeting 3rd stria. Ante-
rior discal pore well behind anterior third, poste-
rior pore well in front of posterior third.
Microreticulation indistinct and highly superfi-
cial, consisting of irregular, rather transverse
meshes. Surface glossy. Wings extremely short-
ened.

Lower surface, Metepisternum about as long as
wide.

Male genitalia. Genital ring almost regularly
triangular, wide, feebly asymmetric, apex rather
narrow. Aedeagus very small, short and compact,
lower surface in apical half strongly curved
down, apex moderately short, widely rounded
off. Parameres moderately elongate, both 4-se-
Tose.

Female genitalia, Stylomere 2 with stout dor-
sal ensiform seta situated rather medially.

Variation, Apart from some differences in rel-
ative shape of pronotum, little variation noted,

DISTRIBUTION
Mt. Bartle Frere, north Queensland. Known
only from that mountain top,

HABITAT

Collected hy pyrethrum knockdown on mossy
tree trunks and logs, and by Berlese extraction of
litter in montane rainforest above 1500m. Col-
lected September, November, and December.

352 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 13. Habitus. A, Philipis striata sp. nov. B, P. planicola sp. nov. C, P. spurgeoni sp. nov. D, P. trunci
(Darlington). A, Philipis tribulationis sp. nov. B, P. thompsoni sp. nov. Lengths: 2.3mm;2.1mm;2.3mm; 2.] mm;
2.0mm; 2.25mm.

REVISION DF PHILIPIS

ETYMOLOGY
Refers to the type locality, Mt. Bartle Frere.

Philipis subtropica sp. nov.
(Figs 6E, 14E, 17A)

MATERIAL EXAMINED

HOLOTYPE: д, 28^14'S, 153°08’E Lamington М.Р.
(O'Reillys) Q, 22-27 Oct 1978, Lawrence & Weir,
under bark rotten logs (ANIC).

PARATYPES: | 9, same data (ANIC); 1 d, 28?16'S,
153^10'E Мі. Bithongabel, 1400m, Lamington Nat.
Pk., Q, 23 Oct 1978, JL & TW, ANIC Berlesate 654,
moss & litter Nothofagus moorei (СВМ); 3 3.3 9,
28°15'S, 153?16' E, Springbrook Repeater, 1000m, 16
Apr 1995, GM, Pyr. tree trunks (CBM, QM).

DIAGNOSIS

Medium-sized to large, quadrimaculate, with
dark pronotum and transverse apical elytral spot,
distinguished from similarly patterned species by
absence of microreticulation on the elytra, pres-
ence of only 3 elytral striae, on lower surface
evenly curved aedeagus with elongate, rather
compact apex, and 5-setose parameres,

DESCRIPTION

Measurements. Length: 2.3-2.8mm; width:
1.0-1.25mm. Ratios: Width pronotum/head:
1.54-1.64; width/length of pronotum: (.42-1.47;
width base/apex of pronotum: 1.43-1.47; width
widest diameter/base of pronotum; 1.13-1.15;
length/width of elytra: 1.35-1.36.

Colour. Reddish-piceous, head slightly darker,
apex of elytra and sometimes suture faintly ligh-
ter. Elytra at shoulder with an inconspicuous, il]
delimited, about triangular, light reddish spot at-
taining about position of 5th stria, In posterior 2/5
of elytra with an equally ill delimited, transverse,
yellowish spot, attaining about 2nd stria. Legs
dark yellow to slightly infuscate,

Head, Median antennomeres c. 1.3 х as long as
wide. Microrcticulation distinct on labrum and
anterior part of clypeus, superficial on frons and
vertex, about isodiamcetric to slightly transverse.
Surface fairly glossy.

Pronotum, Rather transverse, fairly convex,
widest slightly in front of middle, base much
wider than apex. Lateral margin posteriorly al-
most straight, fairly oblique. Posterior angles rec-
tangular, feebly produced over lateral part of
base. Carina al posterior angle fairly clongate,
moderately oblique, slightly incurved, Anterior
transverse suleus rather shallow, more or less
distinctly interrupted. Posterior transverse sulcus
fairly deep, becoming shallow in middle, though

353

without fovea, Anterior lateral seta situated well
behind апіепаг third of margin, Microreticula-
tion absent, surface highly glossy.

Elytra, Moderately short and convex, egg-
shaped, widest slightly behind anterior third. Lat-
eral border evenly curved, not perceptibly serrate
and pilose. Sutural stria crenulate, 2nd stria in
basal half just perceptible as a row of extremely
fine and superficial punctures, outer striae absent.
Ist interval convex, sometimes 2nd interval
faintly convex near base, Only sutural stria trace-
able to apex. 8th stria deeply impressed, anteri-
orly shallower, punctate, though anteriorly
smooth, just attaining posterior marginal pore, or
interrupted shortly in front of it. Recurrent striole
elongate, moderately oblique, meeting position
of 3rd stria. Anterior discal pore well behind
anterior third, posterior pore slightly in front of

sterior third. Microreticulation absent. Surface

ighly glossy. Wings extremely shortened.

Lower surface. Metepisternum e. 1,2 x as long
as wide.

Male genitalia. Genital ring very asymmetric,
not triangular, wide, apex wide. Aedeagus rather
large. fairly short and compact, lower surface
concave, apex rather elongate, wide, protruding,
widely rounded off, Internal sac within folded
sclerite with a conspicuous, strongly sclerotized
peg direcied anteroventrally. Parameres moder-
ately elongate, both 5-setose.

Female genitalia. Stylomere 2 with stout dorsal
ensiform sela situated rather medially.

Variation. Some apparent sexual variation
noted, since the few females tend to be markedly
larger than the males. Also some variation in
relative shape of the pronotuim. Length of male
genital ring and of aedeagus varies also to somc
degrec.

DISTRIBUTION

Lamington Plateau and the adjacent
Springbrook Plateau, southeast Queensland.

HABITAT

Collected in montane, temperate rainforest
above 1000m under bark of rotten logs, by Berl-
est extraction from moss and litter of Nothofagus,
and by pyrethrum knockdown on mossy tree
irupks. Collected April and October.

ETYMOLOGY

Refers to the occurrence in subtropical latitude
in south Queensland.

354
Philipis alutacea sp. nov.
(Figs 6F, 14F, 18D)
MATERIAL EXAMINED

HOLOTYPE: QMT13641; g, Bellenden Ker, МБО,
summit, 10 June 1980, G, B. Monteith.

PARATYPES: 1 9, same data (QM); 1 3, 2 $,
Bellenden Ker Range, МЕО, Summit TV Stn.. 1560m,
17 Oct-5 Nov 1981, EW/QM Pyr. (ANIC, CBM, QM);
1 5, Bellenden Ker Range, МЕО. Summit TV Stn.,
1560m, 1-7 Nov 1981, EW/QM, QM Berlesate No.
346, 17°16'S, 145?51'E, RF, moss on trees & rocks
(QM); | d, Bellenden Ker, МЕО, Centre Peak Summit,
10 Apr 1979, GM, QM Berlesate No. 10, 17°16'S,
145?51'E, RF, 1500m, moss on trees (CBM).

DIAGNOSIS

Medium-sized, quacrimaculate, with dark pro-
notum and transverse apical elytral elytral spot,
distinguished from similarly patterned species by
distinct microreticulation on the elytra, fully, but
very weakly striate clytra, on lower surface
bisinuate aedeagus with very Wide apex, and 5-
setose parameres.

DESCRIPTION

Measurements. Length: 2.25-2.45mm, width:
1.05-1.12mm, Ratios: Width pronotum/head:
1,64-1,68; width/length of pronotum: 1,46-1,49;
width base/apex of pronotum: 1.63-1.67; width
Widest diameter/base of pronotum: 1,04-1,06;
length/width of elytra: 1.32-1.35.

Colour. Reddish-piceous, head and that part of
elytra between the spots and just behind apical
spot slightly darker, Elytra at some distance be-
hind shoulder with an inconspicuous, ill delim-
ited, about tnangular, light reddish spot altaining
about position of 4th stria. In posterior 2/5 nf
elytra with a rather conspicuous, fairly well de-
limited, straight, oblique, light reddish spot or
stripe, surrounded by dark colour, attaining about
3rd stria. Both spots usually well removed from
lateral border. Legs light reddish, tibiae slightly
darker.

Head. Median antennomeres c. 1.6 x as long as
wide. Microreticulation distinct on most of head,
weaker only on vertex, about modiametric to
slightly transverse. Surface somewhat dull.

Pronotum. Transverse, moderately convex,
widest 1n or slightly behind of middle, base much
wider than apex. Lateral margin posteriorly little
curved, but feebly oblique, al posterior angles
faintly incurved. Hence base but slightly nar-
rower than widest part. Posterior angles fairly
obtuse, distinctly produced over lateral part of
base. Carina at posterior angle moderately elon-

MEMOIRS OF THE QUEENSLAND MUSEUM

gate, rather oblique, slightly incurved. Anlenor
transverse sulcus fairly deep, not interrupted.
Posterior transverse sulcus fairly deep, in middle
shallower, but without conspicuous fovea, Ante-
rior lateral seta situated well behind anterior third
of margin. Microreticulation very distinct, almost
isodiametric, surface rather dull.

Elytra. Rather short and convex, egg-shapeu,
widest slightly in front of middle. Lateral border
evenly curved, extremely finely, almost not per-
ceptibly, serrate and pilose. Sutural stria crenu-
late, 2nd stria in basal half slightly impressed,
3rd-Sth striae near base very faintly impressed,
outer striae very inconspicuous, All striae finely
punctate, puncturation becoming even finer later-
ally and apically. Sutural interval convex
throughout, 2nd interval slightly convex in basal
half, outer intervals depressed, Only Ist and 2nd
striae well visible at apex, outer striae barely
perceptible. Sth stria deeply impressed through-
put, punctate, attaining posterior marginal pore.
Recurrent striole elongate, rather oblique, meet-
ing position of 3rd stria. Anterior discal pore well
behind anterior third, posterior pore well in front
of posterior third. Microrcticulation distinct, con-
sisting of but feebly transverse meshes. Surface
rather dull, Wings slightly shorter than elytra.

Lower surface. Mctepisternum feebly longer
than wide,

Male genitalia, Genital ring regularly triangu-
lar, wide, apex rather narrow. Aedeagus rather
large, moderately compact, lower surface gently
bisinuate, apex short and very wide, widely
rounded off. Parameres large, elongate, both 5-
setose,

Female genitalia, Stylomere 2 with stout dor-
sal ensiform seta situated rather medially.

Variation, Some variation noted in relative
shape of pronotum and clytra and in degree of
microreticulation.

DISTRIBUTION
Mt, Bellenden Ker, north Queensland, Known
only from that mountain top.

HABITAT

Collected by pyrethrum knockdown on mossy
trunks and rocks in montane rainforest above
I500m. Collected April, June, October, and No-
vember.

ETYMOLOGY
Refers to the remarkably dul] surface,

REVISION OF PHILIPIS

Philipis rufescens sp. nov.
(Figs 7A, 15A, 18D)

MATERIAL EXAMINED

HOLOTYPE: QMT13645; d, Bellenden Ker Range,
NEQ, Summit TV Stn., 1560m, 17 Oct-5 Nov 1981,
Earthwatch/Qld. Museum pyrethrum knockdown.
PARATYPES: 9 d, 7 9, same data (ANIC, CBM,
QM, USNM, ZSM); 7 4,1 9, Bellenden Ker, Centre
Peak Summit, NEQ, 10 Apr 1979, GM, QM Berlesate
No. 8, 17°16'S, 145°51'E, RF, 1500m, moss on trees
(CBM, ОМ); 1 =, Bellenden Ker summit, МЕО, 10
June 1980, GM (QM); 19 d, 14 9, Bellenden-Ker
Range, МЕО, Summit TV Stn., 1560m, 28 Oct 1983,
GM, DY & GT, Pyr. in RF (ANIC, CBM, QM,
USNM); 1 2.4 2, Bellenden Ker Range, МЕО, Sum-
mit TV Stn., 1560m, 1-7 Nov 1981, EW/OM, QM
Berlesate NO, 337, 340, 341, 343, 344, 17"16'S,
14595 Е, RF, sieved litter, stick brushings, stick &
moss brushings, moss on trees & rocks (QM); 1 5, Mt.
Bellenden Ker, Centre Peak Summit, NEQ, 10-12 Apr
1979, 1500m, GM (QM);9 3,4 9, 17°16'5, 145°52B,
Bellenden Ker summit, NEQ, 8 Oct 1991, 1560m, GM
& HJ, Pyr., trees & logs (CBM, OM, ZSM).

DIAGNOSIS

Medium-sized, reddish-piceous, quadrimacul-
ate, with red pronotum and oblique, slightly sin-
uate apical elytral spot, further distinguished by
fully and deeply striate elytra, distinct
microreticulation of elytra, evenly curved lower
surface of aedeagus with short, convex apex, and
4-setose parameres.

DESCRIPTION

Measurements, Length: 2.15-2.45mm; width:
1.02-1,10mm. Ratios: Width pronotum/head:
1.56-1.63; width/length of pronotum: 1.36-1.45;
width base/apex of pronotum: 1.38-1.41; width
widest diameter/base of pronotum: 1.12-1.18;
length/width of elytra: 1.37-1.38.

Colour. Head and dark parts of elytra reddish-
piceous to piceous, pronotum, base and apex of
elytra and suture more or less dark reddish, Elytra
at shoulder with a medially rather ill delimited,
about triangular, light reddish spot attaining
about position of 4th stria. Tn posterior 2/5 of
elytra with a rather conspicuous, fairly well de-
limited, oblique, though but slightly s-shaped,
light reddish stripe, attaining about 2nd stria.
Legs yellow to light reddish, barely infuscate.

Head. Median antennomeres c. 1.4 x as long as
wide, Microreticulation distinct on most of head,
weaker only on vertex, abont isodiametric to
slightly transverse. Surface somewhat dull.

Pronotum. Transverse, moderately convex,
widest jn front of middle, base much wider than

355

apex. Latetal margin posteriorly little curved,
though rather oblique, at posterior angles percep-
ПЫу incurved. Posterior angles remarkably ob-
tuse, distinctly produced over lateral part of base.
Carina at posterior angle with elongate, little
oblique, slightly incurved. Anterior transverse
sulcus rather shallaw, faintly interrupted in mid-
dle or at least becoming shallower. Posterior
transverse sulcus fairly deep, interrupted by a
rather large fovea, Anterior lateral seta situated
well behind anterior third of margin.
Microreticulation distinct, almost isodiametric,
surfacesomewwhattull.

Elytra. Moderately short and convex, egg-
shaped, widest slightly in front of middle. Lateral
border evenly curved, extremely finely, almost
not perceptibly, serrate and pilose. 1st and 2nd
striae deeply impressed, crenulate, 3rd-5th striae
in anterior half slightly impressed, outer striae
faintly or barely impressed, though all striae eas-
ily visible as rows of rather fine punctures, that
become finer laterally and apically. Ist and 2nd
intervals convex throughout, 3rd-5th intervals
near base morc or less disiinctly convex. Inner
five striae just visible near арех, 8th stria deeply
impressed Ihroughoul, pun: fate, atfaiming poste-
rior marginal pore. Recurrent striole elongate,
rather oblique, meeting 3rd stria. Anterior discal
pore well behind anterior third, posterior pore in
front of posterior third. Microreticulation dis-
linci, though somewhat superficial, consisting of
rather irregular, moderately transverse meshes.
Surface moderately glossy. Wings markedly
shortened.

Lower surface. Metepisternum about as long as
wide.

Male genitalia. Genital ring regularly tnangu-
lar, moderately wide, apex rather narrow.
Aedeagus small, short and compact, lower sur-
face concave, apex very short, widely rounded
off, Parameres moderately elongate, both 4-se-
tose.

Female genitalia, Stylomere 2 with stout dor-
sal ensiform seta situated rather medially,

Variation. Apart from some allometric varia-
tion of relative width and shape of pronotum,
which in large specimens tends to be relatively
wider, little variation noted.

DISTRIBUTION

Mt. Bellenden Ker, north Queensland. Known
only from this mountain top.

356 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 14. Habitus. A, P. vicina sp. nov. B, P. laevis sp. nov. C, Philipis sulcata sp. nov. D, P. frerei sp. nov. E,
P. subtropica sp. nov. Е, P. alutacea sp. nov. Lengths: 2.2mm; 2.25mm; 2.4mm; 2.3mm; 2.75mm; 2.25mm.

REVISION OF PHILIPIS

HABITAT

Collected by pyrethrum knockdown on mossy
trees and logs and by Berlese extraction of litter,
moss, and sticks in montane rainforest above
1500m. Collected April, June, October, and No-
vember,

ETYMOLOGY
Refers to the generally reddish colour of this
species,

Philipis perstriata sp, nov.
(Figs 15B, 19A)

MATERIAL EXAMINED

HOLOTYPE: ОМТ 13726; 9, Mt. Bartle Frere, МЕО,
Sth. Peak Summit, 1620m, 6-8 Nov 1981, Earth-
watch/Qld. Museum, pyrethrum knockdown.

DIAGNOSIS

Small, reddish-piceous, quadrimaculate, with
reddish pronotum and oblique, slightly sinuate
apical elytral spot, distinguished from P.
rufescens sp, noy, by smaller size, narrower
shape, sulcate elytral striae, and weak
microreticulation of elytra,

DESCRIPTION

Measurements, Length: 2.05mm; width:
0.9mm. Ratios: Width pronotum/head: 1,64;
width/length of pronotum: 1.41; width base/apex
of pronotum: 1,37; width widest diameter/base of
pronotum: 1.17; length/width of elytra: 1.39.

Colour. Reddish-piceous, pronotum, base,
apex, suture, and lateral borders of elytra slightly
lighter, reddish. Elytra behind shoulder with à
large, though very inconspicuous, ill delimited,
about triangular, light reddish spot attaining
about 4th stria. In posterior 2/5 of elytra with an
inconspicuous, very ill delimited, slightly
oblique, narrow, light reddish fascia attaining
about 2nd stria. Legs light reddish.

Head. Median antennomeres c. ].2 x as long as
wide. Microreticulation very distinct, coarse,
only in posterior half somewhat superficial, about
isodiametric to slightly transverse. Surface rather
dull.

Pronotum. Transverse, moderately convex,
widest slightly in front of middle, base much
wider than apex. Lateral margin posteriorly al-
most straight, though rather oblique. Base dis-
tinctly narrower than widest part. Posterior angles
almost rectangular, slightly obtuse, slightly pro-
duced over lateral part of base. Carina in posterior
angle moderately elongate, little oblique, slight! у
incurved, Anterior transverse sulcus fairly shal-

357

low, slightly interrupted. Posterior transverse sul-
cus rather deep, interrupted by a large fovea.
Anterior lateral seta situated well behind anterior
third of margin. Microreticulation very distinct
and coarse, slightly transverse, laterally even
forming short, transverse wrinkles, surface mark-
edly dull.

Elytra, Moderately short and convex, egg-
shaped, widest about in middle. Lateral border
evenly curved, not perceptibly serrate and pilose.
Surface in posterior 2/5 just behind posterior fas-
cia with large, transverse impression. All striae
deeply impressed throughout, all intervals mark-
edly convex. Puncturation of striae very fine,
barely visible. 8th stria deeply impressed
throughout, finely punctate, surpassing posterior
marginal pore. Recurrent striole elongate, mark-
edly oblique, anteriorly suddenly incurved, mect-
ing 3rd stria. Anterior discal pore slightly behind
anterior third, posterior pore well in front of pos-
terior third. All marginal setae rather short.
Microreticulation highly superficial, difficult to
see, consisting of irregular, transverse meshes.
Surface highly glossy. Wings markedly short-
ened. Variation unknown.

Lower surface. Metepistermum about as long as
wide.

Genitalia, Male unknown, Female stylomere 2
with stout dorsal ensiform seta situated rather
medially.

DISTRIBUTION
Mt. Bartle Frere, north Queensland, Known
only from this mountain top.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks in montane rainforest above 1600m.
Collected November,

ETYMOLOGY
Refers to the completely and deeply striate
elytra.

Philipis ellioti sp. nov.
(Figs 7B, 15C, 17B)

MATERIAL EXAMINED

HOLOTYPE: QMT13727; d, 19°30°S, 146^57'E, Mt.
Elliot summit, МЕО, 1 150m, 12 May 1991, D. Cook,
pyrethrum, tree & logs.

PARATYPES: 6 d, 4 9, same data (ANIC, CBM, ОМ,
USNM, ZSM» 1 3,1 9, 19?29'S, 14657 E, Mt. Elliot,
NEQ, North Ck,, 27 Mar 1991, 1000m, GM, Pyr., trees
& logs (ОМ); | d, Mt. Elliot Summit, 30km SW

358

Townsville, NEQ, 13 Dec 1990, AG, Hand Collecting
(QM).

DIAGNOSIS

Small, piceous, quadrimaculate, with ill delim-
ited basal, and oblique, only slightly sinuate api-
cal elytral spot, further distinguished by lack of
microreticulation, rather elongate aedeagus with
evenly curved lower surface, and commonly 4-
setose right paramere.

DESCRIPTION

Measurements. Length: 2.0-2.15mm; width:
0.9-1.0mm. Ratios: Width pronotum/head: 1.50-
1.53; width/length of pronotum: 1,38-1,46; width
base/apex of pronotum: 1.28-1.43; width widest
diameter/base of pronotüm; 1,12-1.16;
length/width of elytra: 1.35-1.41.

Colour. Head and dark parts of elytra piceous,
pronotum, base, lateral border, and suture of ely-
tra lighter, reddish. Behind shoulder with an in-
conspicuous, medially very ill delimited,
circular, light reddish spot. attaining about 3rd
stria, but difficult to tell from dark reddish base.
In posternor 2/5 of elytra with a fairly conspicu-
ous, rather well delimited, transverse, yellowish
spot, attaining about 2nd stria, Legs dark yellow
to light piceous.

Head. Median antennomeres barely longer than
wide, Microreticulation distinct on labrum and
anterior part of clypeus, superficial on frons, re-
duced on vertex, about isodiametric to slightly
transverse, Surface moderately glossy.

Pronotum. Moderately transverse, fairly con-
vex, widest about in middle or slightly in front of
it, base much wider than apex. Lateral margin
posteriorly but feebly convex or even straight,
fairly oblique. Posterior angles rectangular, not
produced over lateral part of base. Canina at pos-
terior angle moderately elongate, fairly oblique,
slightly incurved, Anterior transverse sulcus ex-
ceptionally shallow, not interrupted. Posterior
transverse sulcus deep, interrupted by a rather
large fovea. Anterior lateral seta situated behind
anterior third of margin. Microreticulation very
superficial or almost absent, surface glossy.

Elytra. Moderately short and convex. egg-
shaped, widest slightly behind anterior third. Lat-
eral border evenly curved, not perceptibly serrate
and pilose. Sutural and 2nd striae deeply im-
pressed, crenulate. 3rd-Sth striae still visible as
rows of rather coarse punctures at beast in anlerior
half. Outer striae hardly perceptible, Ist and 2nd
intervals convex, sometimes even 3rd interval
slightly convex near base. Only Ist-3rd striae

MEMOIRS OF THE QUEENSLAND MUSEUM

traceable to apex. 8th stria deeply impressed,
anteriorly slightly shallower, finely punctate,
though antenorly smooth, attaining posterior
marginal pore. Recurrent striole elongate, mod-
erately oblique, meeting 3rd stria. Anterior discal
pore well behind anterior third, posterior pore
slightly in front of posterior third. Microreticula-
tion absent. Surface highly glossy. Wings short-
ened,

Lower surface. Metepisternum c. 1.2 x as long
as wide.

Male genitalia. Genital ring triangular, feebly
asymmetric, rather wide, apex narrow. Aedeagus
rather small, moderately elongate, lower surface
concave, apex short, fairly wide, widely rounded
off. Parameres moderately elongate, right 5-se-
tose, left 4- or 5-setose,

Female genitalia. Stylomere 2 with stout dorsal
ensiform seta situated rather medially.
Variation. Little variation noted, apart from
minor differences in shape of pronotum, shape of
posterior elytral fascia, and number of setae on
left male paramere.

DISTRIBUTION
Mt, Elliot south of Townsville, north Queens-
land. Known only from that mountain top.

HABITAT

Collected by pyrethrum knockdown on mossy
trees and logs in montane rainforest above
1000m, only one specimen by “hand collecting",
Collected March, May, and December,

ETYMOLOGY
Refers to the type locality, Мг, Elliot.

Philipis agnicapitis sp. nov.
(Figs 7C, 15D, 19А)

MATERIAL EXAMINED

HOLOTYPE: QMT13736; d, Lambs Head, 10km W
Edmonton, NEQ, 10 Dec 1989, 1200m, Montcith,
Thompson. Janetzki, р rethrum, logs & trees.
РАВАТҮРЕЅ: 1 d, 1 9, same data (CBM); 1 5,2 9,
Lambs Head, 10km W Edmonton, NEQ, 1200m, 11
Dec 1989 (2nd Tower), GM, GT, HJ, Pyr., logs & trees
(QM, ZSM); | d, 1 (sex?), Lambs Head, 10km W
Edmonton, NEQ, 4 Dec 1988, 1200m, GM & GT,
Pyr./loes & trees (QM); 3 d, 2 9, 17°02'S, 145" 40" E,
Lambs Head, МЕО, (East Епа), 29 Nov 1993, 1 180m,
GM & HJ Pyr/itrecs & logs (ANIC, CBM, QM.
USNM).

REVISION OF PHILIPIS

DIAGNOSIS

Small, reddish-piceous, quadrimaculate, with
ill delimited basal, and only slightly oblique,
barely sinuate apical elytral spot, further distin-
guished by lack of microreticulation, narrow pro-
notum, moderately elongate aedeagus with
evenly curved lower surface, and 5-setose right
paramere.

DESCRIPTION

Measurements, Length: 1,85-2.30mm, width:
U.85-1.05mm. Ratios: Width pronotum/head:
1.50-1.57; width/length of pronotum: 1.39-1.42;
width base/apex of pronotum: 1,34-1,37; width
widest diameter/base of pronotum: 1.13-1.15;
length/width of elytra: 1,34-1,39.

Colour. Reddish-piceous to piceous, pronotum,
apex and suture, sometimes also base of elytra
faintly lighter. Elytra at shoulder with an incon-
spicuous, 111 delimited, about tnangular, lightred-
dish spot attaining about position of 4th stria. In
posterior 2/5 of elytra with a slightly more con-
spicuous, moderately well delimited, transverse,
light reddish spot or stripe, attaining about 3rd or
even 2nd stria. Legs yellowish to light reddish,
tibiae sometimes faintly darker.

Head. Median antennomeres c. 1.2 x as long as
wide, Microreticulation distinct, though fairly su-
perficial on most of head, almost absent only on
vertex, about isodiametric to slightly transverse.
Surface moderately glossy,

Pronotum. Moderately transverse and convex,
widest shghtly in front of middle, hase much
wider than apex. Lateral margin posteriorly little
or almost straight, though rather oblique. Base
usually distinctly narrower than widest part. Pos-
terior angles almost rectangular, barely obtuse,
only feebly produced over lateral part of base.
Carina at posterior angle moderately elongate,
slightly oblique, slightly incurved. Anterior
transverse sulcus fairly deep, feebly interrupted.
Posterior transverse sulcus deep, interrupted hy
large fovea. Microreticulation very superficial,
apparently slightly transverse, surface glossy.

Elytra. Rather short and convex, egg-shaped,
widest slightly in front of middle. Lateral border
cvenly curved, extremely finely, almost not per-
ceptibly, serrate and pilose. Ist and 2nd striae
moderately impressed, rather finely punctate.
3rd-Sth striae more ог less distinctly impressed in
basal half, outer striae very fine, barely percepti-
ble. Puncturation generally rather fine. Ist and
2nd intervals gently convex throughoul, ouler
intervals almost depressed, even in basal half.
Only four inner stnae visible at apex. 81h stria

358

deeply impressed throughout, distinctly punctale,
attaining posterior marginal pore. Recurrent
striole elongate, rather oblique, meeting 3rd stria.
Anterior discal pore well behind anterior third,
posterior pore in front of posterior third.
Microreticulation extremely superficial, percep-
tible only laterally and near apex, consisting of
irregular, transverse meshes. Surface glossy.
Wings shortened.

Lower surface. Metepisternum about as long as
wide.

Male genitalia. Genital ring almost regularly
tnangular, moderately wide wide, feebly asym-
metric, apex rather narrow. Aedeagus rather short
and compact, lower surface fairly concave, apex
narrow, short, rounded off. Upper border sinuate
near apex. Parameres rather short, both 5-setose,

Female genitalia. Stylomere 2 with stout dor-
sal ensiform seta situated rather medially.

Variation. Apart from minor differences in
shape of pronotum and distinctness of elytral
pattern, little variation noted.

DISTRIBUTION
Lainbs Head west of Edmonton, north Queens-
land. Known only from that mountain top.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks and logs in montane rainforest at
1200m, Collected December only.

ETYMOLOGY
Refers to the type locality, Lambs Head.

Philipis picta sp. nov.
(Figs 7D, 156, 19A)

MATERIAL EXAMINED

HOLOTYPE: QMT13744; d , Mt. Finnigan Summit via
Helenyale, МЕО, 3-5 Dec 1990, 1050m, Monteith,
Sheridan, Roberts & Thompson, pyrethrum.
PARATYPES: 3 9. same data (ANIC, СВМ, QM); 2 9,
Mt. Finnigan Summit, МЕО, 30 Nov 1985, GM & DC,
QM Berlesate No. 685, 15°48°S, 145^17'E, RF,
1100m, moss on trees (CBM, QM); 1 9. Mt. Finnigan
Summit, NEQ. 29 Nov 1985, GM & DC, QM Berlesate
No. 699, 15?48' S, 145" 1 E, RF, 1 100m, moss on trees
(QM).

DIAGNOSIS

Rather small to medium-sized, piceous,
quadrimaculate, with light apex and base of elytra
and oblique and sinuate apical elytral spot, further
distinguished from P. bicolor sp. nov. by not
contrastingly coloured pronotum, less contrast-

360 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 15. Habitus. A, Philipis rufescens sp. nov. B, P. perstriata sp. nov. C, P. ellioti sp. nov. D, P. agnicapitis
sp. nov. E, Philipis picta sp. nov. F, P. bicolor sp. nov. Lengths: 2.35mm; 2.05mm; 2.1mm; 2.25mm;2.3mm;
2.3mm.

REVISION OF PHILIPIS

ingelytral pattern, less deeply striate and punctate
base of elytra, and short, on lower surface evenly
curved aedeagus with very short apex.

DESCRIPTION

Measurements. Length: 2.18-2.35mm; width-
1.02-1.04mm. Ratios: Width pronotum/head-
1.48-1.51; width/length of pronotum: 1.45-1.49;
width base/apex of pronotum: 1.35-1.40; width
widest diameter/base of pronotum: 1.12-1.16;
length/width of elytra: 1.31-1.34.

Colour. Head and dark parts of elytra dark
piceous, pronotum, base, and apex of elytra red-
dish-piceous. A fairly vague, triangular, laterally
wide spot behind shoulder attaining medially
about 4th stria, a rather well delimited, oblique,
slightly s-shaped fascia in posterior 2/5 medially
attaining about 2nd stria, and posterior half of
suture yellow to light reddish. Legs yellow, tibiae
and tarsi slightly darker,

Head. Median antennomeres c. 1.3 x as long as
wide. Microreticulation distinct, though superfi-
cial, reduced on vertex, about isodiametric to
slightly transverse. Surface moderately glossy.

Pronotum. Transverse, convex, widest about
in middle or slightly in front of it, base much
wider than apex. Lateral margin posteriorly
slightly convex to almost straight, fairly oblique.
Posterior angles not fully rectangular, slightly
obtuse, feebly produced over lateral part of base.
Carina at posterior angle elongate, markedly
oblique, slightly incurved. Anterior transverse
sulcus deep, not interrupted. Posterior transverse
sulcus deep, interrupted by a rather large fovea.
Anterior lateral seta situated well behind anterior
third of margin. Microreticulation present only at
apex and hase, absent on disk, about isodiametric,
surface glossy.

Elytra. Short. convex, egg-shaped, widest
about in middle. Lateral border evenly curved,
extremely finely serrate and pilose. Sutural stria
crenulate. 2nd-3rd striae, sometimes even 4th
stria. faintly impressed and coarsely, but very
widely spaced punctate in basal half, outer striac
very fine throughout, though usually just visible
at high magnification. All, except of sutual and
2nd stria disappearing behind middle. At least
1st-2nd intervals slightly convex in basal half,
sutural interval convex even to apex, 8th stria
deeply impressed, anteriorly shallow, strongly
punctare-crenulate, just attaining posterior mar-
ginal pore. Recurrent striole elongate, rather
oblique, meeting position of 3rd stria, Amerior
discal pore im anterior third, posterior pore
slightly in front of posterior third. Microreticula-

361

tion rather superficial, composed of irregular.
moderately transverse meshes, Surface fairly
glossy. Wings slightly shortened.

Lower surface, Metepisternum slightly longer
than wide.

Male genitalia. Genital ring triangular, slightly
asymmetnc, moderately wide, apex rather nar-
row. Aedeagus short and compact, lower surface
concave, apex very short and wide, widely
rounded off. Parameres moderately elongate,
both 5-setose.

Female genitalia. Stylomere 2 with stout dor-
sal ensiform seta situated rather medially,

Variation. Apart from some minor differences
in relative shape of pronotum and in distinctness
of elytral pattern, little variation noted.

DISTRIBUTION
Mt. Finnigan south of Cooktown, north
Queensland. Known only from this mountain top.

HABITAT

Collected by pyrethrum knockdown and Berl-
ese extraction of moss on trees in montanc
rainforest above 1050m. Collected November
and December.

ETYMOLOGY
Refers to the striking elytràl pattern,

Philipis bicolor sp. nov.
(Figs 3, 7E, 15F, 19А)

MATERIAL EXAMINED

HoLoryen: QMTI3748; 4, Mt. Demi, 7km SW of
Mossman, NEG, 29 Oct 1983, 100m, D. К. Yeates &
G. L Thompson, Pyrethnum knockdown in RF.
РАКАТҮРЕ5; 11 d, 13 9, same data (ANIC, CBM,
QM, USNM, ZSM).

DIAGNOSIS

Rather small to medium sized, black,
quadrimaculate, with light apex and base of elytra
and oblique and sinuate apical elytral spot, further
distinguished from P. picta sp. nov. by con-
tsastingly red pronotum, markedly contrasting
elytral pattern, more deeply striate and punctate
base of elytra, and rather elongate, on lower sur-
face faintly bisinuate acdeagus with fairly elon-

gate apex.

DESCRIPTION

Measurements. Length: 2.0-2.45mm; width:
0,92-1, 15mm. Ratios: Width pronorum/head;
1.50-1.55; width/length of pronotum: 1.53-1.61;
width base/apex of pronotum: 1.44-1.49; width

362

widest diameter/base of pronotum: 1.07-1.09;
length/width of elytra: 1.32-1.36.

Colour. Head and dark parts of elytra very dark
piceous, almost biack. Pronotum, shoulders,
basal third of inner two or three intervals, apex,
suture, and lateral border of elytra, reddish. A
semicircular, laterally wide spot behind shoulder
attaining medially about 4th stria, and a very
oblique, s-shaped fascia in posterior 2/5 medially
attaining about 2nd stria, yellow. Both spots con-
spicuous and well delimited. Legs yellow.

Head. Median antennomeres c. 1.3 x as long as
wide. Microreticulation distinct only on labrum
and anterior border of clvpeus, on frons very
superficial, absent on vertex, about isodiametric
to slightly transverse. Surface highly glossy.

Pronotum. 'Transverse, convex, widest about
in middle or slightly in front of it, base much
wider than apex. Lateral margin posteriorly fee-
bly convex or almost straight, or even faintly
concave in front of posterior angles, fairly
oblique. Postenor angles rectangular, feebly pro-
duced over lateral part of base. Carina at posterior
angle very elongate, markedly oblique, slightly
incurved. Anterior transverse sulcus deep, not
interrüpted. Posterior transverse sulcus deep, in-
terrupted by a rather large fovea. Anterior lateral
seta situated well behind anterior third of margin.
Microreticulation absent on disk, superficial at
apex and base, surface glossy.

Elytra. Short, convex, egg-shaped, widest
about 1n middle. Lateral border evenly curved,
extremely finely serrate and pilose. Sutural stria
crenulate, 2nd-Sth striae, sometimes even 6th
stria faintly impressed and all striae in basal half
coarsely, but very widely spaced punctate. Three
to four inner striae visible near apex. At least
Ist-4th intervals slightly convex in basal half,
sutural interval convex throughout. Bth stria
deeply impressed, anteriorly shallower, strongly
punctate, anteriorly even crenulate, just attaining
posterior marginal pore. Recurrent striole elon-
gate, rather oblique, meeting 3rd stria. Anterior
discal pore behind anterior third, posterior pore
slightly in front of posterior third. Microreticula-
tion absent. Surface highly glossy. Wings slightly
shortened.

Lower surface. Metepisternum slightly longer
than wide.

Male genitalia. Genital ring triangular, rather
asymmetric, moderately wide, apex rather nar-
row. Aedeagus rather short and compact, lower
surface gently bisinuate, apex fairly elongate,
wide, widely rounded off. Parameres moderately
elongate, both S-setose.

MEMOIRS OF THE QUEENSLAND MUSEUM

Female genitalia. Stylomere 2 with stout dor-
sal ensiform seta situated rather medially.

Variation. Apart from some rninor differences
of size, relative shape of pronotum, and siatinct-
ness of pattern, little variation noted.

DISTRIBUTION
Mt. Demi south of Mossman, north Queens-
land. Known only froin that mountain top.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks in montane rainforest at 1 100m. Col-
lected October only.

ETYMOLOGY
Refers to the strikingly bicoloured pattern.

Philipis geoffreyi sp. nov.
(Figs 7B, 16A, 19B)

MATERIAL EXAMINED

Hotorvre: QMT13767; d, 167228, 145? 1 E, 7km
N, Mt, Spurgeon (Camp 2), NEQ, 17-19 Oct 1991,
1250m, Monteith & Janetzki, Pyrethrum, trees & logs.
PARATYPE; | 9, 167248, 145^13' E, Stewart Ck., 4km
NNE Mt. Spurgeon (Camp 1), NEQ, 1250-1300m, 15
Oct 1991, Pyr., GM & HI (СВМ).

DIAGNOSIS

Rather small, dark piceous, quadrimaculate,
with distinct, triangular subbasal, and oblique and
sinuale apical elytral spots, further distinguished
from P. minor sp. nov. by larger size, wider
pronotum, and shorter elytra.

DESCRIPTION

Measurements. Length: 2.12-2. 16mm; width:
1.02-1.04mm. Ratios: Width pronotum/head:
1.51-1.55; width/Iength of pronotum: 1.55-1.57;
width base/apex of pronotum: 1.42-1.46; width
widest diameter/base of pronotum: 1.08-1.09;
length/width of elytra: 1.29-1.31,

Colour. Dark piceous, pronotum and posterior
half of suture of elytra, sometimes also base of
elytra reddish-piccous. Pronotum distinctly ligh-
ter than head and elytra. Lateral border of elytra,
a triangular, laterally wide spot behind shoulder
attaining medially about 4th stria, and a conspic-
uous, very oblique, s-shaped fascia in posterior
2/5 medially attaining about 2nd stria, yellow,
Both spots conspicuous and well delimited. Legs
yellow.

Head. Median antennomeres c. 1.3 x as long as
wide. Microreticulation distinct, though superfi-

REVISION OF PHILIPIS

cial, reduced on vertex, about isodiametric to
slightly transverse. Surface moderately glossy.

Pronotum. Transverse, convex, widest about
in middle, base much wider than apex. Lateral
margin posteriorly slightly convex to almost
straight, fairly oblique. Posterior angles rectangu-
lar, feebly produced over lateral part of base.
Carina at posterior angle elongate, markedly
oblique, slightly incurved. Anterior transverse
sulcus deep, not interrupted. Posterior transverse
sulcus very deep, interrupted by a large fovea,
Anterior lateral seta situated well behind anterior
third of margin. Microreticulation very superfi-
cial, slightly transverse, surface rather glossy.

Elytra. Short, highly convex, egg-shaped, wid-
estabout in middle. Lateral border evenly curved,
extremely finely serrate and pilose, Sutural stria
crenulate. 2nd-Sth stnae slightly impressed and
coarsely punctate at Jeast in basal half, outer striae
very fine even in anterior half. All, except of
sutual and 2nd stria disappearing behind middle,
At least 1st-3rd intervals slightly convex in basal
half, sutural interval convex throughout. 8th stria
deeply impressed, anteriorly shallow, strongly
punctate-crenulate, just attaining posterior mar-
ginal pore. Recurrent striole elongate, rather
oblique, meeting position of 3rd stria. Anterior
discal pore in anterior third, posterior pore
slightly in front of posterior third. Microreticula-
tion distinct, though somewhat superficial, com-
posed of rather irregular, moderately transverse
meshes. Surface moderately glossy, Wings
slightly shortened.

Lower surface. Metepisternum slightly longer
than wide.

Male genitalia. Genital ring asymmetrically
triangular, fairly narrow, apex rather widc.
Aedeagus moderately short and compact, lower
surface almost imperceptibly bisinuate, apex
short, rounded off. Parameres rather elongate,
both 5-setose.

Female genitalia. Stylomere 2 with stout dor-
sal ensiform seta situated rather medially.

Variation, Apart from some minor differences
in depth of ground colour, little variation noted.

DISTRIBUTION
To the north of Mt, Spurgeon, western part of
Carbine Tableland, north Queensland.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks tn montane rainforest above 1250m
Collected October.

363

ETYMOLOGY

Named in honour of the collector of most
specimens of the genus Philipis, Dr. Geoffrey
Monteith of the Queensland Museum.

Philipis minor sp, nov.
(Figs 8A, 16B, 19B)

MATERIAL EXAMINED

HOLOTYPE: QMT13768; d, Mossman Bluff Track.
10km W Mossman, МЕО, 21 Dec 1989, 1180m, Mon»
teith & Thompson, pyrethrum (Site 8).

DIAGNOSIS

Small, dark piceous, quadrimaculate, with dis-
tinct, triangular subbasal and oblique and sinuate
apical elytral spot, further distinguished from P.
geoffrey) sp. nov. by lesser size, narrower pronn-
tum, and longer elytra,

DESCRIPTION

Measurements. Length: 2.0mm; width:
0.96mm. Ratios; Width pronotum/head: 1,45;
width/length of pronotum: 1.47; width base/apex
of pronotum: 1.34; width widest diameter/base of
pronotum: 1.11; length/width of elytra; 1.35.

Colour, Dark piceous, posterior half of suture
and apex of elytra reddish-piceous. Pronotum not
lighter than head and elytra. Lateral border of
elytra, à triangular, laterally wide spot behind
shoulder attaining medially about 4th stria, and a
conspicuous, very oblique, s-shaped fascia in
posterior 2/5 medially attaining about 2nd stria,
yellow. Both spots conspicuous and well delim-
ited. Legs yellow,

Head. Median antennomeres c. 1.3 x as Jong as
wide. Microreticulation superficial, reduced on
vertex, about isodiametric to slightly transverse.
Surface moderately glossy.

Pronotum, Fairly transverse, convex, widest
well in front of middle, base much wider than
apex. Lateral margin posteriorly straight, fairly
oblique. Posterior angles rectangular, feebly pro-
duced over lateral part of base. Carina at posterior
angle elongate, markedly oblique, slightly in-
curved. Anterior transverse sulcus deep, not in-
terrupted. Posterior transverse sulcus very deep,
interrupted by a large fovea. Anterior lateral seta
situated well behind anterior third of margin.
Microreticulation very superficial, slightly trans-
verse, surface rather glossy.

Elytra. Rather short, highly convex, egg-
shaped, widest about in middle, Lateral border
evenly curved, extremely finely serrate and pi-
lose. Sutural stria crenulate, 2nd-Sth striae fairly
impressed and coarsely punctate at least in basal

364 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 16. Habitus. A, P. geoffreyi sp. nov. B, P. minor sp. nov. C, Philipis sinuata sp. nov. D, P. distinguenda sp.
nov. E, P. lustrans sp. nov. F, P. quadraticollis sp. nov. 9 styli. Lengths: 2.65mm; 2.6mm; 2.4mm; 2.15mm;
2.0mm.

REVISION OF PHILIPIS

half, outer striae very fine even in anterior half,
All, except of sutural and 2nd stria disappearing
behind middle. At least 1st-4rd intervals slightly
convex in basal half, sutural interval convex
throughout. 8th stria deeply impressed through-
out, strongly punctate-crenulate, attaining poste-
rior marginal pore. Recurrent stnole elongate,
rather oblique, meeting position of 3rd stria, An-
terior discal pore in anterior third, posterior pore
slightly in front of posterior third. Microreticula-
thon distinct, though somewhat superficial, com-
posed of rather irregular, moderately transverse
meshes. Surface moderately glossy. Wings
slightly shortened. Variation unknown.

Lower surface. Metepisternum slightly longer
than wide,

Genitalia. Male genital ring asymmetrically
triangular, moderately wide, apex rather narrow,
Acdeagus rather short and compact, lower sur-
face distinctly bisinuate, apex short, wide, widely
rounded off. Parameres moderately elongate,
both 5-setose, most posterior seta on lower sur-
face rather removed from penultimate. On upper
side behind last seta with an additional puncture,
but without seta. Female genitalia unknown.

DISTRIBUTION
Mountains west of Mossman, north Queens-
lund. Known only from type locality.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks in montane rainforest at 1180m, Col-
lected December only.

ETYMOLOGY
Refers to the smaller size compared With the
closely related species P. geoffrey.

Philipis sinuata sp. nov.
(Figs 8B, 16C, 19B)

MATERIAL EXAMINED

HOLOTYPE: QMT13769; d, Mt. Bartle-Frere, МЕС,
summit creek, 24 Sept 1981, G. Monteith & D. Cook,
CM Berlesate No. 304, rainforest, 1500m, sieved litter.
PARATYPES: 7 d, 2 9, same data (CBM, QM): 2 d.
Mt Bartle Frere, NEQ, Sth. Peak Summit, 1620m, 6-8
Nov 1981, EW/QM Pyr. (CBM, ОМ); 1 d, Mt Bartle
Frere, NEQ, Campsite, 1500m, 27 Dec 1989, GM, Pyr.,
trees & rocks (QM); 1 9?, Mt, Bartle Frere, МЕО,
NW-Centre Peak, 16 Sept 1982, 1500m, GM & SM
(QM); 2 3,4 9, Bellenden Ker, Centre Peak Summit,
NEQ, 10 Apr 1979, GM, QM Berlesate No, 8, 17°16°S,
145°51'E, RF, 1500m, moss on trees (QM, ZSM); 1 2,
Bellenden Ker Summit, МЕО, 10 June 1980, GM, QM
Berlesate No, 221, 17°16'S, 145?52"E, RF, 1561m,

365

moss on trees (QM): 7 d, 4 9, Bellenden Ker Range,
NEQ, Summit TV Stn, 1560m, 17 Oct-5 Nov 1981,
EW/QM Pyr. (ANIC, СВМ, QM, USNM, ZSM): 2 d.
Bellenden Ker Range, МЕО, Summit TV Stn., 1560m,
25-31 Oct 198), EW/QM, QM Berlesate No. 373,
17°16'S, 145°51°E, RF, moss on trees (ОМУ, 2 &. 1
9, Bellenden Ker Range, NEQ, Summit TV Stn..
1560m, 1-7 Nov 1981, EW/QM, QM Berlesate No,
346, VI^ L6'S, 145"5|' E, RF, moss on trees & rocks
(СВМ, ОМ); 1 d, Bellenden Ker Range, МЕО, Sum-
mit TV Sin, 29 Apr-2 Мау 1983, GM, DY, QM Berles-
ate No. 562, 17^16'S, 145°51'Е, RF, 1560m, moss
(QM); 32 4, 16 2, Bellenden-Ker Range, МЕС), Sum-
mit TV Sin., 1550m, 28 Oct 1983, GM, DY & GT, Pyr,
in RF (CBM, QM); 1 d, 2 9, 17"16'S, 145°52°B,
Bellenden Ker summit, МЕО, 8 Oct 1991. 1560m, GM
& HJ, Pyr., trees & logs (QM).

DIAGNOSIS

Large, dark piceous, quadrimaculale, with
oblique, markedly sinuate apical elytral spot, fur-
ther distinguished by distinct microreticulation
on frons and elytra, wide pronotum with remark-
ably wide base, short aedeagus with faintly
curved lower surface, and S-setose right
paramere,

DESCRIPTION

Measurements. Length: 2.6-2.75mm; width:
1.2-].3mm, Ratios; Width pronotum/head: 1,65-
1.73; width/length of pronotum: 1.51-1.56; width
base/apex of pronotum: 1.51-1.59; width widest
diameter/base of pronotum: 1.09-1.11;
length/width of elytra; 1.34-1.36,

Colour. Dark piceous, pronotum, base of elytra
and sulure slightly lighter. Elytra at shoulder with
a rather ill delimited, triangular, light reddish spot
attaining about position of 4th stria. In posterior
2/5 of elytra with a conspicuous, well delimited,
markedly s-shaped, yellowish stripe attaining
about sutural stria. Mecdio-anteriorly this stripe is
markedly bent and encircles the posterior elytra!
pore. Both elytral pores situated within a small,
light spot. Legs yellow, barely infuscate.

Head, Median antennomeres c. 1.8 x as long as
wide. Microreticulation very distinct on labrum
and anterior part of clypeus, distinct, though
fairly superficial on frons and vertex, about iso-
diamctric to slightly transverse. Surface fairly
glossy.

Pronotum. Transverse, rather convex, widest
behind middle, base much wider than apex. Lat-
eral margin posteriorly little curved, though
rather oblique, al posterior angles perceptibly
incurved. Posterior angles rectangular, though
somewhat obtuse, distinctly produced over lat-
eral part of base. Carina at posterior angle elon-

gate, markedly oblique, slightly incurved. Ante-
riortransverse sulcus rather deep, not interrupted,
Posterior transverse sulcus deep, interrupted by a
large fovea. Anterior lateral seta situated well
behind anterior third of margin. Microreticula-
tion. distinct, though superficial, almost isodia-
metric, surface rather glossy.

Elytra. Moderately short and convex, egg-
shaped, widest at or slightly in front of middle.
Lateral border evenly curved, extremely finely,
almost not perceptibly, serrate and pilose. Sutural
stria crenulate. 2nd stria distinctly impressed in
basal half, 3rd-5th striae at base still faintly im-
pressed, outer striae not impressed, though all
striae easily visible as rows of rather coarse punc-
tures that laterally and apically become finer. Ist
interval convex throughout, 2nd-4th intervals
near base more or less distinctly convex, Inner
four striae just visible near apex. 8th stria deeply
impressed throughout, strongly punctate, attain-
ing posterior marginal pore. Recurrent stnole
elongate, rather oblique, meeting 3rd stria. Ante-
rior discal pore well behind anterior third, poste-
rior pore in front of posterior third,
Microreticulation distinct, though superficial,
consisting of rather irregular, transverse meshes,
Surface moderately glossy. Wings shortened,
though still surpassing elytra,

Lower surface. Metepisternum about as long as
wide.

Male genitalia. Genital ring asymmetrically
triangular, narrow, apex rather wide. Aedeagus
rather large, though short and compact, lower
surface feebly concave, apex very short, widely
rounded off. Parameres moderately elongate,
both 5-setose.

Female genitalia. Stylomere 2 with stout dor-
sal ensiform seta situated rather medially.

Variation. A rather homogenous species, There
is, however, apparently some allometric varia-
tion, since large specimens tend to have relatively
wider pronotum with wider base,

DISTRIBUTION
Bellenden Ker Range, north Queensland.

HABITAT

Collected by pyrethrum knockdown or mossy
trees and by Berlese extraction of moss and litter
in montane rainforest above 1500m. Species oc-
curs only on the very summits of the range-

ETYMOLOGY
Refers to the sinuate, markedly s-shaped pos-
tenor elytra) fascia,

MEMOIRS OF THE QUEENSLAND MUSEUM

Philipis distinguenda sp, nov,
(Figs 8C, 16D, 19B)

MATERIAL EXAMINED

HOLOTYPE: QMT13833; с, Mossman Bluff Track
10km W Mossman, МЕО, 17 Dec 1988, 1200m, Mon-
teith & Thompson, pyrethrum/trees & logs,
PARATYPE: 1 д, Mossman Bluff Track 6km W Moss-
man, МЕО, 16 Dec 1988, 480m, GM & GT, Pyr./trees
& logs (CBM).

DIAGNOSIS

Large, dark piceous, quadrimaculate, with
oblique, markedly sinuate apical elytral spot, fur-
ther distinguished from closely related P. sinuata
sp. nov. by narrower pronotum with narrower
base, longer elytra, and slightly longer aedeagus
with gently bisinuate lower surface and slightly
longer apex.

DESCRIPTION

Measurements, Length: 2,6-2.65mm; width:
1.18-1.24mm. Ratios: Width pronotum/head:
1.59-1.60; width/length of pronotum: 1.45-1.47;
width base/apex of pronotum: 1.45-1.47; width
widest diameter/base of pronotum: 1.10-1.12;
length/width of elytra: 1.38-1.41.

Colour, Head and dark parts of elytra rather
dark piceous, pronotum, basal third, apex, lateral
borders, and suture of elytra slightly lighter, red-
dish-piceous. Elytra at shoulder with a very ill
delimited, triangular, yellowish spot attaining
about position of 4th stria. In posterior 2/5 of
elytra with a conspicuous, well delimited, mark-
edly s-shaped, yellowish stripe medially attaining
about 2nd stria. Medio-anteriorly this stripe is
sharply bent and touches the posterior elytral
pore, Legs yellow, tibiae faintly darker.

Head. Median antennomeres c. 1.8 x as long as
wide. Microreticulation distinct on Jabrum and
anterior part of clypeus, fairly superficial on frons
and vertex, about tsodiametric to slightly trans-
verse. Surface fairly glossy.

Pronotum. Transverse, rather convex, widest
about in middle, base much wider than apex.
Lateral margin posteriorly little curved, though
rather oblique, al posterior angles perceptibly
incurved, Posterior angles rectangular, though at
tip slightly obtuse, distinctly produced over lat-
eral part of base. Carina at posterior angle elon-
gale, markedly oblique, slightly incurved,
Anterior transverse sulcus rather deep, not inter-
rupted. Posterior transverse sulcus deep. inter-
rupted by a large fovea. Anterior lateral seta
situated. well behind anterior third of margin.

REVISION OF PHILIPIS

Microreticulation distinct, though superficial, al-
most isodiametric, surface rather glossy.

Elytra. Moderately short and convex, egg-
shaped, widest at or slightly in front of middle.
Lateral border evenly curved, extremely finely,
almost not perceptibly, serrate and pilose. Ist and
2nd stnae deeply impressed, crenulate. All other
striae also more or less impressed in basal half,
Striae rather coarsely punctate-crenulate, thou
puncturation laterally and apically finer. | st and
2nd intervals convex throughout, 3rd-5th inter-
vals in basal half rather distinctly convex, outer
intervals just feebly convex. Inner four striac
visible near apex. 8th stria deeply impressed
throughout, strongly punctate, attaining posterior
marginal pore. Recurrent striole elongate, rather
oblique, meeting 3rd stria. Anterior discal pore
well behind anterior third, posterior pore in front
of posterior third. Microreticulation distinct,
though superficial, consisting of slightly irregu-
lar, transverse meshes. Surface moderately
glossy. Wings slightly shortened, though still sur-
passing elytra,

Lower surface. Meteptsternum c. 1.2 x as long
as wide.

Genitalia. Male genital ring triangular, narrow,
slightly asymmetric, apex rather wide. Aedeagus
short and compact, lower surface feebly bisinu-
ate, apex very short, widely rounded off,
Parameres, especially right paramere, short and
wide, both 5-setose. Female genitalia unknown.

Variation. Due to few available specimens little
variation noted.

DISTRIBUTION

Track to Mossman Bluff west of Mossman,
north Queensland. Known from only two locali-
ties on this track.

HABITAT

Collected by pyrethrum knockdown on mossy
trees and logs in rainforest at 480m and 1200m
height. Collected December only.

ETYMOLOGY
Refers to the close external similarity with P.
sinaata.

Philipis lustrans sp. nov.
(Figs 8D, I6E, 19B)

MATERIAL EXAMINED

Ног.отүРЕ: QMT13834; 5, North Bell Peak, NEQ,
Malbon Thompson Ra., 20 Nov 1990, 1000m, Mon-
leith & Thompson, pyrethrum-Irees & logs.

367

PARATYPES: 6 С, 3 ©, same data (ANIC, CBM, QM,
USNM, ZSM); 3 4,3 9, Bellenden Ker Range, NEQ,
Cable Tower 3, 1054m, 17 Oct-5 Nov 1981, EW/OM
Pyr, (CBM, QM); 1 d, 17°16°5, 145°50'Е, Mas-
sey/Bellenden Ker saddle, NEQ, 9 Oct 1991, 950m,
GM, HJ & DC, Pyr., trees & logs (QM); 1 d, 1 9,
17°14’S, 145°48°E, Massey Ra., МЕО, 6 km NW of
Bellenden Ker, 1 150m, 11 Oct 1991, Pyr., GM & HJ
(QM); 1 6, Mt Bartle Frere, МЕО, West Side, 1050m,
8 Dec 1990, GM, GT & RS, Pyr.-trees & rocks (ОМ);
4 d, Lambs Head, 10km W Edmonton, МЕО, 10 Dec
1989, 1200m, GM, GT & HJ, Pyr., logs & trees (QM),
4 3,1 9, Lambs Head, 10km W Edmonton, МЕГ),
1200m, 11 Dec 1989, (2nd Tower) GM, GT & HJ, Pyr.,
logs & trees (CBM, QM).

DIAGNOSIS

Medium-sized, dark piceous, quadrimaculate,
with oblique, markedly sinuate apical elytral
spot, further distinguished from related species
by narrower base of the wide pronotum, absence
of microreticulation on frons and elytra, and short
aedeagus with gently bisinuate lower surface,

DESCRIPTION

Measurements. Length; 2.2-2,5mm; width:
1,0-1.16mm. Ratios: Width pronotum/head:
1.56-1.62; width/length of pronotum: 1.51-1.57;
width base/apex of pronotum; 1.46-1.51; width
widest diameter/base of pronotum: 1.09-1.13;
length/width of elytra: 1.33-1.37.

Colour. Fairly dark piceous, suture, sometimes
also base and apex of elytra and even pronotum
faintly lighter. Elytra behind shoulder with a
moderately conspicuous, medially fairly ill de-
limited, about triangular, light reddish spot attain-
ing about position of 4th stria. In posterior 2/5 of
elytra with a conspicuous, well delimited,
oblique, markedly s-shaped, yellowish or light
reddish fascia attaining sutural stria. The sharply
bent median part of the fascia situated well in
front of posterior discal seta. Legs yellowish to
light reddish,

Head, Median antennomeres c. 1.75 x as long
as wide. Microreticulation very indistinct, highly
superficial on most of head, absent on vertex,
about isodiametric to slightly transverse, Surface
highly glossy.

Pronatum. Rather transverse and convex, wid-
est about in middle, base much wider than apex.
Lateral margin posteriorly little curved, rather
oblique. Base distinctly narrower than widest
part. Postenor angles almost rectangular, slightly
obtuse, faintly produced over lateral part of base.
Carina at posterior angle elongate, rather oblique,
slightly incurved. Anterior transverse sulcus
deep, not interrupted. Posterior transverse sulcus

368 MEMOIRS OF THE QUEENSLAND MUSEUM

a Р. atra thompsoni
^ P. subtropica ield R. vicina
4 laevis
ellioti

CAIRNS
,MOSSMAN

C a
T TA
e
Barron Н

32 other species LY
in the Wet Tropics
(see detailed maps)

1
П
I
|
!
|
!
i
|
!
[j
]
1
i
!

| BRISBANE

quadraticollis
ij Bloamtleld А. ^ P. alticola

m v P. cooki nia
oP. laevigata

_ 7 Daintree А.

фонии
TA

Mt Finnigan
Thornton Pk.
Windsor Thld,
Carbine Tbld
Hann Thid.
Black Min.
Lamb Ra.
Walsh Ra
Hugh Nelson Ra
. Atherton ТЫЧ
Mt. Bellenden Ker
. Malbon Thompson Ra
Graham Ra.
Waller Hill Ra
Kirrama/Cardwell Ra
. Seaview Ra.
. Hinchinbrook Is.

ү а Ji 184 . Paluma/Bluewater Ra.
C july R Mt Elliot

5

1

2.
3,
4.
5.
б,
7.
B

9.

FIG. 17. Distribution of Philipis species. A, Queensland showing P. atra sp. nov. and P. subtropica sp. nov. plus
combined extent of 33 tropical species. B, Wet tropics region showing P. thompsoni sp. nov., P. vicina sp. nov.,
P, laevis sp. nov., P. ellioti sp. nov. and the 19 rainforest survey zones used in the collection program. C, P.
quadraticollis sp. nov., P. alticola sp. nov., P. cooki sp. nov. and P. laevigata sp. nov.

REVISION OF PHILIPIS

deep, interrupted by large fovea. Anterior lateral
seta situated well behind anterior third of margin,
Microreticulation very superficial to barely per-
ceptible, apparently slightly transverse, surface
highly glossy.

Elytra, Moderately short and convex, egg-
shaped, widest slightly in front of middle, Lateral
border evenly curved, extremely finely serrate
and pilose. Ist and 2nd striae moderately im-
pressed throughout. 3rd-Sth striae more or less
distinctly impressed in basal half, even outer
siriae sometimes faintly impressed, and all striae
rather coarsely punctate. 1st and 2nd intervals
gently convex throughout, 3rd-Sth intervals
gently convex in basal half, sometimes even outer
intervals slightly convex, though apically de-
pressed. Only four to five inner striae visible at
apex. 8th stria deeply impressed throughout, dis-
linctly punctate, attaining posterior marginal

, Recurrent striole elongate, rather oblique,
mecting 3rd stria. Anterior discal pore about in
anterior third, posterior pore about in posterior
third. Microreticulation extremely superficial,
just perceptible only laterally and near apex, con-
sisting of irregular, transverse meshes. Surface
highly glossy. Wings slightly shortened, though
yet longer than elytra.

Lower surface, Metepisternum е, 1.2 x as long
as wide.

Male genitalia. Genital ring almost regularly
wiangular, moderately wide, feebly asymmetric,
apex rather narrow. Acdeagus very short and
compact, lower surface basally faintly bisinuate,
1n apical third curved down, apex rather narrow,
short, rounded off. Apical part of aedeagus con-
spicuously covered with large microtrichia.
Parameres rather short, both 5-setose.

Female genitalia. Stylomere 2 with stout dorsal
ensiform seta situated rather medially.

Variation. Apart from minor differences in
shape of pronotum and degree of microrcticula-
tion on pronotum and elytra, little variation noted.

DISTRIBUTION

Mountains at and near the eastern border of
Atherton Tableland in the Lamb, Bellenden Ker,
and Malbon Thompson Ranges, north Queens-
land.

HABITAT

Collected by pyrethrum knockdown on mossy
tree trunks, logs, and rocks in montane rainforest
between 950-1200m.. Collected October to De-
cember.

ETYMOLOGY
Refers to the glossy surface as compared with
related species.

NATURAL HISTORY

Available data indicate that all species of
Philipis are arboreal and inhabit rainforest living
on mossy tree trunks. Most specimens were cap-
tured by pyrethrum spraying of the moss on the
trunks of rainforest trees, Even those specimens
that have been collected by Berlese extraction
were sampled from moss litter from vertical sur-
faces. Almost no specimens have been collected
by hand sampling which is perhaps due to their
small size and their habits under the cover of the
moss. Only one specimen has been so far found
under bark. This could be due to failure of appro-
priate sampling, but more likely it reflects their
absence from thal habitat.

Despite the time, energy and inspiration de-
voted by G, Monteith and his collegues, the num-
ber of available specimens remains small,
excepting a few more common species. Difficul-
tes in obtaining such specimens may be
illustrated by the fact that P. J, Darlington, cer-
tainly a skilful collector in the rainforests of Aus-
tralia, recovered only a single specimen of this
genus during his long stay in the urea (actually his
son collected the specimen). Occurrence of al-
most all specics high on mountain tops, some of
which are rather remote, makes their collection
difficult. In general, collecting work means a
strenuous ascent often through dense montane
rainforest to mountain tops, and careful sampling
was hindered hy wet weather. Perhaps these hec-
tles are actually more common than we know, and
additional appropriate sampling work may bring
to light even more species and much more mate-
rial,

Most collecting took place from October to
December just before onset of the wet season.
Thus the actual main activity period of these
beetles is unclear. According ta С. Monteith
(pers. comm.) they are less common during the
dry season, hence I think that the activity period
of these beetles is the wet season and the few
months before, when storms begin to bring same
rain. No specimens have been so far recorded àt
light, although several species are fully winged
and may be able to fly. According to G. Monteith
no specimen has been found in the large series of
flight intercept traps that were run during the
sample periods, Hence, species of Philipis appar
entlv fly very rarely, But flight may be of no

370 MEMOIRS OF THE QUEENSLAND MUSEUM

, COOKTOWN

heatherae oe a P. ruficollis
^ P. picea | CjsemtesR A P, inermis
"P.castanea "5 F v P. reticulata —

2 P. unicolor Cw OP. striata

Lo

w L {1429 —
B LU жи
o

a P. sulcata
Bomis R. A P, frerei
v P. alutacea ы
2 P. rufescens

= P. planicola

4 P, spurgeoni

Y P. trunci e
2 P, tribulationis

C

CAIRNS

FIG. 18. Distribution of Philipis species. A, P. heatherae sp. nov., P. picea sp. nov., P. castanea sp. nov. and P.
unicolor sp. nov. B, P. ruficollis sp. nov., P. inermis sp. nov., P. reticulata sp. nov. and P. striata sp. nov.. C,
P. planicola sp. nov., P. spurgeoni sp. nov., P. trunci sp. nov. and P. tribulationis sp. nov. D, P. sulcata sp. nov.,
P. frerei sp. nov., P. alutacea sp. nov. and P. rufescens sp. nov.

REVISION OF PHILIPIS

в P. perstriata
^ P. agnicapitis
" P. picta

о P. bicolor

| Bloomfiela Н.

|
\,
X
X

X. j }

A,

Sükms

371

сооктон

n

|
145°

a P. geoffreyi

^ P. minor

v P. sinuata

2 P. distinguenda
P. lustrans

t=

MAREEBA^ ,
Vy

Baron RAL, „

S
À € R.
nay “vere

ont

Br wa

50kms

FIG. 19. Distribution of Philipis species. А, P. perstriata sp. nov., P. agnicapitis sp. nov., P. picta sp. nov. and
P. bicolor sp, nov. B, P. geoffreyi sp. nov., P. minor sp. nov., Р, sinuata sp, nov. and P. lustrans sp. nov.

advantage for beetles living in dense montane
rainforest. Many species, however, possess re-
duced wings and are flightless.

Nothing is known on habits and life cycles, on
diet, enemies, mating and reproduction, and no
larvae have been detected, Since none have been
seen runnning about, nothing is known on the
daily period of activity, perhaps they are rather
nocturnal.

On several occasions, and even within the same
samples, different species have been observed
occurring together. However it is unknown,
whether they were from the same tree, because
most samples include material from several trees.
Nothing is also known about their possible pref-
erence for different tree or moss species.

A striking feature that will be important for the
following biogeographical treatment is the oc-
currmence of almost all species (except for P.
planicola and P. striata) in montane rainforest
mainly above 900m. A further feature is the ex-
tremely limited ranges of most species that have
been observed usually on only a single mountain
top, far less commonly on a mountain range or on
a group of adjacent ranges. Only one species (P.
thompsoni) occupies a really wide range through-

out a larger part of the overall wet tropics region
of north Queensland.

In conclusion it must regrettably be stated that
we still know extremely little about the natural
history of these beetles in spite of the enormous
collecting efforts of Geoffrey Monteith and his
co-workers.

RECONSTRUCTED PHYLOGENY

PHYLOGENETIC STATUS

Philipis is a genus of basically plesiomorphic
tachyine beetles which nevertheless shows some
spezialized or apomorphic characters. Erwin
(1973) when revising the similarly partly arboreal
neotropical Xysfosomus, noted some of these
primitive characters that are likewise present in
the Australian Philipis. In a numerical cladistic
analysis of the higher categories of Bembidiini -
using the Hennig 86 program - Erwin (pers.
comm.) found that Philipis keys out as the sister
group of all the New World xystosomines. This
would affirm the plesiomorphic status and per-
haps also the close relationship of both lineages
because plesiomorphic species of the Xystosomus
lineage exhibit some character states present only

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Character states and their phylogenetic value in Philipis.

[3 [Surfaceofelyra |
4

Number of elytral striae present |all striae

develope

É Depth of striae

Shape of strae

Apical striole meets 3rd stria

Colour of space around discal
elytral punctures

Aedeagus

in Bembidion sensu lato, but generally not in
Tachyini. Should this close relation be proved,
this would have a major impact on biogeographic
and evolutionary questions of the tribe Tachyini
in general.

PHYLOGENETIC RELATIONS

Certainly all species of Philipis are closely
related as indicated by their uniform appearance,
colour, pattern, and external and genitalic mor-
phology. Hence it is difficult to deal with the
phylogenetic relations at the specific level, be-
cause parallel evolution presumably has been a
very common event, and, on the other hand, strik-
ing morphological differences are rare and pres-
ent only in few highly aberrant species. In my

resent and well

| [Character — |plesiomorphie_ — — à — apomorphe — 1 5 |
medium-sized to large Маут

[i [size — j|
Peter fener

short, convex
' - very short and highly convex

impressed in anterior third

Tth-Sth striae reduced, less distinct
^ ~7th-5th striae absent

" - only two inner striae present
4'" - only sutural stria present

LA

|
[5 [Lengthofstriae | striae fully developed striae postenorly shortened |

striae deeply impressed

only innér 4 striae impressed

6' - only inner 3 striae impressed

б - only inner 2 striae impressed
"= on

sutural stria impressed

[7 _[Puncturation of striae [striae punctate [striae impunctate
2nd-7th striae normally shaped |all striae deeply sulcate

elongate, curved inwards,

shortened, not curved inwards, not meeting 3rd
stria

lio | Microreticulation of pronotum | present, distint | reduced or absent

il Microreticulation of elytra present, distinct reduced, su
11" - completely absent

[12 |Colourofpronotum _ |blackish-piceous

m Colour of elytra blackish-piceous 138 r iat ж,
| Elytral pattern present, consisting of an apical |14a- present, consisting of à basal and an apical
macula macula each

14b - absent

15 | Shape of posterior elytra] macula | сігсшаг ог slightly transverse — |elongate, oblique
15' - markedly s-shaped 15*

Distinctness of elytral maculae [rather inconspicuous distinct, sharply bordered

1 б ассраша, set off by a conspicuous dark
ackgroun

Colour of sutural interval contrastingly reddish |

normal shaped, with short
apex, lower surface straight

rficial

19а - apex elongate
19b- lower surface sinuate or bisinuate
I9c - very short, compact, curved
9d - apex elongate, slender, at base incised

attempted evaluation of phylogenetic relations,
several morphoclines from plesiomorphic to
more or less highly apomorphic states were
found, but in many cases it is obvious that evolu-
tion of a comparable or even the same
apomorphic character status has occurred twice
or even several times. In other characters it is still
uncertain, whether apomorphic stages of the
morphocline are synapomorphic, i.e, whether
species exhibiting the same apomorphic charac-
ter states are actually related. Such states which
have been certainly evolved independently are
marked with numbers in the character matrix
below.

For better comparison a summary of the mea-
sures and ratios of all species is included that

REVISION OF PHILIPIS

illustrates some of the differences in size and
body shape (Table 1). Although the characters
used and their presumed plesiomorphic and
apomorphic states are shown in Tables 2 & 3,
some characters, especially those that build up
morphoclines, are explicitly discussed below.

1. Size, Comparison with other genera of
Tuchyint, especially Xystesontus and allies, con-
vinces me that very small size is apomorphic
rather than plesiomorphic. Small size may be
advantageous for a beetle living in the often short
moss growing on tree trunks. Size reduction may
have occurred repeatedly in different lineages of
the genus.

2. Shape of elytra. The more elongate, de-
pressed shape of the elytra in the quadraticollis-
group (P. guudraticollis, P. cooki, P. laevigata,
Р. alticola, P. heatherae, and, to a lesser degree,
also P. castanea and Р, picea) is presumably the
plesiomerphic state, and the short and usually
highly convex elytra of certain species - best
evolved in P. sulcara - are apomorphic, Further-
more the convex shape of the elytra is commonly
combined with reduction of elytral striation and
extensive reduction or even complete loss of
microreticulation, which are both likewise
apomorphic states. The tendency to evolve a
rather globular body shape js perhaps generally
common to moss- and fungus-inhabiting beetles.
But which advantage the smooth, glossy integu-
ment should bear, is uncertain, Perhaps it has à
water-repellent function,

4-8. Development of elytral siriaz. Certainly
the full set of elytral striae is the plesiomorphic
state, as in other genera, Complete reduction of
striae, or shortening at base and apex, or reduction
of depth of striae are therefore apomorphie states
that make up morphoclines to the final state,
where only the sutural stria is present. Originally
the striae are punctate or punctate-crenulate,
therefore reduction of puncturation, but at the
same time channelling of all striae, are different
apomorphic states that may eventually lead to
elytra that have all striae deeply channelled but
impunctate. Reduction of striae is commonly
combined with loss of microrcticulation, but in
some species it is not. All these reductions pre-
sumably occurred independently in different tin-
eages, withoul our exact knowing of how many
times they occurred.

9, Form of apical striole, Although the lateral
position of the apical striole is presumably a
plesiomorphic character of the whole genus, the
plesiomorphic state within the genus may be a
curved apical striole that mects the 3rd stria or its

373

position. The almost straight, barely incurved
striole in some species may be therefore а second-
ary, apomorphic state.

10, 11, Loss of microreticulation. Certainly
apomorphic, but it may be duc to parallel evolu-
tion, like most other reductions, in several spe-
cies,

12, 13. Colour of surface, Uniformly piceous
or blackish colour is perhaps the plesiomorphic
state, whereas vivid colouration of pronotum
and/or elytra are apomorphic states, the advan-
tage of which, however, is obscure,

14-18. Elytral pattern. Itis uncertain, whether
unspotted or uniplagiate elytra are the
plesiomorphic status in Philipis, since there are
good reasons for both opinions, Ovt-group com-
parison of the closest relatives reveal that species
of the Xystesomus lineage are generally unspot-
ted, but in some other tachyine genera presence
of an apical spot seems 10 be the plesiomorphic
status. In the species of the apomorphic species
group of Philipis that combines rather short, con-
vex species, however, both unspotted species (P.
unicolor, P, atra) are in some other respects
rather apomorphic (markedly short, convex hod y
shape, reduction of striation, absence of
microreticulation). In these species at least loss of
elytral pattern is perhaps secondary.

The plesiomorphic state of the elytral pattern
when it is present is certainly the unimaculate
one, with the apical macula about circular of at
most vaguely transverse. The bimaculate patlern
and the various oblique to markedly s-shaped
forms of the apical macula, and the pronounced
colour of the maculae on a dark background are
apomorphic states that in some lincages form
morphoclines. Conspicuous colouralions of su-
ture and of discal punctures are found in single
species only and are special features of these
species.

Therefore, the basic branching of the clado-
gram is rather dependent on deciding whether
absence of elytral pattern is plesiomorphic or
apomorphic. Thus a basic branching different
from that chosen in the cladogram cannot be
completely excluded, although it seems to me less
probable,

19. The aedeagus. Basically very similar
throughout the genus. Some unusual modifica-
tions of shape and form of apex are apomorphic,
but may be special features or autapomophic
states of single species only and thus of little
value for the reconstruction of the phylogenetic
relations.

374

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 3. Character states in the species of Philipis, numbered as in Table 2. -: plesiomorphic state; 1, 2 etc.:
apomorphic stale, different apomorphic states distinguished by lower case letters. States of a morphocline

indicated by: ', ", ""
marked by numbers: !, ? etc.

| quadraticoilis| | |

The cladogram (Fig. 9) erected on the basis of
these character states seems to give quite good
results but extreme caution is required during
interpretation, because it certainly has some par-
allelisms. Also, some of the opinions expressed
below are based on mere similarities rather than
on well founded synapomorphies. On the other

| рь; is
РЕБЕР Ааа CC CC fis пив |

Spurgeoni m

= de] tet | e n [D uw pe ШШ use]
bua [rp а Is e| | — L—E- This ie ]- ее]
Hep E a БЕСЕ Je E du ju T кшш D heels | pss
a Бего ИГУ нана пана инр: esi ГЇ

Г нете БЕ Г УСЕ tec p ГС раа dme ШЕШ TO
a cl AP 111 а
жи LIT III LI e = [Г В Be]
ава | ГЕ БЕ ГЕГИ ГАШ a ГТОЕ e [ГГ иши шиш ип шп Г2
baies | ПЕ ШИ ГЕ ГЕ te LTT ШЕН БЕШ ШЕШ Г77Ш БШШ ШЕШ БЕ ИШ ise]
a | а ра ae se TI
ee OL ede а 1 | [|
Du [Гэ єє [ш ш il [е Пе [[_
bees | bI] je] [| he -] i Tua hs he] T T. —
ы e EL tel hah Bg tee je is fe. | | T
bir | la| la ls le i ue [n fm ] ha ера
kara [Т ole ls lel o ua he he TT I ]
E E i5 hi ТГ]
Pt aisle ш e 11]
ear ШЕГИ ин ШЕГУГЕ БЕ ЕЕ И mL [ bas De

ашта | B | fa fof 1 ии} | а e er |] n]

* Comparable or similar apomorphic states that have been likely convergently evolved are

| he ү)

hand, some branchings are certainly disputable
due to the validity of the supporting synopomorp-
ies. The character of the cladogram is thus highly
speculative, as may also be seen by the existence
of several trichotomies or polytomies. Therefore
some groupings that seem better founded are
explicitly discussed below.

REVISION OF PHILIPIS

1. quadraticollis-group. The five species of this
group apparently combine mostly plesiomorphic
character states, hence it is uncertain whether
they are really closely related, Perhaps they con-
stitute a cluster of more or less primitive species.
Within the group P. cooki, P. laevigata, and P.
alticola are perhaps closely related, and they are
perhaps also related to P. heatherae, P.
quadraticollis, ón the other hand, is presumably
the most plesiomorphic species of this group and
of the whole genus.

2. picea-group. Both unpatterned though still
elongate species P. picea and Р, castanea are
perhaps more highly evolved than the species of
the guadraticollis-group and form a distinct
group of uncertain affinities. АП other species are
perhaps related by their more or less short and
convex shape of the elytra. They divide into three
distinct groups, the unicolor-, trunci-, and
sinuata-groups, respectively, the relationships of
which, however, are fairly obscure.

3. unicolor-group. The two unpatterned spe-
cies Р, unicolor and P. atra form a monophyletic
unit, when absence of pattern is actually an
аротоғрһіс state, Р, ruficollis is a rather isolated
species but may be next related to both,

4. trunci-group. The unimaculate species of the
Irunci-group perhaps form a monophyletic
group, although this is at present based on a rather
weak character. Certainly, however, this group
includes the most plesiomorphic species within
all patterned species excluding those of the
quadraticollis-group, which is Р. reticulata. P.
inermis does not fit well in this group and pos-
Sesses several autapomorphic character stales.
Apart from its unimaculate elytral pattern, if does
not share many characters with the rest of the
species of the trunci-group. These, however, are
all combined by the synapomorphic, s-shaped
form of the elytral spot.

Within this latter assemblage Р. trunci, P.
tribulationis, P. thompsoni, and P. vicina are
closely related and apparently constitute a mono-
phyletic group. P. planicola and P. spurgeoni
may be slightly less apomorphic, but are related
to the above group by some apparent syn-
apomorphic character states, All these species
combine perhaps to be the sister group of P.
striata, and together with P. striata they form the
sister group of P. reticulata which seems to rep-
resent the most primitive species of the whole
trunci-group.

5, sinuata-group, The rest of the species from
Р, laevis apparently form again a monophyletic
unit, though the relationships within this group

are highly speculative. P. laevis and again Р.
sulcata are highly specialized, autapomorphic
TA that may be related, but whose relation-

ips are obscure. P. frerei, Р. alutacea, and Р:
subtropica may form à separate group wich is
perhaps the more primitive sister group af the rest
of the species. P. frerei and P. subtropica share
the apparently synapomorphic shape of the
aedeagus, but the three mentioned species may he
actually not very closely related.

The remaining species presumably form a
fairly well-defined monophyletic group, P. per-
striata and P. rufescens may be related to anc
another, but both are fairly aberrant species, per-
haps rather remotely related to the rest of species,
whereas all other species may form a monophy-
letic unit. Р, ellioti and P. agnicapitis may be in
some respects more plesiomorphie than the re-
maining species which all possess a markedly
sinuate apical elytral spot, Certainly Р, picta and
P, bicolor are closely related due to their similar,
highly complex pattern. P. geoffroyi and P. minor
may form another group, but this is uncertain. The
last three species (P. sinuata, P. distinguenda,
and P. lustrans) are again closely related and are
distinguished only by minor characters of shape,
microreticulation, pattern, and structure of
acdeagus.

To conclude, the present 35 species can pre-
sumably be referred to only five different stocks,
three of which are perhaps more closely related
to each other, than any to the quadraticollis-

group.

DISTRIBUTION

Despite of the large number of recorded spe-
cies, the material is still insufficient for a final
review of the distributions, Many species are still
known from a single locality or mountain top
without our knowing whether this apparent lim-
ited range is due to inappropriate sampling, or
whether it reflects the actual range. Therefore,
any considerations about ranges and why (he
ranges of certain species differ to such large ex-
tent are somewhat hypothetical,

Nevertheless, 1 have tried to point out the
current knowledge of the species ranges in the
following tables which list altitude ranges (Table
4) and the recorded species for each locality
(Table 5). The known localities are also depicted
in maps that are based ор a subdivision of the wet
tropics region of north Queensland in 19 different
mountain/rainforest blocks according to the clas-
sification used by G. Monteith for his survey of
the rainforest fauna of north Queensland (Fig.

376

TABLE 4. Recorded altitude range of known species of
Philipis (altitudes below 500m in bold type).

[Species [altitude [Species [altitude |
| quadraticollis |1000-1180m [vicina [850m — |

alticola [1560т — — levis — [1000-1300 |
cooki 1100-1330m |480-1300m |

laevigata — |1100-1250m |rerei — — |1500-1620m]
[hearherae — (1050т ^ |subiropica |1100-1400m |
picea — |8G0m aluacea — |1500-1560m |
(castanea |780-1300m |rufescens — |1500-1560m |
perstriata [1620m |
Jara [900m [ellioti [1000-1150m]
(ruficollis [R50-1100m [аркар [1200m — |
memi: [1100m [picta — — [1050-1100m |
(reticulata [1000m [bicolor — [1100m |
uri [150600m__|geoffreyi — |1250-1300m |
planicola [5-100m O| minor 1180m
|spurgeoni [1330m [sinuata — |1500-1620m |
munci — — [1000-1300m | distinguenda |480-1200m |
[950-1200 | 1200m
FRI

680-1225m [lustrans — |
[ihempsoni — |]00-1250m | |

17B). Each rainforest block 1s defined by more or

less natural barriers as depicted in the maps. In a
Final list the occurrence of the species in these
rainforest blocks is listed (Table 6). As a sum-
mary of those lists and maps the following con-
clusions may be drawn:

1, 33 of 35 known species occur in northeastern
Queensland, especially in the mountains al the
eastern fringes of the Atherton and Carbine Ta-
blelands, The northern border of the range of the
genus is on Mt. Finnigan, south of Cooktown.
South of the tablelands few species occur on somc
scattered mountain tops only (Mt, Macalister, ML
Elliot, Mt. Macartney, Lamington Plateau,
Spnngbrook Plateau). Surprisingly enough, no
Philipis has been thus far recorded on the promi-
nent plateaus of Mt. Spec south of Ingham and
Eungella west of Mackay, although G. Monteith
has sampled both plateaus, Hence the main centre
of diversity of the genus is certainly the Carbine
'Tableland and the Bellenden Ker Range of north-
eastern Queensland.

2, Almost all species occur in montane rainfor-
est above about 900m, and only two species (P.
planicola, P. striate) have been recorded at or
near see level, whereas another two species ex-
tend as far down as ahout 500m, Bul in the
respective areas this is already at the level where
montane rainforest exisis. Many species occur

MEMOIRS OF THE QUEENSLAND MUSEUM

regularly above 1000m, and surprisingly many
species live only on the highest tops of the respec-
tive mountains, On Bellenden Ker and Bartle
Frere for example, those species apparently do
nnt descend below 1500m. Thus Philipis is pri-
marily a genus of montane species and the few
lowland species have probably reached this level
secondarily.

3. Of 35 recorded species, 23, that is more than
two thirds, arc from a single locality or mountain
top (Table 5). They are especially common on
the isolated tops of Mt, Finnigan, Thornton Peak,
Mt. Spurgeon, on the Carbine Tableland, on Mt.
Bellenden Ker and Mt. Bartle Frere, and again in
the southern part of the range of the genus. On the
Atherton Tableland, on the other hand, only one
widespread species (P. fhompsoni) occurs but
only on the mountains that make up the westem
border of the tableland. A single apparently en-
demic species has been found on the Lamb Range
at the eastern fringe of the Tableland. So the fauna
of the Atherton Tableland is surprisingly poor and
is therefore in sharp contrast to the ranges east of
the tableland with their nich, endemic faunas,

Three species (P. castanea, Р. cooki, Р, sul-
cata) that have been collected at several locali-
ties, nevertheless occur on a single range or
tableland only (Cape Tribulation- Thornton Peak
arca, and Carbine Tableland area, respectively).
A1 the current state of knowledge they are en-
demic to the respective range.

Three further species (P, striata, P.
tribulationis, P. sinuata) occur on two adjacent
ranges or tablelands which are separated hy a
river or creek valley thal reaches below 500m.
The first wo species live in the Cape Tribulation
and Windsor Tahleland, and Cape Tribulation
and Carbine Tableland areas, respectively, which
are both separated by the Daintree River, P,
sinuata occurs on Mt. Bellenden Ker and Mt.
Bartle Frere that belong to a common range, but
are separated hy the deeply incised valley of
Babinda Creek in the Babinda area. Actually P.
Sinuata occurs only at high altitude (above

1500m) an both mountain tops, A further species,
P. quadraticollis, is somewhat enigmatic, be-
cause it has been found on the Carbine Tableland
and again in the Massey Range at the western
slope of the Bellenden Ker Range. This is indeed
a very wide gap, but the single Iwo known spec-
imens of this species are both females, therefore
| am not entirely convinced that they are actually
conspecific.

Another two species (Р. lustrans, Р. unicolor)
occur on several adjacent ranges (Malbon

REVISION OF PHILIPIS

Thompson Range, Lamb Range, Massey Range,
Bellenden Ker Range, or Malbon Thompson
Range, Bellenden Ker Range, and Mt. Bartle
Frere, respectively) that are separated by the Mul-
grave River, Little Mulgrave River, and Babinda
Creck valleys, respectively, but nevertheless are
part of a former common tableland.

One species, P. planicola, is outstanding, be-
cause it has been recorded only at very low alti-
tude in the eastern foothills of the Bellenden Ker
Range, where it is an inhabitant of lowland
rainforest, Even so this species seems to be local-
ized in this area.

Only a single species (P. thompsoni) is really
wide-ranging. It occurs on almost all ranges sur-
rounding the Atherton Tableland from Kuranda
in the north, but not in the Bellenden Ker and
Bartle Frere ranges at the eastern border of the
tableland. It extends beyond the Atherton Table-
land to the south in the Tully area and further
south in the Card well Range.

In conclusion it should be stressed that almost
all species are endemic to one mountain top or
range, or limited to a svstem of adjacent ranges
or tablelands that certainly formed previously à
common range. However, only one species is
actually widespread, and another, somewhat du-
bious species occurs apparently on two widely
separated ranges. Ву far most species occurin the
Carbine and Bellenden Ker/Bartle Frere rainfor-
est blocks (Fig. 10), remarkably fewer in the
isolated Finnigan and Thornton blocks, in thc
Lamb Range block west of Cainrs, and on the
coastal Malbon Thompson Range, but single spe-
cies only are scattered through the other rainfor-
esl zones.

A similar general distribution pattern has been
found in flightless, ground-living carabids that
inhabit montane rainforest, namely of the genera
Notonomus and Trichesternus (Darlington
1961a, b), and according to G, Monteith (pers.
comm.) also in the (likewise flightless) Ten-
ebrionidac of the subfamily Adeliinae, and in
Aradidae. These are all groups of low agility that
inhabit either the forest floor or tree trunks and
logs, In all groups the Bellenden Ker Range and
the Carbine Tableland possess by far the richest
fauna of endemic species, whereas some species
of the Atherton Tableland are widespread and
occur on several mountain tops. Hence this dis-
tribution pattern and the underlying biogeograph-
ical events are very similar in several insect
groups of low vagility including Philipis.

3T

TABLE 5. List of localities and the respective recorded
species (species occurring on a single locality or
mountain top printed in bold types).

[Mt.Finnigan (ruficollis, inermis, picta

o c—
castanea, trunci, laevis
tribulationis
Carbine Tableland сан
| Mt. Spurgeon

Devils Thumb 7 аа. naki: edi.
Plane Crash Site

Mossman Bluff & cooki, picea, striata, sulcata,
Track minor, distingue.

| Pauls Luck [sulcata

(м. Demi EN cooki, bicolor

[Mt Lewis cooky |
[мите Sout [etemo |
[Lambs Hess
Cs |
мрз jim —
расу Hills _|hentherne,thompront_]
TETTE ws soni

alticola, из reticulata,

| Mt. Bellenden Ker alutacea, rufescens, sinuaia,
lustrans

quadraticollis, thompsoni, lustrans

| m Ker = Range колор

[Rossen River а]
pat Вале нав no parei pei, at rerei, perstriata wen |
fins atest |
=

Mt. Fisher — С=С |
ы == =
Bipper tatei СЕ ерні = =)
ee _—

| Mt. Macartne ee Ü
E ptor Plateau Subrropica

rngbrook Plateau |subrropica —

BIOGEOGRAPHY

Philipis is essentially tropical. However, it is a
phylogenetically plesiomorphic and, therefore,
old genus, perhaps closely related to the, likewise
tropical South American Xystosomus lineage. If
that phylogenctic status 1s right, according to its

378

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 6. Mountain/rainforest blocks of north Queensland and the recorded species of Philipis (for names of
zones see Fig. 17B. Two species (P. atra and P. subtropica) do not occur within the tabulated area.

uadraicolis | — | — |
LL |

uem

кезе ГГ
—L

spurgeoni

| trunci

mai |
bei | X x |
| [ LJ

| rufescens

pes
agnicapilis

peoffreyi
minor
sinuata

mum

distribution Philipis may constitute a part of the
ancient tropical-subtropical "old Gondwanan
faunal element" (Howden, 1981; Baehr, 1990b).
Nevertheless, it is unknown where and in which
environment the original stock of Philipis sur-
vived in Australia during the very long time of
changing environmental conditions of the late
Mesozoic and early Tertiary periods.

Although the genus is widely distributed, the
most plesiomorphic species still occur in north-

m eee

vicina «Г ү ү у |

| 1] | L Инн ШЕШ ИШ ШЕН
cL MEE cote a a =
е" ит |
=, а.
жый кг Г LE E
i

sl SR I PATT | ПЕЕ E

eastem Queensland. Hence, the genus may al-
ways have lived in more or less tropical rainfor-
est. It is unknown, however, whether the
moss-inhabiting way of life is the original one, or
has been adopted later. It is likewise unknown,
whether the genus has been always as montane as
it is today, since most ranges and tablelands that
it inhabits at present have been built up or uplifted
comparative recently, during the Pliocene and
Pleistocene.

REVISION OF PHILIPIS

Phylogenetic evidence reveals the existence of
two main present centres of diversity and species
richness, namely the Carbine Tableland and the
eastern fringe of the Atherton Tableland (espe-
cially the Bellenden Ker and Bartle Frere ranges),
with several pairs of rather closely related species
occurring in both areas (P. cooki-P. alticola, P.
striata-P. planicola, Р. tribulationis-P.
thampsoni, P. distinguenda-P. sinuata). The spe-
cies that are presumably most plesiomorphic also
occur in these areas, Within both areas the degree
of endemicy is fairly low because several species
occur on adjacent ranges or tablelands, but almost
no species occurs in both areas. The single excep-
lion is P, quadraticollis ihat occurs on the Car-
bine Tableland and in the Bellenden Ker Range.
It is perhaps the most plesiomorphic species of
the whole genus, but the specific status of both
populations (that are both known by only a single
female specimen) is not yet fully seitled.

From the Carbine Tableland the rather isolated
monutain tops of Mt. Spurgeon, Thornton Peak,
and Mt. Finnigan (as well as Mt. Misery) have
been colonized, All three tops are inhabited by
three species endemic to the respective mountain,
but the fauna of the most isolated Mt. Finnigan is
phylogenetically most isolated, because P.
ruficollis and P. inermis do not posses a close
relative, and the third species, P. picta, is related
to P. bicolor of Mt. Demi only. The faunas of
Thornton Peak and Mt. Spurgeon, on the other
hand, are more vclosely related to those of the
eastern parts of Carbine Tableland. The species
which occur on all three tops however belong to
different stocks, thus showing that there have
been three different colonisation or speciation
eVents on each top alihough those on Mt,
Finnigan were perhaps the oldest.

In the ranges at the eastern fringe of the
Atherton Tableland the situation is rather simple
with: (1). Some species inhabiting several adja-
cent ranges (Lamb Range, Malbon Thompson
Range, Bellenden Ker Range); (2). Additional
species with a slightly more limited range; (3).
Some species occurring only on the Bellenden
Ker Range: and (4). Several species endemic to
cither Mt. Bellenden Ker or Mt. Bartle Frere. This
distribution pattern may reflect populations of
different age that have been isolated to a different
degree. In the southern part of the range of the
genus the species densily is low, but as in the
northem part of the range, all species are endemic
to à single mountain top and most aré more or less
closely relaied іо a species occurring further
north.

379

Although the phylogenetic analysis is not con-
clusive, it seems as if the faunas of both Carbine
Tableland and of the ranges at the eastern fringe
of Atherton Tableland are composed of both
fairly wide-ranging, rather plesiomorphic specics
and usually endemic, apomorphic species with
limited ranges. With regard to the existence of
some pairs of closely related species in both
areas, it may be suggested that the former ranges
of the stocks of several recent species have been
wide and have been rather recently interrupted.

According to the patterns of distribution, the
phylogenetic evidence, and the known paleogeo-
graphic, climatic, and floristic history of north-
eastern Australia, the history of the genus may be
thus hypothetized as following: Philipists an old
and in certain respects very plesiomorphic genus
that lived presumably always under subtropical
and perhaps even tropical conditions in rainforest
and may be a part of the so-called "Old
Gondwanan element", because its nearest relati ve
is perhaps the tropical South American
Xystosomus lineage. However, most species
scem to be young according to their very similar
extemal and genitalic structures, The high species
diversity of the pe: may be thus a rather recent
event and may be caused by the combination of
three geologic and climatic factors:

(1) Uplift of much of the Great Dividing Range
during Pliocene and Pleistocene and its following
disintegration into more or less isolated tops and
tablelands due to rapid erosion.

(2) Gradual climate change of Australia to a
warmer and dner climate with more pronounced
dry seasons as the Australian block drifted north-
wards. While lowlands became drier and hotter
and less favourable for rainforest living animals,
mountain tops remained cooler and wetter be-
cause of the altitude and of orographic rains from
the nearby sea.

(3) Repeated and more radical climatic
changes during the “ice age" when rainforests
retreated up the mountains to form small, iso-
lated, pockets during glacials when the sea level
was low and the mountains were far away from
the coast, and descended again during the inter-
glacials to become more or less continuous for-
ests,

The effects of all three events were superirn-
posed оп з gradual retreat of the rainforest to the
mountain tops and a general isolation of rainfor-
est pockets during late Tertiary and the
Pleistocene glacials, interrupted only by a tempo-
rary but repeated spreading of rainforest during
the Pleistocene interglacials. lt seems thal these

380

climatic and floristic events repeatedly separated
and rejoined formerly wide species ranges, that
can be postulated for example for the Carbine
Tableland and the mountains east to the Atherton
Tableland on the basis of several pairs of closely
related species occurring in both areas, and thus
supported allopatric speciation on single moun-
tain tops or ranges by geographical isolation. As
aconsequence, more than two thirds of the extant
Species are endemic to a single mountain top.
According to phylogenetic data, the present 35
species can be referred to only 5 original stocks,
representatives of which occur in both main cen-
tres. Hence speciation presumably occurred
mainly by vicariance of populations separated by
uplifting of ranges or tablelands, either due to
development of barners through rapid erosion of
formerly more homogenous areas, or 10 dismem-
bering and isolation of formerly continuous
rainforest areas due to climatic changes.

At the northern and souther borders of the
Tange however, some speciation may have oc-
curred by colonisation of isolated tops rather than
by parainsular reproductive isolation,

Main speciation events (producing the present
species) perhaps occurred as late as during
Pleistocene, whereas less ample speciation
(which perhaps led to the founding species of the
main species groups or of the main branchings
within the species groups) occurred in late Terti-
ary.

In this context the question arises, whether the
species of this genus had been formerly as "mon-
tane" as they are at present, or whether thay have
trapped on the mountain tops, where they still
persist. If Philipis is actually an old genus and at
the same time an "Old Gondwanan element", then
it lived for a long period under warm temperate
to subtropical circumstances, until Australia -
presumably at the end of Miocene - finally arrived
at its present position. Only then the genus or the
original stocks may have adapted to tropical cli-
mate. Therefore, it is conceivable that the genus
escaped full tropical conditions in refugia on the
highest mountain tops with their more temperate
climatic conditions. On these reasons I believe,
that the genus has been montane for a long time,
and Өз its present montane, moss-living habits
are old.

ACKNOWLEDGEMENTS

| thank Mr. C, Vogt (Cambridge/
Massaachusetts) and Mr. T. Weir (Canberra) for
the kind loan of types and specimens. Special

MEMOIRS OF THE QUEENSLAND MUSEUM

thanks are due to Dr. G. Montcith (Brisbane)
who, assisted by his colleagues Doug Cook,
Heather Janetzki, and Geoff Thompson, collected
almost all specimens studied and kindly loancd
them for examination. Geoff Monteith provided
critical comments on the manuscript while Kann
Koch assisted in its editing. Geoff Thompson
carried out the excellent full illustrations in Figs
2 and 3.

LITERATURE CITED

ANDREWES, H. E. 1925, A revision of the Oriental
species of the genus Tachys, Annali del Museo
civico di Storia naturale Giacomo Dona, Genova
51: 327-502.

BAEHR, M. 1986. Review of thew Australian species
of the genus Tachyta Kirby (Coleoptera, Car-
abidae, Bembidiinae), Entomofauna 7: 305-314.

1987, A review of the Australian Tachyine beetles
of the subgenera Tachyura Motschoulsky and
Sphaerotachys Müller, with special regard to the
tropical fauna (Insecta, Coleoptera, Carabidae,
Bembidiinac). Spixiana 10: 225-269,

1989. A new species of the Tachys ectromoldes-
group from Western Australia (Coleoptera, Car-
abidae, Bembidiinae). Spixiana 12; 279-283.

19908. Revision of the Australian Groand-beetle
Genus Tasmanitachoides Erwin (Insecta: Col-
eoptera: Carabidae: Bembidiinae), with Special
Regard to the Tropical Species. Invertebrate Tax-
олоту 4: 867-894,

1990h, Grundzüge der Faunenzusammensetzung
und Faunengeschichte Australiens, dargestellt
am Beispiel der Laufküfer (Coleoptera, Car-
abidac). Mitteilungen der Deutschen Gesell-
schaft für Allgemeine und Angewandte
Entomologie 7:619-625,

1991. Tachys windyorensis, spec. nov. from North
Queensland, a further new species of the Tachys
ectromoides-group (Insecta, Coleoptera, Car-
abidae, Bembidiinae). Spixiana 14: 189-192.

BRUNDIN, L. 1966. Transantarctic relationships and
iheir significance, as evidenced by chironomid
midges, with a monograph of the subfamilies
Podonominae and Aphroteniinae and the austral
Heptagyinae. Kunglisk svenska Vetenskan
Akadademi Handlingar 1 1: 1-472.

DARLINGTON, P. J. Jr. 1957. Zoo phy: The
geographical distribution of animals. (Wiley: New
York).

196ta. Australian Carabid beetles VIT.
Trichosternus, especially the tropical species.
Psyche, Cambridge 68: 113-130.

196]b. Australian Carabid beetles IX. The tropical
Notonomus. Breviora. Museum of Comparative
Zoology 148: 1-14.

1962. Australian Carabid beetles XI. Some Tachys,
Psyche, Cambridge 69: 117-128.

REVISION OF PHILIPIS

1963. Australian Carabid beetles XII. More Tachys.
Psyche, Cambridge 70: 22-33.

ERWIN, T. L. 1970. A reclassification of bombardier
beetles and a taxonomic revision of the North and
Middle American species (Carabidae:
Brachinida). Quaestiones entomologiae 6: 4-215.

1972. Two new genera of Bembidiine Carabid Bee-
tles from Australia and South America with
Notes on their phylogenetic and zoogeographic
Significance (Coleoptera). Breviora. Museum of
Comparative Zoology 383: 1-19.

1973. Studies of the Subtribe Tachyina (Coleoptera:
Carabidae: Bembidiini), Part I: A Revision of the
Neotropical Genus Xystosomus Schaum.
Smithsonian Contributions to Zoology 140: 1-39.

1974a. Studies of the Subtribe Tachyina (Col-
eoptera: Carabidae: Bembidiini), Part П: A Revi-
sion of the New World-Australian Genus
Pericompsus LeConte. Smithsonian Contribu-
tions to Zoology 162: 1-96.

1974b. Studies of the subtribe Tachyina (Col-
eoptera: Carabidae: Bembidiini). Supplement A:
Lectotype designation for New World species,
two new genera, and notes on generic concepts.
Proceedings of the Entomological Society of
Washington 76: 123-156.

1975. Studies of the Subtribe Tachyina (Coleoptera:
Carabidae: Bembidiini), Part II: Systematics,

381

Phylogeny, and Zoogeography of the Genus
Tachyta Kirby. Smithsonian Contributions to Zo-
ology 208: 1-68.

1994, Arboreal beetles of tropical forests: The
Xystosomini group, subtribe Xystosomina (Col-
eoptera: Carabidae;: Bembidiini). Part I. Charac-
ter analysis, taxonomy, and distribution. The
Canadian Entomologist 126: 549-666.

HENNIG, W. 1966. Phylogenetic Systematics. (Uni-
versity of Illinois Press: Urbana).

HOWDEN, H. F. 1981. Zoogeography of some Austra-
lian Coleoptera as exemplified by the
Scarabaeoidea. In: A. Keast (ed.): Ecological Bio-
geography of Australia, 35: 1007-1033. (W. Junk:
Leiden).

ROSS, H. H. 1974. Biological systematics. (Addison-
Wesley Publishing Co.: Reading).

SLOANE, T. G. 1921. Revisional notes on Australian
Carabidae. Part VI. Tribe Bembidiini. Proceed-
ings of the Linnean Society of New South Wales
46: 192-208. WATROUS, L. E. & Q. D.

WHEELER. 1981. The out-group comparison method
of character analysis. Systematic Zoology 30: 1-
11.

WILEY, E. О. 1981. Phylogenetics. The Theory апа
Practice of Phylogenetic Sytematics. (Wiley: New
York).

COMMUNAL NESTING IN THE SMALL SKINK,
LAMPROPHOLIS ADONIS. Memoirs of the Queensland
Museum 38(2) 382. 1995:- Communal nesting is well-known
amongst Lampropholis spp. It has been observed frequently
in two species, L. delicata and L. guichenoti (Greer, 1989 &
references therein). Its occurrence in the recently described L.
adonis Ingram, 1991, is therefore predictable.

On 12 November, 1994, a communal nest (Fig. 1) contain-
ing 53 skink eggs was found in Bulburin State Forest
(24?31'S, 151?29'E). The eggs had been laid in a pocket of
insect frass and decaying vegetation between sheets of corru-
gated iron. The ‘nest’ was in a small, well-shaded clearing in
complex notophyll vine forest, 5m from the forest's edge. L.
adonis was the most abundant skink species at this site.
Several specimens of this species were seen beneath the
corrugated iron, and had been observed at this locality pre-
viously, associated with a communal nest (S. Wilson, pers.
comm. ). To verify that L. adonis had laid the eggs in the ‘nest’,
seven eggs were taken from the edges and centre of the egg
cache and incubated at room temperature in moist vermiculite.

The eggs varied in length from 9.54mm to 10.76mm (n-7,
mean-9.91), in width from 7.07mm to 7.94mm (n=7,
mean-7.61) and weighed between 0.30g and 0.38g (n-7,
mean-0.33). They hatched over a 24 hour period on 25th (4
eggs) - 26th (3 eggs) January, 1995. All seven hatchlings were
identified as L. adonis. As the eggs were collected widely

MEMOIRS OF THE QUEENSLAND MUSEUM

across the nest and hatched over a short time frame, it is
reasonable to assume that the entire egg cache was deposited
by L. adonis, and that the eggs had been laid more or less
simultaneously. The maximum clutch size recorded for
Lampropholis (L. delicata) is seven eggs (Greer, 1989: Table
9). Presuming L. adonis produces similar sized clutches to L.
delicata, a cache of 53 eggs would represent the nesting effort
of at least eight females.

The hatchlings (QMJ59313-59315, QMJ59331-59334)
measured 17.12mm to 20.36mm SVL (n=7, mean = 18.86)
and had a total length of 39.47mm to 43.18mm (n=7, mean =
41.57). They weighed (preserved weight) between 0.17g and
0.20g (n=7, mean=0.18). All hatchlings were similarly
marked to adults of this species (Ingram, 1991),

Literature Cited

Greer, A.E., 1989. ‘The biology and evolution of Australian
Lizards.’ (Surrey Beatty & Sons: Chipping Norton.
264pp.

Ingram, G.J., 1991. Five new skinks from Queensland
rainforests. Memoirs of the Queensland Museum
30(3):443-453.

Couper, P.J., Queensland Museum, PO Вох 3300, South

Brisbane, Queensland 4101, Australia; Schneider, C.J., Zo-

ology Dept., University of Queensland, St Lucia, Queensland

4072, Australia; 25 August 1995.

FIG. 1. À communal nest containing 53 L. adonis eggs,
Bulburin SF.

A NEW ISCHNACANTHID ACANTHODIAN FROM THE LATE SILURIAN
(LUDLOW, PLOECKENSIS ZONE) JACK FORMATION,
NORTH QUEENSLAND

C.J. BURROW & A.J. SIMPSON

Burrow, C.J. & Simpson, AJ, 1995 12 01: A new ischnacanthid acanthodian from the Late
Silurian (Ludlow, ploeckensis Zone) Jack Formation, north Queensland. Memoirs of the
Queensland Museum 38(2) 383-395. Brisbane. ISSN 0079-8835,

Scales, fin spine fragments, two symphysial tooth whorls, a dentigerous jaw bone fragment,
and dentition cones of a new acanthodian species comprise the total acanthodian component
from samples of the Jack Formation. The morphological variety of scales includes three
normal body scale types, two specializes head scale types, pore scales and tesserac. The
range of scale types, the shape of the fin spine fragments, and the form of the dental elements,
indicate they are from а new species of ischnacanthid acanthodian. [ ] Acanthodii, Jack
Formation, Gomphonchus, ischnacanthid, Ludlow, microvertebrates, scales.

CJ. Burrow, Department of Zoology, University of Queensland, Australia 4072; A.J,

Simpson, Geology Museum, University of Queensland 4072, Australia; 1 October 1995.

Acanthodian scales are common in
microVertebrate assemblages from Upper Silur-
ian and Devonian limestones throughout Aus-
tralia, but until recently there have been no
systematic descriptions of them. Burrow (1995a)
described severa) new taxa from the late
Lochkovian/early Pragian Connemarra Forma-
tion and Gleninga Formation (including the
Jerula Limestone Member) of central New South
Wales. Tumer & Pickett (1982) were the first to
report Acanthodian scales from the Silurian of
Australia. Turner (1991, 1993) recorded various
occurences of Acanthodian remains from Silur-
ian deposits in eastern Australia, including mate-
rial described in this paper. The present paper
describes the microvertebrate assemblage in an
acetic acid-treated samples from the Jack Forma-
tion of northem Queensland. The acanthodian
component appears to derive from a single spe-
cies, The only other vertebrate element in the
section is a scale of the primitive osteichthyan
Lophosteus cf. L. superbus, the type material of
which is from Pridoli deposits in the Baltic. Ma-
terial is lodged in the Geology Museum, Univer-
sity of Queensland (UQY).

STRATIGRAPHY

Vertebrate remains were recovered from sam-
les processed to extract conodonts from the Jack
ormation (Simpson, 1983, 1994b), the upper-

most unit of the Graveyard Creek Group, Broken
River Province, north Queensland (Fig. 1), The
Group consists of the predominately siliciclastic
Crooked Creck Conglomerate, Quinton Forma-

tion and Poley Cow Formation, and siliciclastics
and carbonates of the Jack Formation.

The Jack Formation, originally the Jack Lime-
stone Member of the Graveyard Creek Formation
(White, 1959), previously included only carbon-
ate lithologies exposed in the area around the
Broken River where it is cut through by the Jack
Hills Gorge. Remapping of the area by the Geo-
logical Survey of Queensland raised the unit to
formation-status (Withnall, 1989) and broadened
its concept to incorporate siliciclastic rocks, in-
terpreted as lateral equivalents of the carbonates
in areas to the north and south (Withnall et al.,
1993). Recent work (Sloan et al., 1995) has
shown that same of these lateral equivalents may
represent allochthonous deposits post-dating
some autochthonous outcrop tracts of the Jack
Formation.

The Jack Formation conformably overlies and
interfingers with the Poley Cow Formation
(Fleming. 1986; Withnall & Fleming in Withnall
et al., 1993). It is disconformably overlain by
siliciclastics of the Early Devonian Shield Creck
Formation.

Specimens described here are from sections
along Bullock Creck through the western limb of
an unnamed, southwesterly plunging syncline,
located 0.5 km to the south west where the Pan-
danus to Wandovale road crosses the Broken
River, east of the axis of the Broken River Anti-
cline (Withnall & Lang, 1992). The core of this
syncline consists of two distinct lithologies of the
Jack Formation. This region was mapped as un-
differentiated Jack Formation by Withnall &
Lang (1992), Brief notes on the lithology are

364 MEMOIRS OF THE QUEENSLAND MUSEUM

ALa

AN

— Unconformity
k Section

Upper Limastona Unit
JACK FORMATION . $ро! Sample
Lower Limestone Unit

G Graptolite Locality

Judsa Bed
Graywacka/Siltstone VOLEY, REA, [13] weg 0097

interbedded FORMATION Trojamitas

Austrolion Mapping
7843000 Grid Coordinates

FIG. 1, Map of syncline near Broken River crossing showing location of sections through the Jack Formation,
north Queensland. Acanthodian remains are from Section BRO7 (Upper Bullock Creek Section), in near basal
samples BRO7-01 and BR07-02.

SILURIAN ACANTHODIAN

given below, as the sequence in the syncline
differs from the type section 4km to the west, in
the Jack Hills Gorge (Withnall & Fleming in
Withnall et al., 1993),

Tn the syncline, the lower part of the Formation
consists of thin-bedded siltstone and fine-
grained, micaceous arenite interbedded with a
range of carbonate lithologies, many with abun-
dant corals in growth positions; a highly variable
sequence colloquially known as the "coral gar-
dens" (lower limestone unit, Fig. 1). The basal
sequence of the "coral gardens" unii has, in
places, discontinuous, lenticular beds of calcinid-
ite, which crop-out sporadically around the nose
of the syncline. One of these has been interpreted
as a massive debris flow, possibly part of a chan-
nel cutting through autochthonous carbonates
and clastics of the сога] gardens unit (Sloan et al,,
1995: 52). As well as conodonts of the plaecken-
sis and siluricus Zones, and acanthodian material,
carbonates of the coral gardens unit have also
yielded foraminifera, algae, sponge spicules, by-
ronids and phyllocarid remains (Simpson, 1994).

The "coral gardens" unit at the Broken River
crossing is overlain by а massive- to thinly-bed-

upper limestone unit (Fig. 1). Thicker to
massive beds consist of fine-grained muddy and
dolomitic limestone containing minor corals and
stromatoporoids. Thinner beds consist of a vari-
ety of lithologies, including beds of richly
bioclastic detritus. These may consist of a range
of skeletal allochems or may be dominated by a
single allochem. Dominant allochems include
large articulated crinoid stems, some with
oncolitic envelopes, large, low-spired gastro-
pods, and abundant brachiopods. The upper lime-
stone unit has yielded the conodont Jcriodus
woschmidti hesperius Klapper & Murphy some
45m above the base, indicating the approximate
position of the Silurian-Deyonian boundary. A
broad study of the Jack Formation (Simpson,
1995) shows that it extends into the Devonian
only in this area, In all other regions including the
type section the uppermost beds of the formation
all terminate within the Silurian.

A review of biostratigraphic data pertaining to
all the constituent units of the Graveyard Creck
Group has been recently presented (Jell, et al., in
Withnall et al., 1993),

Samples yielding acanthodian remains were all
restricted to the "coral gardens" unit of the Jack
Formation from the Upper Bullock Creek section
(Simpson, in press). Acanthodian elements were
most abundant in samples BROT-01 and BROT-
(2, some 7m and 10m stratigraphically above Ihe

385

base of the Jack Formation, but were also recov-

ered much higher in the section, just below the

first appearance of Polygnathoides siluricus. The

age of these samples is discussed separately
ow.

AGE OF THE SAMPLES

Basal samples of the Upper Bullock Creck
section through the "coral gardens" unit of the
Jack Formation yield the Ludlow zonal index
conodont Ancoradella ploeckensis (Walliser,
1964). The species ranges through the lower 95m
of section equalling approximately the lower
75% thickness of the "coral gardens" unit (Simp-
son, in press). The succeeding Ludlow zonal
index, P. siluricus occurs in small numbers over
а narrow stratigraphic range in the upper part of
the range of A. ploeckensis. The stratigraphic
overlap of these two species is well established
and has been discussed by а number of authors
(Klapper & Murphy, 1975; Simpson & Talent,
1995) and has been recorded previously in Aus-
tralia (Link & Druce, 1972).

Although recent work (Simpson & Talent,
1995) has shown that A. ploeckensis probably
extends back into the late Wenlock, there is no
evidence 1o suggest that basal samples of the Jack
Formation (in section BRO7) are any older than
Ludlow. The associated conodont fauna is typi-
cally Ludlow in aspect. Furthermore, morpho-
metric analysis of Pa elements of A. ploeckensis
(Simpson, 1994a; Simpson & Talent, 1995) sug-
gest Ludlow rather than late Wenlock examples.

As noted above, acanthodian specimens docu-
mented here were all recovered from strata pre-
dating the appearance of P. siluricus, hence
permitting a fairly narrow age diagnosis for the
vertebrate fauna.

The siluricus Zone is generally equated with
the basal Ludfordian leinrwardinensis graptolite
Zone (Cocks & Nowlan, 1993), Jeppsson (1983)
discussed the age of P. siluricus and concluded il
was late? Leintwardinian or late? carly Ludford-
ian. Other authors give a slightly older range-basc
for the taxon: Kleffner (1989; fig. 6) indicated а
latest Gorstian age for the first appearance of the
species.

Asthe Jack Formation Acanthodian specimens
were 211 recovered from strata of the ploeckensis
Zone pre-dating the siluricus Zone, from a sec-
tion where there are no obvious stratigraphic or
chronological discontinuities, the age range of the
fauna can be considered, on available data, to
extend from the early to middle Ludlow (Gorst-

386 MEMOIRS OF THE QUEENSLAND MUSEUM

SILURIAN ACANTHODIAN

ian), with youngest examples being probably по
younger than latest Gorstian in age.

Although Australian data indicate a Gorstian
age, a single element was recovered from the
Lorentz River, Irian Jaya (Turner etal., 1995; fig,
53). This locality produced a late Ludlow crispa
Zone conodont fauna (Van den Boogaard, 1990),
indicating species range throughout the Ludlow.

SYSTEMATIC PALAEONTOLOGY

Class ACANTHODII Berg, 1922
Order ISCHNACANTHIDA Berg, 1940
Family ISCHNACANTHIDAE Woodward,
1891

Gomphonchus Gross 1971

TYPE SPECIES
Gomphonchus sandelensis (Pander) 1856.

DIAGNOSIS (after Denison, 1979: 39)

Scales with a low or convex base, low or ele-
vated crown that may be smooth or ornamented
with radiating ribs. Base is cellular bone, rarely
penetrated by fine canals from the inner surface.
The crown is dentinous with thin layers of en-
ameloid on top, without a well developed canal
system, but with long dentine tubules that rise in
the neck and turn toward the centre of the crown
top, giving off side branches. Stellate platelets
from the head have apposed rather than super-
posed growth zones. Tooth spirals carry teeth
with a large cusp and small side cusps. Fin spines
are slender, nearly straight, and usually or-
namented with smooth, longitudinal ribs of which
the anterior rib is largest. Spines are composed
entirely of dentine except for a thin basal layer of
bone.

Gomphonchus? turnerae sp. nov.
(Figs 2-6)

1982 "Vertebrate fragments’ Turner & Pickett: 314.
199] “Scales, spines, jaws and, tooth whorls of
Acanthodians’ Turner: 446-7.

387

1993 Scales cf Nostolepis and Gomphonchus Turner:
179.
71995 ‘Acanthodii indet. Tumer et al.: fig. 5].

DIAGNOSIS

An ischnacanthid acanthodian with a range of
scale types comprising: small, ‘normal’ body
scales with two to six sharp, longitudinal crown
ribs and a median longitudinal furrow; typical
ischnacanthid head scales with a broad, flat, ir-
regularly shaped base, and apposed growth zones
in the crown, and also special head scales with a
long crown bearing a jagged, irregular ornament;
pore scales with four to seven short, anterior
crown ribs, of about four pore canal openings
under the posterior crown, and up to 10 pores nn
the posterior crown surface; symphysial tooth
whorl with four or five teeth comprising from one
to five cusps, which bear longitudinal, branching,
noded ridges.

MATERIAL EXAMINED

HOLOTYPE: body scale morphotype I, UQY 7692 (Fig.
2C): lower part of the Jack Formation, Graveyard
Creek Group, Ludlow, ploeckensis Zone, exposed in
Bullock Creek, on the western flank of the unnamed
syncline Ikm southwest of the Broken River crossing
on the Wandovale-Pandanus Creek road,

PARATYPES: body scale morphotype П, UQY7653
(Fig. 2D): body scale, morphotype III, UQY 7694 (Fig.
2E,F); head scale, morphotype IV, UQY7695 (Fig,
4A,B); head/sensory line scale, morphotype V,
UQY7696 (Fig. 4C,D); symphysial tooth whorl,
UQY7697 (Fig. 6A).

OTHER MATERIAL; 303 scales, two tesserae, seven
dentition cones (including UQY7698, Fig. 6E), one
small dentigerous jaw bone fragment UQY 7699 (Fig.
6D), two fin spine fragments (including UQY7715,
Fig. 7A), and ground thin sections UQY7701-5 (Figs
3A-D, 7B.C, in order), from samples BR07-01 and
BRO7-02, Upper Bullock Creek section, Jack Forma-
lion,

ETYMOLOGY

In recognition of Dr Susan Turner's contribu-
tion to IGCP 328: Palacozoic Microvertebrates.

FIG. 2. Normal body scale types of G.? turnerae sp. nov. from sample BR07-02, Jack Formation, north
Qucensland. А,В, Morphotype ! scale, UQY7707 in anterior view (А), and latero-basal view (B). C, Crown
view of morphotype I scale, holotype UQY7692. D, Paratype scale, morphotype П, UQY 7693, latero-crown
view, E,F, Paratype scale, morphotype III, UQY 7694, in lateral view (E), and crown view (Е). С.Н, morphotype
Ш scale, UQY 7708, in crown view (С), and close-up of posterior section of scale (Н). 1.7, Asymmetrical scale,
morphotype ПІ, UQY 7709, in crown view (I), and anterior view (J). К, morphotype TII scale. UQY 7710, lateral
view. Arrow points in a rostral direction; scale bar = 0.1 mm.

388

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. Ground thin sections of scales of G.? turnerae sp. nov. from the sample BR07-02, Jack Formation. A,B,
Horizontal section of crown, morphotype I scale, UQY 7701; (B)is a magnified area of (A). C, Verticaltransverse
section of morphotype I scale, UQY7702. D, Vertical longitudinal section of morphotype III scale, UOY 7703.
Arrow points in rostral direction; scale bar = 0.1 mm unless stated.

DISTRIBUTION

Jack Formation, Broken River Province, north-
ern Queensland (Ludlow, ploeckensis Zone); un-
differentiated Palaeozoic rocks, Lorentz River,
east Irian Jaya (late Ludlow, crispa Zone).

DESCRIPTION

Body scales, morphotype I (Figs 2A-C, 3A-C)
range from 0.1 to 0.4 mm wide, and 0.1 to 0.4mm
long; most scales are towards the lower size limit.
They are bilaterally symmetrical, with a sub-
rhombic, horizontal crown, which has a sharply
marked anterior edge (Fig. 2C). Two high,
strongly developed, sub-parallel ridges lead back

from the anterior edge, decreasing in height to-
wards the posterior crown edge (Fig. 2A,C).
These ridges enclose a rounded, central furrow,
and are flanked by relatively flat, lateral areas.
Crown width and length are usually slightly less
than those of the base, though the posterior crown
point extends slightly beyond the base on some
scales (Fig. 2C). The neck slopes away from the
crown on all sides, and may have a buttressed
appearance from the vertical slits, which indicate
the position of canal openings on the latero-pos-
terior neck area (Fig. 2A,B). The neck/base junc-
tion is a well-marked rim, with a lemon-shaped
outline. The base has a typically ‘gomphonchid’

SILURIAN ACANTHODIAN

389

FIG. 4. Head scales of G.? turnerae sp. nov. from sample BRO7-02, Jack Formation. A,B, paratype head scale,
morphotype IV, UQY7695. A, in crown view. B, anterior view. C,D, paratype head/sensory line scale,
morphotype V, UQY7696. C, crown view, D, latero-crown view. Arrow points in a rostral direction; scale bar

= 0.1 mm; o = vascular canal opening.

shape, being moderately vaulted, with the swell-
ing pushed forwards, and flattening out, or be-
coming slightly concave, towards the posterior
rim (Fig. 2B).

Fungal hyphae and remineralization have ob-
scured much of the histological detail of the
scales; however, some details of the pattern of
dentine tubules, and the structure of the base, are
discernible (Fig. 3A-C). No bone cell lacunae are
preserved in the base or the crown; nor are there
any wide vascular canals. Small patches of den-
tine that are preserved (Fig. 3A,B) comprise net-
works of tubules without lacunae. The Sharpey's
fibres of the base form a simple cone, radiating
from the base apex (Fig. 3C); the bone lamellae
of the base are highlighted by the interruption of
the remineralized fibres.

Body scales, morphotype II (Fig. 2D) have a
similar size range to morphotype I scales, and
also have the distinchve central crown furrow,
hut differ in being more vertically compressed,
and in the ornamentation of the lateral crown
zones. The latter areas have several somewhat
sinuous ridges leading up and back from ihe
anterior or lateral crown edge, directed towards
the central ridges (Fig, 2D). The neck is short, and
the base is only slightly vaulted.

Body scales, morphotype III (Figs 2E-K, 3D)
are slightly larger than those of morphotypes 1
and П, being 0.4 to 0.6mm wide. Their crowns

usually have the same central furrow, however
the ridges lining it are usually shorter and lower
than on morphotype 1 and II scales. The anterior
crown bears two to four additional shortribs (Fig.
2E,G,K). Many of the scales of this type exhibit
‘cracks’ delineating the growth lines of the pos-
terior crown, but no discrete pore canal openings
ate Visible (Fig. 2H). The posterior crown edge is
denticulated on some scales, and overhangs the
base (Fig, 2F). The posterior neck area is deeper
than on morphotype I scales, and is marked by the
same vertical slits (Fig. 2K). The anterior neck
area has small, regularly spaced, circular pore
openings (Fig. 2K). As on morphotype I scales,
the base swelling is pushed forwards (Fig. 2E,K).

Head scales, morphotype IV (Figs 4A,B,
5C,D) are 0.3 to 0.4mm wide, and up to 0.6mm
long. Some less abraded scales of this type have
a micro-ornament of noded ridges (Fig. 5C.D).
The crown is approximately horizontal, and its
apposed growth yields an upper surface compris-
ing a jagged, irregular ornament, which merges
into the neck anteriorly (Fig. АВ). The crown 18
up to three times as long as the base. Small,
regularly spaced, circular canal openings pierce
the short, anterior neck (Fig. 4B). The baseis very
low or flat, and is wider than long.

Head/sensory line scales, morphotype V (Fig.
4C.D) c. 0.6mm wide and 0.5mm long. Along the
relatively straight anterior edge, the crown rises

390 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Special scales of G.? turnerae sp. nov. from BR07-02, Jack Formation. A, crown view, morphotype VI
scale, with pore canal openings in posterior crown, UQY7711. B, latero-basal view, morphotype VI scale,
UQY7712. C,D, small head scale, morphotype IV, UQY7713. C, latero-crown view. D, close-up view, showing
nodose micro-ornament. E, anterior view of short, wide scale, UQY7714. F, crown view, tessera, UOY7700.
Scale bar = 0.1mm.

SILURIAN ACANTHODIAN

up sharply from the base, then curves backwards,
so that the rest of the crown slopes up only
slightly towards the postenor (Fig. 4D). The
crown exhibits areal, rather than superpositional,
growth zones, marked by ridges running parallel
to the semi-circular outer crown edge (Fig,
4C,D). No pore canal openings are detectable on
the main crown surface. The crown is much
longer than the base, which is two to three times
wider than long, and flat or with a shallow vault,

Pore scales, morphotype VI (Fig. 5A,B), A
small number of scales (c. 2%) have several large
pore canal openings under the posterior crown,
and up to 10 (or possibly more) pores on the
posterior crown surface (Fig. 5A). The scales are
0,4 10 0, 5mm long, and 0,2 to 0.3mm wide, Short,
rounded, sub-paralle] ridges adorn the anterior
crown, rarely extending to the posterior half of
the crown surface. The posterior crown over-
hangs the base (Fig. 3B). Depth of the neck can
vary markedly, being negligible below the ante-
rior crown edge on some scales (e.g., Fig. 5A),
and quite deep on others. The base/neck junction
does not form a sharp rim; the bases of most of
these scales lack the typical *gomphonchid"
shape of the other morphotypes. Instead, the base
is relatively symmetrical, with a rounded, lumpy
surface (Fig. 5B).

Several small tesserae (Fig. 5F) of
' уер type sensu Gross, 1971 (pl, 2, fig,
27) were in the assemblage, and are also pre-
sumed to belong to G.? turnerae sp. nov.

The two symphysial tooth whorls (Fig. 6A-C)
found in the sample have a base that arches from
antenor to posterior, and are about 0.6mm long,
The whorls bear four (UQY 7697) and five teeth
(UQYT706) respectively (Fig. 6A,B), increasing
in size from anterior to posterior. A small, single-
cusped tooth is foremost on whor] UQY 7706, and
the following teeth are all three-cusped, with the
central cusps larger than the side cusps. The cen-
tral cusp (with a broken tip) of the posterior,
largest tooth of whorl UQY7706 (Fig. 6B.C) is
0.5mm high, with side cusps 0.15mm high. All
cusps bear longitudinal ridges: the noded, branch-
ing ridges of UQY7697 are particularly well pre-
served (Fig. 6A).

Seven dentition cones (Fig. 6E) have the same
form as some of those of Peracanthodes menneri
desenbed and figured by Valiukevicius (1992; pl,
2, figs 2,3, pl. 3, fig. 2): namely, an elongated,
hollow cone, with a row of denticulations and/or
randomly positioned denticles. Burrow (in press,
h) characterised these elements, from Lochkov-

39]

jan microvertebrate assemblages of central New
South Wales, as dentition cones type I and П.

One small dentigerous fragment (UQY7699
Fig. 6D) of the anterior section of a jaw bone was
preserved. The base has the lalero-medial con-
cavity characteristic of acanthodian jaw bones,
marking the position of the jaw cartilage (Fig.
6D). The occlusal surface has a medial ridge, and
a higher lateral ridge, separated by a shallow
groove. Only one cuspis preserved, on the broken
posterior extremity of the fragment.

Several small fin spine fragments (Fig. 7A-C)
up to 2mm long were in the assemblage; some of
these were sectioned (Fig, 7B,C). The spines are
ornamented with relatively smooth and rounded,
parallel, longitudinal ribs, The anterior rib, form-
ing the leading edge of the spine, is widest, fol-
lowed by three ribs each about 0.] mm wide. All
ribs are separated by rounded grooves of equal
width. A wider groove separates these nbs from
the posterior area of the spine, which varies in
width depending on the age of the spine,

REMARKS

The material described comprises the sum of
the determinable acanthodian elements in the
sample, and by comparison with the range of
elements in articulated specimens of
Poracanthedes menneri Valiukevicius, 1992, is
interpreted as belonging to a single species. Gross
(1971), in attributing scales, tooth whorls, dentig-
erous jaw bones, and fin spines to Gomphonchus
sandelensis, and Gomphonchus sp., considered
that Poracanihodes (Brotzen, 1934) was not an
independent genus, and believed poracanthodid
scales to he modificd lateral line scales of G.
sundelensis and С. hoppei. Whereas
Poracanthodes i$ now recognized as a valid
genus, no articulated specimens of Gomphonchus
have yet been found. As Grass did not attribute
any dental elements or fin spines to
Poracanthodes, il is probable that some of the
elements he assigned to Gomphonchus are refer-
able to Poracanthodes (see Burrow, in press, b,
for further discussion of this topic).

Unfortunately, remineralization and fungal hy-
phac have obscured histological detail of the Jack
Formation scales, precluding a firm generic clas-
sification based on the scales alone. Morpholog-
ically, the posterior crown section of morphotype
TII scales appear to show areal rather than super-
positional growth (Fig. 2F-H), as in scales of
Poracanthodes menneri. Pore canal networks are.
not detected, however, in ground thin sections of
G.? lurnerae sp. nov, scales. The growth zone

392 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 6. Symphysial tooth whorls, dentigerous jaw bone fragment, and dentition cone of G.? turnerae sp. nov.,
from the Jack Formation. A-D are from sample BR07-02, E is from BRO7-01. A, occlusal view of paratype tooth
whorl, UQY7697, with branching, noded ridges. B,C, tooth whorl UQY7706. B, in lateral view. C, occlusal
view. D, concave base of dentigerous jaw bone fragment UQY 7699. E, lateral view, dentition cone UQY7698,
Arrow points in a rostral direction. Scale bar = 0.2mm.

SILURIAN ACANTHODIAN

393

FIG. 7. Fin spine fragments of G.? turnerae sp. nov., from BRO7-02, Jack Formation. A, lateral view, UQY7715.
B, cross-section of a juvenile fin spine, with open cavity, UQY7704. C, Cross-section of fin spine with two
growth zones, UQY7705. fc = longitudinal fin spine cavity, gl = growth line delimiting first growth zone. Scale

bar = 0.2 mm.

‘cracks’ on these scales are probably an artifact
of preservation, caused by the thinness of the
growth zones in the posterior crown, abrasion,
and fungal invasion. This theory is enhanced by
the lack of pore canal openings on the crown of
morphotype V head scales, as comparable scales
on P. menneri have many such openings
(Valiukevicius, 1992; pl. 9, fig. 6a,b). Morpho-
type V head/sensory line scales are commonly
found in microvertebrate assemblages (e.g., Bur-
TOW, in press, a; pl. 3, fig. 14; and possibly Mader,
1985; pl. 3, fig. 2), and are similar to the sensory
line scales on the head of Jschnacanthus gracilis,

in Miles (1966; fig. 11) and Bernacsek & Dineley
(1977; pl. 6, fig. 4). Morphotype IV head scales
appear the same as the special head scales of P.
menneri (Valiukevicius, 1992; pl. 9, figs 2-4).
Only the poorly represented morphotype VI
scales assigned to G.? turnerae have the pore
canal openings on the underside of the posterior
crown, which are typical of Poracanthodes
scales.

The symphysial tooth whorls (Fig. 6A-C) re-
semble those attributed to "Gomphodus"
sandelensis by Gross (1957; fig. 1), and differ
from the lone, small, tri-cusped whorl assigned to

394

Р. menneri (Valiukevicius, 1992; figs. Sa,b).
However, the latter whorl was broken, and the
number of teeth per whorl is probably variable,
as illustrated by Gross (1957; figs 1,2) for
"Gomphodus" sandelensis. The G.? [urnerae
whorls differ to all previously described exam-
ples by their distinctive, branching, longitudinal
ridges (Fig. 6A); this ornament gives the cusps а
‘tubercular’ appearance.

The G.? turnerae fin spine fragments (e. g., Fig.
7A) resemble those of Р. menneri (e.g.
Valiukevicius, 1992; text fig. | 1B). Fin spines of
the ischnacanthids Jschnacanthus gracilis, P.
menneri, and Gomphonchus sp, conform to a
common pattem of smooth, longitudinal ribs,
with the leading rib wider than the following ribs.
The number of the narrower ribs varies with the
age of fish, and the position of the spine on the
body (Valiukevictus, 1992; 205), Gross (1971;
figs 24E,F, 25А,С,р,Е, 26А) illustrated cross-
sections of young and old spines, showing the
increasing number of growth zones correspond-
ing to the age of the spine, which leads to the
variation in their external morphology. Fin spine
omamentation, thus, is as yet of limited use in
assigning fin spines to any of these ischnacanthid
genera,

A comparable suite of acanthodian elements,
incorporating all elements listed for G.? turnerae
except the morphotype IV scales, was observed
in microvertebrate assemblages from several
sites in the Lower Devonian Trundle beds of
central New South Wales (Burrow, in press, a, b),
This particular range of elements was tentatively
attributed tò а new taxon. Only a small percentage
of the Trundle beds scales of this taxon had pore
canal openings on the upper surface of the crown,
just as in the G.? turnerae sample. Determining
if the new ischnacanthid should be assigned to
Poracanthodes, от Gomphanchus, or a new
genus, is difficult without scales which have been
well preserved histologically. Pamarily on the
basis of the lack of a discemible pore canal sys-
tem, and the apparent absence of bone cell lacu-
nae in the base, they are assigned to
Gomphonchas? rather than Poracanthodes be-
cause of the lack of good histological detail, they
are not assigned to a new genus. A scale figured
by Tumer et al. (1995; fig, 53) from an erratic
block af the Lorentz River, east Irian Jaya (late
Ludlow age), appears to bea morphotype Ш scale
of G. turnerae. Gomphonchus? turnerae is the
oldest acanthodian described from Australian as-
semblages.

MEMOIRS OF THE QUEENSLAND MUSEUM

ACKNOWLEDGEMENTS

We thank Dr Susan Turner (Queensland Mu-
seum) for her initial investigations of the mate-
rial, and for critical appraisal of the manuscript;
and Dr John Long (Western Australian Museum)
for SEM photography and ground thin sections of
some of the material, Specimens were recovered
during an honours project undertaken by one of
us (AJS) at Macquarie University. Ruth Mawson
and John Talent (Macquarie University) are
thanked for the use of facilities, encouragement
and discussion on stratigraphy. John Jell (Univer-
sity of Queensland) is thanked for his field exper-
tise and imparting his knowledge of the
stratigraphy of the region. Fig, 1 was drafted by
Joanne Simpson with customary skill. This is a
contribution tò IGCP328: Palaeozoic
Microvertebrates.

LITERATURE CITED

BERNACSEK, G.M. & DINELEY, D.L. 1977, New
acanthodians from the Delorme Formation
(Lower Devonian) of N.W.T., Canada. Pal-
acantographica A 158: 1-25.

BURROW, C.J, (in press, a). Microvertebrate assem-
blages from the Lower Devoniàn (pesavis/sul-
catus) of central New South Wales, Australia.
Modern Geology.

(їл press, b), Acanthodian dental elements from the
Trundle beds (Lower Devonian) of New South
Wales, Records of the Western Australian Mu-
scum

COCKS, L.R.M, & NOWLAN, G.S, 1993, New left
hand side for correlation diagrams, Silurian Times
1: 6-8

DENISON, R.H. 1979, Acanthodii, Pt, 5. In Schulize,
H-P. (Ed.), "Handbook of Paleoichthyology".
62p, (Gustav Fischer Verlag: Stuttgart, New
York).

FLEMING, P.J.G. 1986. Sections through the Silurian
to Devonian Jack Limestone Member, Broken
River area, north Queensland. Queensland Gov-
emment Mining Journal 87: 141-144,

GROSS, W. 1957. Mundzühne und Hautzähne der
Acanthodier und Arthrodiren. Pal
acontolographica A 109: 1-40,

1971. Downtonische und ditlonische Acanthodier-
Reste des Ostseegebieles. Palacontographica А
136: 1-82.

JELL, J.S., SIMPSON, A.J., MAWSON, R. & TAL-
ENT, J.A. 1993, Biostratigraphic summary. In
Withnall, 1.W. & Lang, S.C. (Eds), ‘Geology of
the Broken River Province, north Queensland",
Queensland Geology 4: 239-245.

JEPPSSON, L. 1983, Silurian conodont faunas from
Gowland. Fossils and Strata 15: 121-144.

SILURIAN ACANTHODIAN

KLAPPER, G. & MURPHY, M.A, 1975. Silurian -
Lower Devonian conodont sequence in the Rob-
erts Mountains formation of central Nevada. Uni-
versity of California Publication, Geological
Sciences 111: 1-62.

KLEFFNER, M.A. 1989. A conodont-based Silurian
chronostraugraphy. Bulletin, Geological Society
of America 101: 904-912.

LINK, A.G. & DRUCE, E.C. 1972. Ludiovian and
Gedinnian conodont stratigraphy of the Yass
Basin, New South Wales, Bureau of Mineral Re-
sources, Australia, Bulletin 134: 1-136.

MADER, Н. 1986. Schuppen und Zähne von
Acanthodiern und Elasmobranchiern ans dem
Unter-Devon Spaniens (Pisces), Gétinger
Arbeiten Geologie und Paläontologie 28: 1-59.

MILES. R.S. 1966. The acanthodian fishes of the
Devonian Plattenkalk of the Paffrath Trough in the
Rhineland. Arkiv för Zoologiel 18(9); 147-194,

SHERWIN, L. 1994, Palaeozoic Stratigraphy of the
Narromine 1: 250000 Sheet area. Geological Sur-
vey of New South Wales, Quarterly Notes 96:1-
37

SIMPSON, A. 1983. Silurian to basal Devonian conod-
ont biostratigraphy of the Broken River area, north
Queensland. BA(Hons,) Thesis, School of Earth
Sciences, Macquarie University, (unpublished).

1994a. Variations in the Ludlow conodont genus
Ancoradella, and their potentia] chronological
significance. Australasian Palacontological Con-
vention, 1994, Abstracts and Programme, Mac-
quarie University Centre for Ecostratigraphy and
Palaeobiology: 54.

1994b. Silurian to basal Devonian conodonts and
other microfossils from the Graveyard Creek
Group, Broken River crossing, north Qucens-
land. Australasian Palaeontological Convention,
1994, Abstracts and Programme, Macquarie Uni-
versity Centre for Ecostratigraphy and Palaeobio-
logy: 88.

1995. Silurian conodont studies in eastern Australia.
PhD thesis, Department of Earth Sciences, Uni-
versity of Queensland (unpublished).

IN PRESS, Silurian 10 basal Devonian conodonts
from the Broken River Crossing, north Queens-
land. Journal of Paleantolog

SIMPSON, A.J. & TALENT, J. А. 1995. Silurian con-
odonts from the headwaters of the Indi (upper
Murray) and Buchan rivers, southeasiem Aus-
tralia, and their implications, Courier
Forschungsinstitut Senckenberg 182: 79-215.

SLOANE, T.R,, TALENT, ЈА. MAWSON, R SIMP-
SON, AJ., BROCK, G.A.. ENGELBRETSEN,
M. JELL, J.S., AUNG, A.K., PFAFFENRIT:
TER, C, TROTTER, J. & WITHNALL, LW.
1995, Conodont data from Silurian - Middle
Devonian carbonate fans, debris flows, al-
lochihonous blocks ‘and adjacent autochthonous

395

platform margins: Broken River and Camel Creek
areas, north Queensland, Australia, Courier
Forschungsinstitut Senckenberg 182: 1-77.

TURNER, S. 1991. Palacozoic vertebrate microfossils
in Australia. Pp.429-464. In Vickers-Rich. P.,
Monaghan, J.N.. Вагі, R.F. & Rich, T.H. (eds),
"Vertebrate palaeontolo op in Australasia".
(Poineer Design Studios with Monash University,
Publications Committee; Melbourne),

TURNER, S, 1993, Palaeozoic microvertebrate
biostratigraphy of Eastern Gondwana. Pp. 174-
207. In Long, J.A. (Ed.), "Palaeozoic vertebrate
hiostraligraphy and biogeography". (Bellhaven
Press: London). о!

TURNER S. & PICKETT, J. 1982. Silurian vertebrates
in Australia, Scarch 13; 314-315.

TURNER, S.. VERGOOSSEN, J.MJ. & YOUNG,
G,C. 1995, Fish microfossils from Irian Jaya.
Memoirs of the Association of Australasian Рај.
acontologists 18; 165-178.

VALIUKEVICIUS, J.J. 1992, First articulated
Poracanthodes from the Lower Devonian of
Severnya Zemlya, p. 193-214. In Mark-Kurik, Е.
(Ed.), "Fossil Fishes as Living Animals". (Acad-
emy of Sciences of Estonia: Tallinn).

VAN DEN BOOGAARD, M. 1990. A Ludlow conod-
om fauna from Inan Jaya (Indonesia), Scripta
Geologica 92; 1-27.

WALLISER, O.H. 1964, Conodonten des Silurs.
Abhandlungen des hessischen Landesanstes fur
Bandeforschung, Wiesbaden 41: 106pp.

WHITE, D.A. 1959. New names in Queensland strag-
raphy, Parts 2, 3 and 4, Oil and Gas Journal 5(9):
31-36; 5(10); 31-36; 5(1 1): 26-28.

WITHNALL, LW. 1989. Revision of the straligraphy
of the Broken River area, north Queensland -
Ordovician and Silurian units. Queensland Gove
emment Mining Journal 90: 213-218.

WITHNALL, LW., FIELDING, C.R., LANG, S.C. &
FLEMING, PJ.G. 1993. Stratigraphy and sedi-
mentology of the Silurian to Early Devonian
Graveyard Creek Group and Shield Creek Forma-
tion, In Withnall, IW. & Lang. S.C, (Eds), 'Geol-
ogy af the Broken River Province, north
Queensland’, Queensland Geology 4: 55-78,

WITHNALL, LW. & LANG, S.C. 1992. Broken River
Special, 1:100,000 scale map. (Department of
Resource Industries; Queensland).

WITHNALL, I.W.. LANG, S.C.. JELL, J.S.,
MCLENNAN, Т.РТ., TALENT, J-A„ MAW-
SON, К. FLEMING, P.J.G.. LAW, SR;
MACANSH, J.D., SAVORY, P, KAY, J.R. &
DRAPER, J.J, 1988, Stratigraphy, sedimentolngy
bipstratigraphy and tectonics of the Ordovician to
Carboniferous, Broken River Province, north
Queensland. Australasian Sedimentologists
Group Field Guide Series 5: 1-200.

396

SAP FEEDING BY THE AUSTRALIAN GECKO
GEHYRA DUBIA, Memoirs of the Queensland Museum
38(2):396. 1995;- Although Australian geckos are generally
opportunistic arthropod feeders (Greer, 1989), several species
will also feed on plant nectar and sap. Christinus guentheri
and Rhacedactylus lindneri lick nectar from blossoms (Cog-
ger etal., 1983; King & Horner, 1993), белуга australis licks
decaying, or pulpy fruit (King & Horner, 1993); and
Christinus marmoratus, Heteronotia binoei and
Lepidodactylus lugubris will feed in captivity on sugar- based
substances (after Greer, 1989). Only two records of geckos
feeding on sap are known. Gehyra variegata feeds on the sap
of small Acacia shrubs in Western Australia (Dell, 1985);
Strophrurus spinigerus was seen by one of us (SKW) feeding
on Acacia sap near Perth, WA, in spring, 1981.

On 2] March, 1995, in open forest 15km south of Yuleba
(26"43'26"S, 149°19'Е) SEQ, a specimen of белуга dubia
was observed al approximately 1930h head-down, apparently
licking sap from the trunk of a small tree, Acacia leiocalyx
leiocalyx. Small black ants were present, so it was difficult to
determine whether the gecko was feeding on these or the
Acacia sap.

Later in the evening 2040h ikm south of the initial obser-
valion site, several specimens of G. dubia (2-3 per tree) were
seen near ihe bases of trunks of small Acacia trees. Six
specimens were feeding on sap. Two were observed for 18
minutes, The first was feeding 15cm above the ground, on a
2,7 metre high Acacia l. leiocalyx tree with a trunk diameter
of 5cm. The gecko was head-down, licking ata lcm ‘bead’ of
sap on which a solid crust had formed. The gecko's tongue
penetrated this outer crust to extract the softer, almost liquid,
sap. The second gecko was: feeding in the same way, also on
sap of an А. I leiocalyx tree (3m high, trunk diameter 8cm).
The gecko was about 1 lcm from the ground, and was licking
a 2cm weeping ‘wound’ on the Acacia trunk. Both geckos
were collected (QMJ59560-61), killed and preserved im-
mediately. The full duration of this behaviour was not re-
corded. At this site, geckos were also feeding on sap from trees
of two other species - A. confería and A. decora. Similar
behaviour by С. dubia was observed in Barakula State Forest
(26°15'5, 150?30' E) in December, 1982 (SKW), on an un-
identified Acacia tree.

Both QMJ59560 -61 are adult males, with enlarged testes.
They have full stomachs, That of QMJ59560 (SVL 63,0mm)
contains fragments of a large katydid (Tettigoniidae). The

MEMOIRS OF THE QUEENSLAND MUSEUM

stomach of QMJ59561 (SVL 64.7mm) contains earwig and
cockroach fragments, No recognisable Acacia sap is present.
There are two possible explanations for the apparent absence
of sap from both stomach contents: relatively small amounts
of it were ingested; it is likely that sugary fluids are rapidly
and completely digestible. The presence of insect remains in
both gut samples shows that arthropods remain an important
food source for G. dubia specimens, even when they feed on
sap.

Po ap-feeding by Gehyra cf. baliola and Rhacodactylus
australis has been observed (SKW) recently. à specimen of
the former Was photographed (QMNP754) feeding on the sap
of an Acacia sp. tree, 500m from the tip of Cape York
Peninsula (10*41'S, 142°33'Е). The R. australis was seen
feeding on the sap of an unidentified rainforest tree in the
Lockerbie Scrub (10°47'S, 142"28' E).

Literature Cited

Cogger, H., Sadlier, R. & Cameron, E., 1983, The terrestrial
reptiles of Australia's island territories, Australian Na-
tional Parks and Wildlife Service. Special publication

11; 80pp.

Dell, J., 1985. Arboreal geckos feeding on plant sap. Western
Australian Naturalist 16(4): 69-70.

Greer, A.E., 1989, "The biology and evolution of Australian
lizards’. (Surrey Веацу & sons Pty Ltd: Chipping Nor-
lon, NSW) 264pp.

King, M. & Horner, P., 1993. Family Gekkonidae. Pp 221-
233. In Glasby, C.J., Ross, G.J.B, & Beesley, P.L. (eds),
Fauna of Australia vol. 2a, Amphibia & Reptilia. (Aus-
tralian Government Publishing Service: Canberra).
439pp.

Acknowledgements

This discovery was made on field work funded by the
Australian Nature Conservation Agency (Endangered Species
Program). We thank Trevor Beetson (DPI) for identifying the
Acacia spp. on which the geckos were feeding, and Dr Geoff
Monteith (QM) for identifying arthropod remains in the gut
samples.
P.J. Couper, J.A. Covacevich, S.K. Wilson, Queensland Mu-
m Box 3300, South Brisbane, 4101 Australia; 28 August
1995.

CRASPEDELLINAE BAER, 1931 (PLATYHELMINTHES: TEMNOCEPHALIDA)
ECTOSYMBIONTS FROM THE BRANCHIAL CHAMBER OF AUSTRALIAN
CRAYFISH (CRUSTACEA: PARASTACIDAE)

LESTER R.G. CANNON AND KIM B. SEWELL

Cannon, L.R.G, & Sewell, К.В. 1995 12 01: Craspedellinae Baer, 1931 (Platyhelminthes;
Temnocephalida) ectosymbionis from the branchial chamber of Australian crayfish
(Crustacea: Parastacidae), Memoirs of the Queensland Museum 38(2): 397-418. ISSN
0079-8835.

Craspedellinae is recognised to include taxa of temnocephalids which live in the branchial
chamber of Australian crayfish (Parastacidae) and arc characterised by possession of one or
таге transverse papillate ridges across the dorsal body and crenulate (papillate) tentacles,
Le. 6 species of Craspedella Haswell, 1893, 5 of them new, all from eastern Australian
Cherax spp., and 3 new ies of a new genus all from Western Australian Cherax spp.,
and а new genus and species from Euastacus bispinosus from Victoria. [ ] Temnocephalida,
Craspedellinae, Craspedella, crayfish, ectosymbionts, taxonomy-

Lester R.G. Cannon & Kim B. Sewell, Queensland Museum, PO Box 3300, South Brisbane,

Queensland 4101, Australia; 20 October 1995.

Craspedella spenceri Haswell, 1893, described
from the branchial chamber of Astacopsis
bicarinatus Gray, 1845 [i.e. Cherax destructor
Clark, 1936] from easter Australia, is currently
the only described species in the genus, Haswell
(1893) characterised the species thus “In the pos-
terior half of the body, on the dorsal surface, are
three transverse lamellae, 0.05mm in breadth.
divided into about 15-20 rounded lobes, each
tipped with a few papillac....'. The only other
temnocephalid known to have dorsal papillae is
Notodacrylus handschini Baer, 1953 which oc-
curs on the carapace of Cherax spp. from Aus-
tralia and New Guinea. In N. handschini the
papillae are not raised on ridges or lamellae and
are retractile between the peculiar dorsal surface
scales unique to this species (Jennings, Cannon
& Hick, 1992). One scutariellid, Bubulecercus
sketi, from cave shrimps in Slovenia and Croatia
is reported to have papillae ‘strewn’ all over the
body (Matjasic, 1990).

The habitat of C. spenceri within the branchial
chamber is sheltered and this small 'less than
2mm', delicate and non-pigmented worm can
easily go unnoticed; this is especially so when
contrasted with the much larger ‘(about 0.5ст)'
Temnocephala dendyi Haswell, 1893 which also
occurs in the branchial chamber of Cherax de-
structor, or the large, and often pigmented, tem-
nocephalids obvious on the external exoskeleton
of many of Australia's crayfish.

The crayfish branchial chamber often supports
a rich fauna (Alderman & Polglase, 1988; Can-
non & Jennings, 1987; Jennings, 1988) including
temnocephalids: the Scutariellidae are found

there in shrimps from Europe (Matjasic, 1990)
and Asia (Baer, 1953); in Australia the monotypic
Actinodactylellidae is found in the branchial
chamber of burrowing crayfish, and several
members of the Temnocephalidae are now
known from shrimps and crabs (Cannon, 1993).
Although С, spenceri remains the only described
specics in the genus, researchers have examined
small branchial chamber dwelling tem-
nocephalids from Cherax spp. and have referred
to them as either Craspedella sp. (see Rohde,
1987a,b; Jones & Lester, 1992) or C. spenceri
(see Cannon & Jennings. 1987).

Examination of the branchial chamber of cray-
fish Cherax spp. and Evastacus spp. collected
from around Australia during 1990-1992 as part
of an Australian Biological Resource Study of the
Temnocephalida revealed new taxa which arc
described here.

MATERIALS AND METHODS

Live crayfish were captured in collapsible min-
now traps baited with fatty mutton or canned cat
food, and occasionally by dip netting, and were
maintained for up to several hours in water from
the habitat before they were dissected. To obtain
living worms where captured crayfish were plen-
tiful, the carapace was detached using strong
forceps inserted anteriorly through the articular
membrane and under the dorsal carapace, and the
carapace and carcass placed into a shallow vessel
containing water from the habitat. The inner sur-
face of each branchiostegite (1.6, the branchios-
tegal membrane), the gills and the body wall were

398

searched with the aid of a dissecting microscope.
Worms were collected on wooden points and
transferred lo a watch glass containing water. To
obtain living worms where crayfish were rare,
and needed intact for confirmation of host iden-
tification, several podobranchs were detached
from the bases of the walking legs and placed in
a watch glass containing water. Otherwise, the
crayfish and any worms were killed and fixed by
immersion in near boiling water (HW). All dis-
membered hosts, and when possible, additional
intact host specimens were stored after HW fixa-
tion in 70% ethanol (Al) for subsequent taxo-
nomic host identification and as a source of
additional worm specimens. Where possible
worms were drawn alive with the aid of a camera
lucida, although for many species observations
on living worms in the ficld was not possible.

For histology, living worms were flooded with
fixative, usually cold 10% buffered formalin
(Form), sometimes Bouin's fluid (Bouin) or ace-
tic, formol, alcohol (AFA): some were fixed with
Berland's fluid (10% formalin in glacial acetic
acid) Some living worms removed from their
host were fixed with hot water and transferred
immediately to 10% phosphate buffered formalin
(HW/Form), Where the crayfish host was col-
lected by other than the collector of the worms the
labelling convention - host collector/worm col-
lector - is observed. Wholemounts were prepared
hy staining with cither Mayer's or Harris' Hae-
matoxylin (Hx) and mounting in Canada balsam.
Serial sections were prepared from worms em-
bedded in 56°C Paraplast, cut at 6-7%m and
stained with Mayer's haematoxylin and eosin
(H&E) or (rarely) using either Mallory's
Тпсһготе (MT), Heidenhain's Azan (HA) or
Heidenhain's Iron Haematoxylin (HIH) and
mounted in Depex. All measurements were made
with the aid of a camera lucida.

For scanning electron microscopy (SEM}
warms were fixed by flooding with either (1) hot
water (approx, 90°C), then transferred to 10%
phosphate buffered formalin (HW/Form), or (2)
cold 3% glutaraldehyde (4°С). Fixed worms were
washed severul times in distilled water to remove
surface contamination, dehydrated in a graded
alcohol series, critical point dried, mounted on
stubs, coated with gold, and examined with a
Hitachi S-530 SEM operating at 20 or 25kY,

For examination of the cimus, worms were
placed on a slide in a drop of de Faure's (deF)
mounting medium (distilled water 501; chloral
hydrate 50g; glycerol 20ml and gum arabic 30g)
on a microscope slide either alive, or in the case

MEMGIRS OF THE QUEENSLAND MUSEUM

of fixed specimens after rinsing in distilled water
for 2-5 days to remove fixative, covered wilh а
coverslip and examined using bright field and
N 1 microscopy. This technique clears the
soft body tissue of the worms and provided much
clearer images of the sclerotic male copulatory
organ than those from stained wholemounts.
Fixed worms cleared less effectively than live
worms when placed in de Faure's medium. The
shape of the vaginal cavity and its pattern of
ridges and folds were also revealed by Faure's
medium.

Taxonomic descriptions of worm species were
prepared initially with the aid of the DELTA
program (Dallwitz & Paine, 1986) and based on
measurements from the type series. Terminology
follows that discussed by Cannon (1993). Mate-
nal is deposited in the collections of the Queens-
land Museum (QM) and wholemounts are
designaled (WM), de Faure’s cirrus preparations
(CP) and serial sections (LS, TS or FS) -longiti-
dinal, transverse or facial sections: the number of
slides in the series is given in brackets, Camera
lucida drawings were scanned and used as tem-
plates for illustrations which were prepared using
Adobe Jilustrator, and photomicrographs were
scanned from 35mm slide or negative film onto
Kodak Photo CD, edited and assembled into
plates using Adobe Photoshop.

MEASUREMENTS AND TERMINOLOGY

Considerable variation in the relative size,
shape and position of internal structures was ob-
served in living worms. This plasticity of shape
and further distortion caused by the effecis of
fixative indicate that measurements are valuable
only as a guide to the size and shape of the worms
and their internal structures. Thus, the measure-
ments we provide for soft structures and the cirrus
are taken only from the large mature worms
which comprise respectively the type series and
de Faure's cirrus preparation series,

The terminology we use to describe the male
reproductive structures essentially follows Can-
non (1993), However, some reiteration and re-
finement of the terms are necessary. Following
Cannon (1993), we term a cirrus the entire scle-
тойс male copulatory organ comprised of an in-
trovert (flexible distal eversible region armed
with spines) and sbaft (rigid, tapering. proximal.
tubular region). The shaft tapers proximal to dis-
tal and is further classified to have the shape of
either a funnel, a goblet or a cone. Funnel or
gobjet shaped shafts have a wide proximal region
which tapers rapidly to form a narrow, tubular

CRAYFISH ECTOSYMBIONTS

distal region. Light microscopy reveals that the
introvert is comprised of (1) an inner wall of thin
sclerotic material which appears altached to, and
effectively makes a continuation of, the shaft and
from which the spines project inward and distally
when not everted and (2) an outer layer of pre-
sumably sclerotic material optically distinct from
the inner wall i.e., the swelling or enlargement
described by Cannon (1993). The swelling varies
in thickness over the length of the introvert being
thinnest distally and continues proximally for a
short distance past the base of the introvert to the
distal region of the shalt. Our descriptions of the
cirmus refer to the inverted state of the organ.

Measurements of cirrus length were made
along the outside of the shaft wall but inside the
introvert swelling. Measurements of total cirrus
length were made along the longest side of the
organ and include the introvert. Measurements of
the width of the introvert base include the thick-
ness of the shaft walls but exclude the introvert
swelling. Measurements for the copulatory bulb
exclude the cirrus, The arrangement and orienta-
tion of the prostate duct reservoirs within the
copulatory bulb is characterised as approximately
either parallel, diagonal, or at right angles in
relation to the longitudinal axis of the copulatory
bulb.

SYSTEMATICS
CRASPEDELLINAE Baer, 1931

DIAGNOSIS

Temnocephalidae with oval or elliptical body
slightly dorso-ventrally compressed, without lat-
cral flanges and with five similar anterior tenta-
cles. Tentacles with an annulate or crenulate
appearance derived from prominent, conical, cil-
jated papillae arranged in near regular rows along
and around a central axis. Rhabdite tracts to all
tentacles, but most conspicuous in central three
tentacles, with pore openings most concentrated
in the ventro-distal region just posterior to the tip.
Pigment confined to a single dorsal pair of eyes
at the base of the tentacles. Posterior sucker pe-
dunculate, strongly muscled, with a circular ad-
hesive disc and marginal valve. Dorsal surface
without imhricating scales, but with one or more
transverse body ridges, bearing raised papillae;
with dorsal body ridges arranged radially poste-
rior to the mos! posterior transverse row also with
raised papillae. Without locomotory cilia. Buccal
cavity or pre-pharynx inconspicuous; pharynx
directed antero-ventrally, rudimentary, undi-

399

vided; pharynx sphincters equal; oesophagus in-
conspicuous; gut without colour or septa. Gut as
wide as long and usually seen extending posteri-
orly further on the right. Major nerve trunks in-
conspicuous; eyes present, spherical to ovoid,
discrete but close; pigment granules irregular,
black-brown and mainly small. Longitudinal and
circular muscles of body wall usually of equal
strength; body догѕо-уепіга] muscles weak. Át-
tachment muscles of pharynx weak. Muscles con-
trolling the male organ and about seminal vesicle
and copulatory bulb strong. Gonopore mid-ven-
tral in posterior third of body; genital atrium
commodinus; without bursa copulatrix (but
atrium may function às a bursa). Ovary spherical
to ovoid; vesicula resorbiens usually spherical to
ovoid/kidney-shaped. Vagina muscular, usually
with bulbous sphincter distally; vaginal cavity
without teeth but delineated by prominent longi-
tudinally oriented ridges and folds; seminal re-
ceptácle single; vitellaria scattered. Testes, two
pairs anterior and posterior, spherical to ovoid,
positioned lateral and posterior to gut respec-
tively, smooth to follicular. Ejaculatory sac
where present contained in copulatory bulb, Pros-
tate secretion fills copulatory bulb around ejacu-
latory duct. Disc glands scattered across dorsal
body at about the posterior of posterior testes,
Small, slender worms up to 2mm in vivo. Inhah-
iting the branchial chamber of their hosts.

REMARKS

Cannon (1986) recognised three families
within the Temhocephalida - the monotypic
Actinodactylellidae from Australia, the
Scutanellidae from and Asia, and the
large and diverse Temnocephalidae from Mada-
gascar, Asia, Australasia, and South and Central
America.

Several genera of the Temnocephalidae are
monotypic, Craspedella spenceri being unique in
having papillae raised on dorsal ridges (Haswell,
1893). Baer (1931) originally proposed a separate
family for this species, giving the following diag-
nosis:

"Temnocephalids of rather small size with five
mobile tentacles at the anterior end. In the poste-
rior region of the worm are three pairs of small
tentacles situated on the edges of the animal and
joined, two by two, on the dorsal surface, hy a
type of small dentate fringe, Behind the last
fringe, al the posterior end, are four more small
tentacles. The pharynx is rudimentary. The other
anatomical! characters are identical with those of
the Temnocephalidae."

400

MEMOIRS OF THE QUEENSLAND MUSEUM

CRAYFISH ECTOSYMBIONTS

Baer (1931) obviously misinterpreted the de-
scription of Haswell (1893) and believed the
edges of the papillate ridges (fringes) were small
lateral tentacles. In fact the papillae are not nec-
essarily larger than those elsewhere on the ridges
and are not ‘trois paires de petits tentacles situés
sur les bords de l'animal’. Regardless, Bresslau
& Reisinger (1933) considered these characters
insufficient to create a new family for this one
species and Hyman (1951) and Baer (1961) ac-
cepted this. The discovery of new taxa, all sharing
similar morphology, prompts the delineation of
this group from the remainder of the Tem-
nocephalidae.

Type genus: Craspedella Haswell, 1893

Other genera: Heptacraspedella gen. nov.,
Zygopella gen. nov.

KEY TO CRASPEDELLINAE BAER, 1931

1. With more than one transverse papillate ridge on
dorsal body and without pits, but with four papil-
late ridges which radiate towards the body mar-
gin behind the most posterior transverse ridge .

With a single transverse papillate ridge on poste-
rior dorsal body and a pair of posterior pits de-
fined dorsally by a papillate ridge

ОРЦА ЕЕ Zygopella gen, поу, 3
2. With 3 transverse papillate dorsal ridges... .
сос Shea bat £6 32А Craspedella

With 7 transverse papillale dorsal ridges . . . .

.. . Heptacraspedella peratus gen. et sp. nov.

3, Cirrus small, cone shaped and lacking large spines
4

ot o9 on 9 on on

Cirrus large, cone-shaped with large spines
Z. deimala sp. nov.

++ «+ + om к= ии

4, Cirrus with wide introvert opening and junction
between introvert and shaft oblique
bine eh eb qu mr Z. pista sp. nov.
Cirrus with narrow introvert opening; tentacles
уетугиозе......... 7. stenota sp. nov.

5. Cirrus shaft distal region not reflexed; introvert
not permanently everied, armed with spines . .

Bom а = 4 ecc om 4 o4 к» э = bu >м we. v ө

Cirrus shaft distal region reflexed; introvert per-
manently everted, spines reduced to be flat, over-
laid and 'plate-like' |... . . C. pedum sp. nov.

5. Cirrus not slender, thick-walled

401

7. Cirrus shaft not cone-shaped and without thick
proximal rim
Cirrus shaft cone-shaped with thick proximal rim

"itii С. sharti sp. nov.

8, Cirrus shaft goblet-shaped; vagina with distinct

"lateral pocket"
Cirrus shaft funnel-shaped with wide proximal

opening; vagina without distinct ‘lateral pocket"

C. spencert

9. Cirrus shaft: ratio of length of tubular distal re-
gion to length of introvert about equal and never
lessthan tI oe ereton C. yabba sp. nov,

Cirrus shaft: ratio of length of tubular distal re-
gion to length of introvert about 0.7:1 and al-
ways less than 1... . . C. simulator sp. nov.

Craspedella Haswell, 1893

DIAGNOSIS

Craspedellinae with three dorsal papillate
ridges in the posterior half of the body and, behind
the last ridge, four short posterior papillate ridges
radiating towards the posterior body margin. Ex-
cretory ampullae cojled, spherical to ovoid but
not elongate. Vagina muscular, with distinct dis-
tal sphincter.

TYPE SPECIES
Craspedella spenceri Haswell, 1893

OTHER SPECIES
Craspedella gracilis sp, nev.
Craspedella pedum sp. nov.
Craspedella shorti sp. nov.
Craspedella simulator sp. nov,
Craspedella yabba sp. nov.

ETYMOLOGY

Haswell (1893) provided no derivation of the
name. Clearly it is from kraspedon (Greek; mas-
culine; edge). The diminutive does not change the
gender, so Craspedella is masculine meaning
‘little edge’, a reference to the dorsal papillate
ridges.

FIG. 1. Scanning electron micrographs of genera of Craspedellinae from type localities. Specimens fixed
HW/Form. A, Craspedella spenceri Haswell, 1893. Note prominent lamellae. Scale = 500рт.В,
Heptacraspedella peratus gen. et. sp. nov. Scale = 500um. C, Zygopella pista gen. & sp. nov. Scale = 2008m:
Specimens fixed 3% glutaraldehyde. D, dorsal view of left lateral tentacle of Zygopella stenota sp. nov. showing
rugose crenulations. Scale = 201m. E, posterior end of Zygopella pista showing paired posterior pits. Scale =

50шт.

Craspedella spenceri Haswell, 1893
(Figs 1A,2,3, 4A, 10A,E)

MATERIAL EXAMINED

TYPE SPECIMENS (Lost, see remarks).

NEoTYPE: ОМСІ.19489 (WM), ex Cherax destructor
Clark, 1936 from Condamine R., Warwick, QLD
(28"11.39'8; 151°57.50'Е) 24/0ct/1992 Sewell К.В.
& Sewell S.G. HW/Form/Hx.

OTHER MATERIAL: from QLD:- ex Cherax destructor
same data as neotype HW/Form/H&F QMGL18$490-
18491 (LS[1.1 p, 4/Aug/1994 Sewell К.В. & Joffe B.
HWideP QMGL18610 cirrus inverted (CP[2], 2 adult
specimens), QMGLI8611 cirrus everted (CP[2], 2
adult specimens), 1/5ер/1994 Sewell К.В. HW/deF
QMGL18612 cirrus inverted (CP[8], 8 adult speci-
mens) from Western R., 1.5km from Winton on
Jundah Rd., QLD (22°24,2'S; 143'02.2'E)
22/Nov/1990 Cook S/Cannon L.R.G. Form/Hx
QMGL18492 (WM): from Thompson R. al Longreach
Waterhole, Longreach, QLD (23°24.7'S; 144°13.8'E)
2/0cU1990 Cannon L.R.G. & Sewell К.В.
HW/Form/Hx QMGL18493-18498, Bouin/H&E
QMGL18499-] 8500 (LS[1,1])- from Marlong Ck., Mt
Moffat N.P., QLD (25'02'S; 14754" E) 26/Sep/1986
Monteith N.C. AVHx QMGLI18501-18503 у:
from Dawson R., Taroom, QLD (25'39'5; 149°48'Е)
3/Dec/1986 Cannon L.R.G. & Jennings J. Form/Hx
OMGL18504 (WM): from Bungil Ck., Roma, OLD
(26'30'S; 148'48' E) 2/Dec/1991 Cannon LRG, &
Jennings J. Form/H&E QMGLIS505 (WM),
FAA/H&E QMGL18506 (LS[5]: From Eukey, dam at,
QLD (2846.25; 151°59.2`Е) 18/Apr/1990 Cook S./
Cannon L.R.G. Form/Hx QMGL 18507-18508 (WM),
АРГО Form/Hx QMGL18509 (WM),
QMGL18510 (LS[1]: from Willows gemfield, in dam
beside road, OLD (23'45'S; 14725 E) 20/Sep/1990
Cook S./Cannon L.R.G. Form/H&E QMGLI18511-
18513 (LS[1.1,1]:- ex Cherax depressus complex
sensu Riek, 1951 from Wallaby Ck., on Henderson Rd.
0.2km from Mt. Cotton Rd. junction, Sheldon, QLD
(27734,218; 153°12,78°E) 22/Sep/1994 Sewell K.B.,
Joffe B., Solove: J.V. & Solovei S.B. Form/Hx
QMGLIS514-18516 (WM):

From NSW:- ex Cherax desiructor, from Lake
Madgwick, LI. N.E, campus Armidale, NSW (30'31'8;
15140 E) 23/May/1991 Zoology Dept U.N.E.
Bouin/Hx QMGL18517-18523 (WM), Boum/H&E
QMGL18524 (flattened specimen LS [2]),
QMGL18325-18527 (LS[1,1,2]k from Yarunga Ck.
irib.. L.2km NW Fitzroy Falls, Morton N.P., NSW
(4738,48; 150728.4' E) 19/Sep/1991 Cannon L.R.G,
& Sewell К.В, Form/Hx QMGL18528-38529 (WM),
QMGL1I8530-18531 (LS[L,1), HW/Form/H&E
QMGLIS532 (15111):

From SA:- (all collected by Beveridge I., fixed in
Berland' s fluid and stained jn Mayer's Haemalum as
wholemounes); ex Cherax destructor from Lake Al-
exandrina, via Clayton (35'25'S; 139"10'Е)
20/Jan/1989 QMGL1I8533: fram Cooper Ck.

MEMOIRS OF THE QUEENSLAND MUSEUM

26/Nov/1988 QMGL18534: from the Narrows, via
Clayton 2/Dec/1988 QMGL 18535: from Bordertown
(3618'S; 140'46' E) 31/Oct/1988 QMGL 18536; from
Lake Merreti (34'01'S; 140°46'E) 3/Dec/1988
QMGLGL 18537; from Avenue Ra. 18/Oct/1988
QMGL18538: from Mt. Benson (37°02'S; 139.49'E)
I3/0ct/1988 QMGL18539: from Bool Lagoon
(STOSS; 140743'E) 4/Nov/1989 QMGL 18540,

DESCRIPTION

External. Body from posterior margin 10 tip of
tentacles 1109um, to eyes 76 шт long and
522um wide. Posterior disc 304pm in diameter;
peduncle 162um in diameter, Transverse body
ridges form pronounced lamellae divided into
lobes tipped with clusters of papillae. Eptdermis
about 2-3um high dorsally and ventrally,

General Anatomy. Pharynx 65um across. Ex-
cretory ampullae about 7Ojim across, Eyes about
lum across.

Reproductive System. Female. Ovary 90m
across. Vesicula resorbiens 70um long, 35um
wide, wall about LOum thick, embedded m gut
wall. Seminal receptacle about 100um long and
27um wide. Vagina long.

Male. Anterior testes about 1 104m long, 70m
wide, Posterior testes about 110m long, 70um
wide, Seminal Vesicle 80pm long, 50pm wide.
Copulatory bulb 69jim long, 62pm wide, with
ejaculatory sac. Prostate duct reservoirs parallel.
Cirrus (based on 10 adult fully inverted speci-
mens fixed HW/deF ex QMGL18611 and
GL18612) 176-196um (x=185um) long in total,
Shaft funnel-shaped, curved, thick walled, with
distal region less than length of introvert; proxi-
mal opening 46-761m (x-60pum) wide, with nar-
row rim. Introvert strongly. curved, 13-171m
(x=]5um) wide at base, longer side 71-33m
(x=78um) long. shorter side 38-494 m (х=43шт)
long (i.e. introvert about 5 times longer than
width of introvert base), with clearly asymmetri-
cal swelling 1,e, much wider on longer side, distal
opening 20-27um (x-24um) wide.

Hosts, Cherax destructor Clark, 1936; Cherax
depressus group sensu Riek, 195] : Parastacidae,

Lacality. Murray-Darling river system and east
coast streams.

REMARKS

Type specimens were not located despite a
careful search of the collections al (be Australian
Museum, Sydney and at the McLeay Museum at
the University of Sydney. Haswell worked in
both institutions, but evidently failed to lodge
type specimens of the worms he described, A
search for data which might indicate the localities

CRAYFISH ECTOSYMBIONTS

FIG 2. Craspedella spenceri Haswell, 1893, Neotype. Scale = 250рт.

from which he obtained crayfish hosts and thus
indicate type localities failed to provide useful
information. In Haswell's time the crayfish As-
tacopsis bicarinatus Gray, 1845 was the name for
what is now known to be several species of
Cherax found in eastern Australia. Cherax de-
structor, however, is by far the most common and
wide spread of species and is found throughout
the Murray-Darling system. The specimens we
obtained from this host conform closely with the
majority of Haswell's description and thus con-
firm the validity of the species. However,

Rhabdite
tracts

Eyes

Excretory
ampulla

403

Haswell (1893) apparently confused
the cirrus of C. spenceri, in our view
the most important taxonomic char-
acter, with that of another closely
related species described below (see
Haswell, 1893 Plate XIII, Fig. 21).
Moreover, the widespread distribu-
tion of C. spenceri may result in
some variation in body size and form
between localities and hosts (see
below section on Craspedella sp.
from Cherax albidus Clark, 1936).
Since nomenclatural instability
could potentially develop and thus
circumstances are somewhat excep-
tional, a neotype has been erected

Pharynx
Rhabdite

glands Craspedella gracilis sp. nov.

(Fig. 4B)

MATERIAL EXAMINED

HorLoTvPE: QMGL18429 (WM), ex
Cherax depressus from Marlborough.
créek beside caravan park, QLD
(22°49.2'S; 149°53.2'Е) 21/Sep/1990
Cannon L.R.G. & Sewell K.B.
HW/Form/Hx.

PARATYPES: Same data as holotype,
QMGL18430-18431 (WM), QMGL
18432 (LS[1]).

OTHER MATERIAL: Same data as holo-
type, HW/Form/Hx QMGL 18433-
18435 (WM), QMGL18436 (LS[1}),
HW/AFA/H&E QMGL18437 (WM),
HW/Form/deF QMGL18613 cimus in-
verted (CP[2], 4 adult, 1 young speci-
mens).

DESCRIPTION

External: Body from posterior
margin to up of tentacles 510-
605m (х=550Нт), to eyes 323-
391m (x-354um) long and
170-198um (x-187pm) wide; Pos-
terior disc 102-115ит (х=106рт)
in diameter; peduncle 58-68um
(х=62рт) in diameter. Transverse body ridges do
not form lamellae. Epidermis about 2-3шт high
dorsally and ventrally.

General Anatomy, Pharynx 27-34,1m
(x-30jum) long, 30-34рт (x-33pm) wide, Gas-
trodermis about 251m high. Excretory ampullae
24-34um (x-31jm) long, 10-17um (x=15p1m)
wide. Eyes about 8um across.

Reproductive System. Female. Ovary 30-344m
(х=31рт) across. Vesicula resorbiens 45-68)1m

(x=$8um) across, wall 3-5um thick. Seminal re-
ceptacle about Збут long, 941m wide.

Male, Anterior testes about 34-41um
(x=39,1m) long, 25-30um (x=281m) wide, Pos-
tenor testes about 52-68um (x=60um) long and
29-41um (x=35um} wide. Seminal vesicle 54-
60m (x-561im) long, 25-3611m (x=3 lum) wide.
Copulatory bulb, 27-34um (xX=31lum) long, 30-
37pm (x-33um) wide, with ejaculatory sac.
Prostate duct reservoirs parallel. Cirrus (based on
4 fully inverted adult specimens ex
QMGLIS6]3) 157-168um (х=163рт) long in
total. Shaft narrow, goblet-shaped, curved, thin-
walled, with distal region about same length as
introvert; proximal opening 22-29um (x-24um)
wide, with narrow rim. Introvert slightly curved,
6-7рт (x= Tum) wide at base, longer side 61-
6lum (x=5lum) long, shorter side 44-52jtm
(X=48}im) long (з.е. introvert about 9 times longer
than width of introvert base), with narrow asym-
metrical swelling i.e. wider on longer side, distal
opening angled, about 6-8um (x=7 um} wide,

Hosts. Cherax depressus complex sensu Rick,
1951: Parastacidae.

Locality. Marlborough, central eastern QLD.

ETYMOLOGY
Latin, gracilis, slender, refernng to the slender-
ness of the cirrus.

REMARKS

The slender, delicate cirrus clearly dis-
tinguishes this species. In wholemounted speci-
mens the cirrus shaft is often severely contorted,
and the introvert considerably narrowed, presum-
ably as a result of ihe effects of fixation,

Craspedella pedum sp. nov.
(Figs 4C, 10C)

Craspedella sp, Sewell & Саппоп,1995: 151; Sewell
& Whittington, 1995: 1121; Watson, Rohde &
Sewell, 1995: 131.

MATERIAL EXAMINED

HOLOTYPE: QMGLIB846] (WM), ex Cherax
quadricarinatus (von Martens, 1868) from University
of Queensland aquaculture ponds, Pinjarra Hills, OLD
(27°32.38'S; 15255, 1 8' E) 15/May/i992 Sewell К.В.
HW/Form/Hx,

PaRATYPES:Same data as holotype, QMGL 18462
(WM), 19/Арг/1991 HW/Form/Hx QMGL18463
(WM), 30/Mar/1992 Bonin/ H&E QMGL18464
Ө), 22/Dec/1992 Bouin/MT QMGLI8465
1.511]).

MEMOIRS OF THE QUEENSLAND MUSEUM

OTHER MATERIAL: ex Cherax guadricarinatus (Von
Manens, 1868) from University of Queensland aqua:
culture ponds, Pinjarra Hills, QLD (27°32.38'S;
152'55.]8'E) 1991-1993 — HW/Form/Hx
QMGL 18466-18472 (WM), Bouin/Hx QMGL18473
(LS[1]), Form/HA QMGL18474 (LS[1]), Form/HIH
QMGL18475 (LS[1]), HW/deF QMGL18626 cirrus
everted (CP[5| 10 adults, 15 juveniles or damaged):
(Mitchell R. stock) from Walkamin, D.P.1. Research
Station aquaculture ponds, QLD (17°08"S; 14525" E)
25/Sep/1990 Cannon L.R.G. & Sewell К.В. AFA/Hx
QMGL18476 (WM), Bouin/Hx QMGL 18477 (WM):
from East Leichardt R., 50km SW Mi. Isa, QLD
(20°56'S; 139°45°E 3/May/1993) Monteith G.B.
Al/Hx QMGLI18&478-1 8488 (WM).

DESCRIPTION

External. Body from posterior margin to tip of
tentacles 739-896pm (x-800pim), to eyes 497-
6l6um (X=544um) long and 223-346um
(x=278pm) wide. Posterior disc 157-186дт
(х=171рт) in diameter; peduncle 69-107рт
(x=83,im) in diameter, Transverse body ridges do
not form lamellae. Epidermis about 2-3m high
dorsally and ventrally,

General Anatomy. Pharynx 37-48pm
(x=43um) long, 27-390 m (х=33рт) wide, Gas-
trodermis about 30pm high. Excretory ampullae
about 44-62um long (x-51um) long, 27-451m
(x-34um) wide, Eyes about 141m across.

Reproductive System. Female. Ovary 31-55um
(х=45рт) long, 27-47ит (х=37ит wide). Ves-
icula resorbiens 91-130pm (x-106jum) long, 56-
39um (x-721m) wide, wall about 10pm thick.
Seminal receptacle about 34um long, 11mm
wide.

Male. Anterior testes 46-67рт (х=5&т) long,
31-53um (х=45рт) wide. Posterior testes 72-
S4um (x=81 um) long, 34-50um (x—44jum) wide.
Seminal vesicle strongly muscled 105-144ит
(x=118um) long, 41-58,1m (x=48,im) wide. Cop-
ulatory bulb 73-861m (x=79um) long, 73-9811m
(x283jum) wide, with ejaculatory sac, Prostate
duct reservoirs parallel. Cirrus (based on 10 adult
specimens ex QMGL18266) 236-283um
(x=264um) long in total. Shaft funnel-shaped,
strongly curved (reflexed) distally; proximal
opening 69-1 121m (x=96uLm) wide, with narrow
rim; outside wall of reflex widened at point of
introvert ‘eversion’ muscle insertion. Introvert
permanently everted, forming rigid flange 48-
59um (х=54шт) long; without obvious swelling.
Spines reduced to be flat, overlaid, "welded
together’ and ‘plate-like’.

Hosts. Cherax quadricarinatus (Von Martens,
1868): Parastacidae.

Locality. Queensland.

CRAYFISH ECTOSYMBIONTS

" Vesicula resorbiens

: Seminal receptacle
Ni

Ejaculatory sac eS

Common genital i
atium
Copulatory bulb

FIG, 3. Reproductive structures of Craspedella spenceri. Scale = 50m,

ETYMOLOGY

Latin, pedum, shepherd's crook, referring to the
u-shaped reflex of the distal portion of the cirrus
shaft. This is a noun in apposition.

REMARKS

Morphology of the male cirrus is unique and
serves to distinguish this species. The muscle
insertion point of the introvert 'eversion" muscle
on the reflex becomes thicker with age. The ab-
errant form of the apparently permanently ever-
ted introvert required that the standard
measurement protocol had to be modified.

Craspedella shorti sp. nov.
(Figs 4D, 10D)

MATERIAL EXAMINED

HOLOTYPE: QMGL18450 (WM), ex Cherax
depressus from roadside gully on Mumford Rd., Nar-
angbah, QLD (27°12.34°5; 152°57.42’E) 18/Mar/1992
Short J./Sewell К.В, HW/Form/Hx.

PARATYPES: Same data as holotype, QMGL.18451,
(WM): ex Cherax depressus from Wallaby СК, on
Henderson Rd, 0.2km from Mt. Cotton Rd. junction,
Sheldon Brisbane, QLD (27°34.21'S; 153712.78'E)
Cannon L.R.G & Joffe B/Sewell К.В, 14/Sep/1994
Form/Hx QMGL18452 (WM), Bouin/H&E
QMGL18453 (LS[1/1]).

OTHER MATERIAL: Same data as holotype,
HW/Form/Hx QMGL18454-18455 (WM); ex Cherax
depressus from Wallaby Ck., on Henderson Rd. 0.2km
from Mt. Cotton Rd. junction, Sheldon Brishane, QLD
(27'34.21'S; 153°12.78'Е) Cannon L.R.G & Joffe

Vitelline ducts

"Сын,

405
B./Sewell K.B. 14/Sep/1994
Form/Hx QMGL18456-18460

(WM), HW/deF QMGL18614 cimus
inverted (CP[3], 3 adults),
QMGL18615 cirrus everted (CP[5], 5
adult specimens).

DESCRIPTION

External. Body from posterior
margin to tip of tentacles 414-
761um (x=608pim), to eyes 352-
Si5um (x=427um) long and
311-435um (x-327pm) wide.
Posterior disc 137-140um
(X=138um) in diameter: peduncle
65-904m (X=78um) in diameter.
Transverse body ridges do not
form lamellae. Epidermis about
2um high dorsally and ventrally.

General Anatomy. Pharynx 32-
72um (x=56um) long, 43-EOjtm
(x=58um) wide. Gastrodermis
about 27pm high. Excretory am-
pullae 45-60um (x= 53pm) long,
21-30um (x-261m) wide. Eyes about 13jim
across. Posterior glands present and discharging
in two postero-lateral regions (see remarks).

Reproductive System. Female, Ovary 40-434am
(x=4 11m) across, Vesicula resorbiens 93-126
(x=1] lum) long, 93-724um (x=78mpm) wide,
wall 8-1lpm thick, Seminal receptacle about
50um long, lum wide.

Male. Anterior testes about 54-68um
(х=60рт) long, 36-47um (х=43ит) wide. Pos-
terior testes about 74-91um (x=81um) long, 40-
50m (x24Sym) wide. Seminal vesicle 97-1 124m
(x=103pm) long, 40-49um (x-43u m) wide. Cop-
ulatory bulb 83-10] um (x-93pm) long, 62-75um
(х=68рт) wide, without ejaculatory sac. Prostate
duct reservoirs at nght angles (except in most
proximal region of the bulb). Cirrus (based on 7
adult specimens (3 inverted, 4 everted) ex
QMGL18614 and QMGL18615) 153-1691m
(х=161рт) long in total. Shaft cone-shaped,
curved, thick walled; proximal opening 29-38m
(x=32,1m) wide, with very thick rim. Introvert not
curved, 10- 15um(1 lim) wide at base, both sides
almost same length, longer side 45-
47um(x-46lm] long, shorter side 42-
47um(x=44um) long (i.e. introvert about 4 times
longer than width of introvert base), with near
symmetrical swelling. distal opening 13-
174m(x=15tim) wide.

Hosts. Cherax depressus complex sensu Riek,
1951: Parastacidae.
Locality. south castern OLD.

Gonopore

406

ETYMOLOGY

For John Short, Crustacea Section QM who
collected the host from which the first specimen
was recognised

REMARKS

The prostate duct reservoirs in the copulatory
bulb being at right angles to the long axis of the
bulb form a distinctly baccate pattern which
serves to distinguish this species, as does the thick
proximal rim of the cirrus shaft. Postero-lateral
glands similar to those observed for this species
have been described previously (see Cannon,
1993). The nature of similar glands from Tem-
nocephala minor are the subject of à separate
study by Cannon & Watson (in press).

Craspedella simulator sp. nov.
(Figs 4E, 10Р)

Craspedella spenceri; Cannon & Jennings, 1987

MATERIAL EXAMINED

HoLorvreg QMGL1354] (WM), ex Cherax dispar
from Woodgate Lagoons, beside Childers tà Woodgate
Rd.. QLD (25°07.4'S; 152°30.6'E) 6/ Apr/ 991 Sewell
K.B. HW/Form/Hx,

PARATYPES: Same data as holotype, QMGL18542-
18543 (WM); QMGL 18544 (LS[1].

OTHER MATERIAL: From QLD; ex Cherax dispar
same data as holotype, QMGL18545-18553 (WM),
QMGL18554(LS[1]), 18/Sep/1990 Cannon L.R.G. &
Sewell К.В, HW/Form/Hx QMGLI8555 (WM),
HWiForm'deF ОМОГ 18816 cimus inverted (CP[2]. 5
adult, 2 young specimens); also ex Cherax dispar fram
Gap Ck., in Gap Ck. Reserve picnic nd, Brook-
Dekt QLD (27-28. 68°S;152°55.71'E) J/Sep/1984 Hall
N. & Francis S. Bouin/Hx OMGL 18556-18557 (WM),
Bouin/H&E QMGL18558 (TS[2]). 12/Dec/1984 Can-
non L.R,G. FAA/Hx QMGL18559-18563 (WM),
FAA/H&E QMGL18564-18565 (LS[1.1]: from Wal-
laby Ck., on Henderson Rd. 0.2km from Mt. Colton
Rd. junction, Sheldon Brisbane, OLD (27°34.21'S;

153*12,78' E) Sewell K.B., Joffe B., Solovei 1, V. &
Solovei S.B. 19/Scp/1994 Form/Hx QMGL18566-
18573 (WM), 14/Sep/1994 Cannon L.R,G. & Joffe B./
Sewell К.В. deF QMGLIS6/7 cirrus everted (CP[1],
1 adult specimen): ex Cherax depressus from Wallaby
Ck. on Henderson Rd, 0,2km fram Mt, Cotton Rd.

junction, Sheldon, Brisbane, QLD (27734.21'S;

15312.78 E) Sewell K.B., Joffe B., Solovei LV. Ж
Solovei S.B. 19/Sep/1994 Form/Hx QMGLI&S74-
18576 (WM): ex Cherax destrucsor from Thompson
R. at Longreach Waterhole, Longreach, QLD
(22°24.7'S; 134713.8 E) 2/0cU/1990 Cannon L. К.б. &
Sewell K.B. FAA/Hx QMGLI8577 (WM),
HW/Form/Hx OMGLISS78 (WM): from Longreach,
QLD (23278; 144" I5 E) L3/Dec/1990 Cook S./Can-
non L.R.G. Form/H&E QMGL18579-18580 (WM):

MEMOIRS OF THE QUEENSLAND MUSEUM

from Condamine R,, Warwick, QLD (28'11.39'8;
1557.50" E) 24/0ct/1992 Sewell К.В, & Sewell S.G.
HW/Form/Hx QMGL18581-18589 (WM).

From NSW:- ex Cherax destructor from a tributary of
Yarunga Ck, 1.2km NW Fitzroy Falls, Morton N.P.,
NSW (34'38,4'S; 150728.4'E) 19/Sep/1991 Cannon
L.R.G. & Sewell К.В. HW/Form/Hx QMGLI8590-
18591 (WM), QMGL18592 (LS[1]: ex Cherax
cuspidatus from Wauchope, roadside swamp beside
‘Timbertown'. NSW (31'28.4'S; 152'42.8'E)
15/Feb/1992 Sewell К.В. & Sewell S. G. HW/Form/Hx
QMGL18593-18594, Form/Hx QMGL18595-18598
(WM), QMGL18599-[8600 (LS[1,1 ]).

From SA: (all collected by Beveridge 1, fixed in
Berland's fluid and stained in Mayer's Haemalum as
wholemounts); ex Cherax destructor from Tod К.

(34°30'S; 135°51°E) 9/Mar/1989 QMGL18601- from
Springbett, Gawler (34°36°S; 138'45' E) 23/Jan/1989
QMGL1t8602: from Murray R., Murray Bridge
(35'07'8; 139" 16" E) 10/Feb/1989 QMGLLI8603: from
Onkaparinga R. (35°00'S; 138'49'E) 22/Nov./1989
QMGLi8604: from Light R., Kapunda (34°21°S;
138'55' E) 1/Mar/1989, QMGL18605: from Inman К.
(38'30'S; ]38'3]'E) 19/Nov./1988 QMGL 18606;
from Strawberry Hill Springs (34°31°S; 135'43'B)
YMan 1989 OQMGLI8607: from Broughton R., Spal-

ding (33°30'S; 138°37°E) 39/Mar/1989" QMGLi8608:

from Gerard Yabbie Farm, Berri (34°17'S; 140°36'Е)
19/Jan/1989 QMGL18609.

DESCRIPTION

External. Body from posterior margin to tip of
tentacles 476-506um (x=486,im), lo eyes 313-
34Tum (X-33lum) long and 205-210um
(х=207рт) wide; Posterior disc 95-1 10рт
(x-103jm) in diameter; peduncle 51-58 т

(x-55um) in diameter. Transverse body ridges do
not form lamellae. Epidermis about 2umhigh
dorsally and 3pm high ventrally.

General Anatomy. Pharynx 30-3
(x=3 11m) long, 34-36um (x-35|m) wide, Gas-
trodermis about 20pm high. Excretory ampullae
about 40m long, 20um wide. Eyes about 9um
астозз.

Reproductive System. Female. Ovary 32-3бит
(x=34ym) across. Vesicula resorbiens 54-60um
(х=56рт) across, wall 5-1Ourm thick. Seminal
receptacle about 584m long, 191m wide. Vagina
with pronounced lateral pocket,

Male, Anlerior testes about 30-440т
x-38pm) long, 18-400 (x23lIjm) wide. Pos-
terior lestes about 38-780 m (х=5бит) long and
27-35um (х=31ит) wide. Seminal vesicle 54-
63um (x=58um) long, 23-260 m (x-225jun) wide,
Copulatory bulb, 34-40um (х=38итп) long, 44-
49um (x=47 ит wide, with ejaculatory sac. Pros-
tate duct reservoirs parallel, Cirrus (based on 5
adult specimens ex QMGL18816) 176-186um

CRAYFISH ECTOSYMBIONTS 407

FIG 4. Nomarski interference photomicrographs of cirri of adult worms ex type host and locality unless stated:
A, Craspedella spenceri (i) inverted; (ii) partially everted. B, Craspedella gracilis sp. nov., inverted; C,
Craspedella pedum sp. nov.; D, Craspedella shorti sp. nov. (i) inverted, (ii) partially everted; Craspedella
simulator sp. nov. (i) inverted, (ii) inverted ex Cherax dispar from Wallaby Ck., Sheldon, QLD, (iii) partially
everted ex Cherax dispar from Wallaby Ck, Sheldon, QLD. Scale = 100m.

(x=180pm) long in total. Shaft, goblet-shaped, —trovert curved, 11-13рт (х=12рт) wide at base,

slightly curved, thick-walled, with distal region longer side 70-771m (х=72рт) long, shorter side

shorter than length of introvert; proximal opening 64-70ит (x=66pm) long (i.e. introvert about 6

38-43um (x=41p1m ) wide, with narrow rim. In- times longer than width of introvert base), with

408 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Nomarski interference photomicrographs of cirri of adult worms ex type host and locality unless stated:
A, Craspedella yabba sp. nov., (i) inverted (ii) partially everted; B, Craspedella sp., inverted; C,
Heptacraspedella peratus gen. & sp. nov., part everted; D, Zygopella pista gen. & sp. nov., partially everted; E,
Zygopella deimata sp. nov. partially everted; Е, Zygopella stenota sp. nov., inverted, Scale = 100um.

clearly asymmetrical swelling і.е. much wider on Locality. eastern Australia.
longer side, distal opening 11-17рт (x=1441m) wide.

Hosts. Cherax cuspidatus Riek, 1969; Cherax ETYMOLOGY
depressus complex sensu Riek, 1951; Cherax Latin, simulator, an imitator which refers to the
destructor Clark, 1936; Cherax dispar Riek, close resemblance of this species to that of C.
1951: Parastacidae. spenceri. This is a noun in apposition.

CRAYFISH ECTOSYMBIONTS

REMARKS

Although superficially resembling C. spenceri
with which it frequently co-inhabits the branchial
chamber this worm is generally smaller, has a
more elongate body shape, much less pronounced
transverse body ridges, ie. they do not form 1а-
mellae, a goblet-shaped cirrus with a narrower
introvert, and a smaller vagina which has a pro-
nounced lateral pocket (Fig. 10F).

Craspedella yabba sp. nov.
(Fig. 5A)

MATERIAL EXAMINED

HoLorvYe QMGL18438 (WM), ex Cherax
depressus from Conondale Ra,, State Forest 792, QLD
(26'47'S; 15232? E) 16/Sep/1994 Smith G/Scwell
K.B. Bouin/Hx.

PARATYPES: Same data as holotype, Form/Hx
QMGL18439 (WM), Bouin/H&E QMGL18440 (FS
111), GL 18441 (LS[1]).

OTHER MATERIAL: Same data as holotype, Form/Hx
QMGL18442 (WM), Bouin/H&E QMGL18443
(LS(1]), HW/deF QMGL 18618 cirrus inverted (CP[1],
I adult specimen), QMGL 18619 cirrus everted (CP[2],
2 adult specimens): ex Cherax dispar from Booloumba
Ck. at first road crossing E of Little Yabba Ck. Rest
Area (26°37.0'S; 152'39.1'E) 23/Mar/1990 Cannon
L.R.G. & Sewell K.B. HW/AFA/Hx QMGL18444
(WM), Bouin/H&E QMGL18446-18447 (LS[1,1,):
from Conondale Ra., QLD (26'44'S; 152°43'E)
17/Nov./1983 Cannon L.R.G. Form Hx QMGL 18445
(WM), FAA/H&E QMGLIS448 (LSII]),
QMGL18449 (LS[2]).

DESCRIPTION

External. Body from posterior margin to tip of
tentacles 336-380рт (x —358,un), 10 eyes 255-
285um (x=270um) long and 209-22 :ит

x-211um) wide. Posterior disc 951m in diame-
ter; peduncle about 58m in diameter. Transverse
body ridges do not form lamellae. Epidermis
about 2-5um high dorsally and ventrally-

General Anatomy. Pharynx 25-27um
(x=26um) long, 25-27um (х=2бит) wide, Gas-
trodermis about 37-444 m high. Excretory ampul-
lae about 284m long, 20um wide. Eyes about
1211m across.

Reproductive System. Female. Ovary 40-41 um
(x=41 pm) long, 18-211 (x=20pm) wide. Ves-
icula resorbiens 36-544um (x=45pm) across, wall
6-9um thick. Seminal receptacle about 9-1 Цит
across. Vagina with distinct lateral pocket.

Male. Anterior testes about 27-43um
(¥=35um) long, 31-37um (х=34рт) wide. Pos-
terior testes about 58-66jum (х=621т) long. 27-
28um (x=28um) wide. Seminal vesicle 43-45um

(x=44um) long, 23-25um (x=24{1m) wide. Cop-
ulatory bulb, 31-31 um (x=3 lum) long, 39-40pm
(x-40um) wide, with ejaculatory sac. Prostate
duct reservoirs parallel. Cirrus (based on 3 adult
specimens (1 inverted, 2 everted) cx
QMGL18618) 188-19511m (x-1911m) long in
total. Shaft, goblet-shaped, curved, me-
dium/thick-walled, with distal region longer than
length of introvert; proximal opening 43-45um
(x=44um) wide, with narrow rim. Introvert
slightly curved, 12-13рт (134m) wide al base,
longer side 65-67um (x=66p1m) long, shorter side
57-57шт (x=57pm) long (i.e. introvert about 5
times longer than width of introvert base), with
asymmetrical swelling 1.e. wider on longer side,
distal opening about 15m wide.

Hosts. Cherax dispar Riek, 1951; Cherax
depressus complex sensu Riek, 1951:
Parastacidae.

Locality. Conondale Ra., QLD.

ETYMOLOGY

A noun in apposition, yabba means forestin the
local Waka language of the Aboriginal people
and refers to the type locality, the Conondale
Ranges,

REMARKS

C. yabba is very close to C. simulator in general
body form and the morphology of the cirrus.
However, the cirrus of С. yabba has a shorter
introvert and the ratio of the length of the tubular
distal region of the shaft to the length of the
introvert is never less than 1:1 compared to that
for C. simulator which 15 clearly less than 1:1.
Moreover the intravert swelling on the longer and
shorter sides of the introvert are more equal in
thickness than those of C. simulator.

Craspedella sp. 1
(Fig. 5B)

MATERIAL EXAMINED

Ex Cherax albidus Clark, 1936 fram Dwyers Ck, 2km
SSW Mirrantawa, VIC (37°24°S; 142'23'E)
5/Oct/1991 Cannon L.R.G. & Sewell К.В.
HW/Form/Hx, QMGL 18420-18425 (WM), Form/Hx
QMGLI8426 (WM), HW/Form/deF QMGL 8620 cir-
rus inverted (CP[1 J).

REMARKS

The distinctiveness of the crayfish species
Cherax albidus, the host of these worms, from
Cherax destructor is uncertain (see Morrissy &
Cassells. 1992). The worms appear close to C.
simulator in general anatomy including the form

410

Rhabdlite
tracts

Eyes

Excretory
ampulla

Pharynx

Vitelline Rhabdite

glands

Gut

Glands

FIG. 6. Heptacraspedella peratus gen. et sp. nov. Ho-
lotype. Scale = 2504m.

of the transverse and posterior ridges, however,
they show some slight differences in the shape of
the cirrus which in the shaft at least (Fig. 5B)
resembles that of C. spenceri. The worms may
provea further distinct species. The small number
of specimens collected and the large geographical
range of C. destructor/albidus makes us hesitant
to erect a new species of Craspedella at this time.

Heptacraspedella gen nov.

DIAGNOSIS

Craspedeilinae with seven dorsal transverse pa-
pillate ridges, and behind the last ridge, four short
posterior papillate ridges consisting of raised

MEMOIRS OF THE QUEENSLAND MUSEUM

points radiating towards the posterior body mar-
gin. Excretory ampullae strongly coiled and elon-
gate. Muscles about seminal vesicle and
copulatory bulb relatively very strong. Distal
vaginal sphincter not pronounced.

TYPE SPECIES
Heptacraspedella peratus sp. nov.

ETYMOLOGY
Greek, hepta, seven, a reference to the number
of ridges or edges.

REMARKS

The number and form of the dorsal ridges are
quite different from those of Craspedella
spenceri and other new members of that genus.
The host, Euastacus, is also of a different genus;
Craspedella spp. have only been found on mem-
bers of Cherax. Parenchymal gland cell bodies
(?mucus glands) were observed positioned later-
ally between the anterior and posterior testes (Fig.
6).

Heptacraspedella peratus sp. nov.
(Figs 1B,5C.6,7, 10B)

MATERIAL EXAMINED

HoLorvPE: QMGL18403 (WM), ex Euastacus
bispinosus from Jimmys Ck., Grampians, at picnic
ground 6.5km WNW Mafeking, VIC (37°23'S;
142°34°Б) 5/Ocv1991 Cannon L.R.G. & Sewell К.В.
HW/Form/Hx.

PARATYPES: Same data as holotype, QMGL18404-
18405 (WM), Form/H&E QMGL18406-18407
(LS[1,1].

OTHER MATERIAL: Same data as holotype.
QMGL18408-18419 (WM), HW/Form/deF
QMGLLI8621 cirrus everted (CP[4], 9 adult (1 bent), 1
young specimen(s)),

DESCRIPTION

Exiernal. Body from posterior margin to tip of
tentacles 848-870um (х=862рт), to eyes 565-
580um (x-575um) long and 290-3044im
(x=300um) wide. Posterior disc 137-152um
(x=142um) in diameter; peduncle 72-72um
(x=72um) in diameter. Epidermis about 4m
high dorsally, about 5рт high ventrally.

General Anatomy. Pharynx 33-501m
(x=40um) long, 33-50um (x=40um) wide. Gas-
trodermis about 354m high. Excretory ampullae
elongate 65-б5рт (x=65um ) long and 25-251m
(х=25рт) wide. Eyes about 15 іт across.

Reproductive System. Female. Ovary about
SSum across. Vesicula resorbiens 116-123,1m

CRAYFISH ECTOSYMBIONTS

Gonopore Commoh genital
atrium

Vesicula resorbiens

i Seminal receptacle

4\1

Zygopella gen. nov.

DIAGNOSIS

Craspedellinae with a single trans-
verse papillate ridge lacking lohes
across the posterior body; and with a
pair of pits behind it each defined dor-
sally by a horseshoe-shaped papillate
ridge. Vagina musculature with dis-
tinct distal sphincter.

TYPE SPECIES
Zygopella pista sp. nov.

OTHER SPECIES
Zygopella deimata sp. nov.
Zygopella stenota sp. nov.

FIG. 7. Reproductive structures of Heptacraspedella peratus gen.

et sp. nov. Scale = 50um.

(х=118рт) across, wal] about [5.1m thick, em-
bedded in gut wall, not open to gut. Seminal
receptacle at least 121m long.

Male. Anterior testes about 70-80um
x=76j1m) across. Posterior testes about 108-
156m (x=131 4m) long and 40-50џрт (х=44рт)
wide. Seminal vesicle strongly muscled 58-
101ит (x=75pm) long, 43-45um (x-44um)
wide. Copulatory bulb 101-130um (x=118pm)
long, 80-87j1m (x=82p1m) wide, without ejacula-
tory sac. Prostate duct reservoirs oblique (except
in most proximal region of bulb). Cirrus (based
on 8 adult specimens (all everted) ex
QMGL18621) 76-104um (x-93um) long in
total. Shaft cone-shaped, not curved, thick
walled; proximal opening 51-894m (x-75um)
wide, with slightly thickened rim. Introvert 32-
35um (х=33рт) wide at base, sides about equal
length 38-42um (x-40pm) long (i.e. introvert
slightly longer than width of introvert base), dis-
tal opening unable to be determined accurately
due to lack of fully inverted specimens (about
10um wide), with wide, near symmetrical swell-
ing.
Hosts. Euastacus bispinosus Clark, 1941:
Parastacidae.

Locality. Jimmys Ck., Grampians, МІС,

ETYMOLOGY

Latin, peratus, western/edge, refering to having
3 host species at the western most edge of the
range of its genus (Euastacus).

ETYMOLOGY

Greek, zygon, yoke and Greek (feminine),
pella, cup, i.e, symmetrical cups.

REMARKS

The possession of only a single papillate ridge
and, instead of 4 posterior ridges, a pair of pits
each defined by a horseshoe-shaped papillate
ridge, clearly distinguishes Zygopella from
Craspedella and Heptacraspedella.

Zygopella pista sp. nov.
(Figs 1C,E,5D,8,9)

MATERIAL EXAMINED

HOLOTYPE: OQMGL18326 (WM), ex Cherax ten-
uimanus from Margaret R., at water catchment weir,
WA (33°57.0'S; 115°05'2E) 23/Jan/1992 Cannon
L.R.G. & Sewell K.B. HW/Form/Hx.

PARATYPES: Same data as holotype, QMGL 18327
(WM), QMGLI8328 (WM), Bouin/H&E
QMGL18329-30 (LS[1,1], Bouin/H&E QMGL 18331
(FS[3]).

OTHER MATERIAL: Same data as holotype,
QMGL18332-63 (WM), Bouin /H&E QMGL18564
(LS[1]), QMGL18365 (LS[2]), QMGL18366 (LS[1]).
QMG18367 (LS[1]), QMGL18368 (LS[I]».
HW/Form/deF QMGL18622 cirrus everted (CP[5]. 5
adult specimens): also ex Cherax tenuimanus from
"Margaret R. Marron Farm’, 10km SE Margaret R.,
WA Australia (34700.8'S; 115°09,5°E) 23/Jan/1992
Cannon L.R.G. & Sewell K.B. HW/Form/Hx
QMGL18369-18373 (WM), Bouin/H&E
QMGL18374 (FS[2]), QMGL18375 (LS[2]).
Bouin/H&E QMGLI!8376-18380 (LS[1]).

Rnabdite
tracts

Pharynx
Rhabdite
glands

Gut

Vitelline
glands

FIG. 8. Zygopella pista gen. & sp, nov, Holotype. Scale = 250ит,

DESCRIPTION
External. Body from posterior margin to tip of
tentacles 667-884um (х=748ит), to eyes 457-

MEMOIRS OF THE QUEENSLAND MUSEUM

high. Excretory ampullae 40-45um
(x242um) across. Eyes about
15pm across with short, narrow
pigment tails that descend ven-
trally.

Reproductive System. Female.
Ovary about 47um across. Ves-
icula resorbiens 100-120um
(х=107рт) across; wall about 3-
5pm thick, embedded in gut wall,
not open to gut. Seminal receptacle
about 10рт long.

Male. Anterior testes about 73-
871m (x-80jim) across. Posterior
testes about 73-108um (x-85um)
across. Seminal vesicle 80-110um
(x-99um) long, 36-50um
(x-243um) wide, strongly muscled.
Copulatory bulb 100-130Jum
(x-119um) long, 36-50um
(х=44рт) wide; with ejaculatory
sac. Prostate duct reservoirs
oblique. Cirrus (based on 5 adult
specimens (all everted) ex
QMGL18622) 54-661m
(x-61um) long in total. Shaft,
cone-shaped, curved, thin walled:
proximal opening 25-29um
(x-271m) wide with narrow rim.
Introvert 10-10um (10pm) wide at
base with junction between intro-
vert and shaft oblique, longer side
20-22um (х=21рт) long, shorter
side 16-17um (x=17um) long (i.e,
introvert about two times longer
than width of base), with thin,
asymmetrical swelling, i.e, wider
on longer side of introvert, distal
opening width not able to be deter-
mined accurately due to lack of
fully inverted specimens (about
7um wide).

Hosts. Cherax tenuimanus
(Smith, 1912): Parastacidae.

Locality. Margaret R., WA (wild
and in culture).

616um (x-5i3um) long, and 348-417um
(x=378um) wide. Posterior disc 137-150рт
(x=144um) in diameter; peduncle 72-87um
(x=78um) in diameter. Epidermis about 21m
high dorsally and ventrally.

General Anatomy. Pharynx 38-50um
(x=44um) across, Gastrodermis about 45um

ETYMOLOGY

Greek, pistos, faithful and refers to the retention
of this species with its host even when placed into
culture, a situation which sees the external natural
temnocephalid replaced with a translocated spe-
cies (Cannon & Sewell, 1994).

CRAYFISH ECTOSYMBIONTS

//
Seminal vesicle Р V

75

4& Prostate secretion

Ejaculatory sac ;

Со fet bull
pulatory bulb Gondpore Common genital

atrium

FIG. 9. Reproductive structures of Zygopella pista gen. et sp. nov. Seale

. Vesicula resorbiens

413

Walled; proximal opening 82-
86um (x=84jim) wide with thick-
ened rim. Introvert 16-17ym
(x=17um) wide at base, both sides
about 34um long (i.e. introvert
about two times longer than width
of base), distal opening unable to
be determined accurately due to
lack of fully inverted specimens,
with near symmetrical swelling.
Proximal spines very large, with at
least three large dagger-shaped
spines, about 30um long posi-
tioned centrally and apparently at-
tached to introvert base.

Hostis. C. rTenuimanus (Smith,
1912): Parastacidae.

= 50um. Locality. Inlet R., WA.
Zygopella deimata sp. nov. ETYMOLOGY .
(Fig. SE) Greek, deimatos, a fearful thing, referring to the
robust nature of the cirrus and its spines.
MATERIAL EXAMINED

HoLorvPE: QMGL18381 (WM), ex Cherax tën-
uimanus from Inlet R., on South Western Highway to
Walpole, WA (34°55.2'S; 116'34.2'E) 25/Jan/1992
Cannon L.R.G. & Sewell К.В, HW/Form/Hx.
PARATYPE: Same data as holotype, QMGL18428
(LS[2]).

OTHER MATERIAL: Same data as holotype,
HW/Form/deF QMGL 18623 cirrus cverted (2 adult
specimens).

DESCRIPTION

External. Body from posterior margin to tip of
tentacles 792рт, to eyes 540um long and 306m
wide. Posterior disc 122um in diameter; peduncle
65m in diameter. Epidermis about 2-3um dor-
sally and ventrally.

General Anatomy. Pharynx 35m long, 501m
wide; Gastrodermis about 60m high. Excretory
ampullae thick walled about 471m across. Eyes
about 1 21m across.

Reproductive System. Female. Ovary 50um
long, 21pum wide. Vesicula resorbiens 144um
across, wall 2 lum thick. Vagina with pronounced
distal sphincter and very prominent proximal lon-
gitudinal folds.

Male. Anterior testes about 831m across. Pos-
terior testes about 90jtm across. Seminal vesicle
90j.m Jong, 63m wide, strongly muscular. Cop-
ulatory bulb 100um long, 631m wide, without
ejaculatory sac. Prostate duct reservoirs oblique.
Cirrus (based on 2 everted specimens ex
QMGL18623) 119-121um (x=120pm) long in
total, Shaft cone-shaped, not curved, thick-

REMARKS

Only a few specimens were collected. How-
ever, the species is distinct, as the cirrus is large
and quite unlike the modest cone found in 2.
pista.

Zygopella stenota sp. nov.
(Figs 1D, 5F)

MATERIAL EXAMINED

HoLorYPE: QMGL18382 (WM), ex Cherax c.f.
quinquicarinalus from Inlet R., on South Western
Highway to Walpole, WA (34'55.2'S; 116342" E)
25/Jan/1992 Cannon L.R.G. & Sewell K.B. Bouin/Hx.
PARATYPES: Same data as holotype, OMGL18383-
18384 (WM), Bouin/H&E QMGL18385-18386
(LS[1.1].

OTHER MATERIAL: Same data as holotype.
QMGL18387-18389 (WM), Bouin/deF QMGL18624
(citrus inverted (CP[2]. 2 adult specimens),
QMGLÍ8625 cimus everted (CP[6], 6 adult speci-
mens): ex Cherax tenuitnanus from Inlet R., on South
Western Highway to Walpole, WA (34 55.2'5;
116°34.2'Е) 25/Jan/1992 Cannon L.R.G. & Sewell
K.B. HW/Form/Hx QMGL18390-18392 (WM),
QMGL18427 HW/Form/H&E (LS[1]): ex Cherax c.f.
bicarinatus from Deep К, tributary, circa 4km N Mt,
Pingerup, WA (3450.48; 116'32.4'E) 25/Jan/1992
Cannon L.R.G. & Sewell K.B. HW/Form/Hx
QMGL 18393-18402 (WM).

DESCRIPTION
External. Body from posterior margin to tip of
tentacles 476-570um (х=515рт), to eyes 335-

414 MEMOIRS OF THE QUEENSLAND MUSEUM

CRAYFISH ECTOSYMBIONTS

407um (x-3561m) long, and 251-272um
(x2261|im) wide. Posterior disc 103-122uUm
(x=1 llum) in diameter; peduncle 44-63um
(х=53шт} in diameter, Tentacles with very ru-
gose crenulations.

General Anatomy, Pharynx 18-25um
(x-22um) long, 25-374 m (x=3 lpm) wide, Gas-
trodermis about 35-+441m high. Excretory ampul-
lae 28-404 m across. Eyes about 9um across.

Reproductive System. Female, Ovary about
25m across. Vesicula resorbiens kidney-shaped
63-96ит (xz82yum) long, 43-59 (x=S1 um) wide,
walls about 8m thick. Seminal receptacle about
10pm long,

Male. Anterior testes about 70-100um
(x-861m) long, 37-49um (x=42um) wide. Pos-
terior lestes about 70-941 m (x-82y m) long, 44-
57pm (х=50Шт) wide. Seminal vesicle 59-7 11m
(х=65рт) long, 36-45um (x-4lum wide),
strongly muscled, it and copulatory bulb show a
distinct semi-spiral arrangement of muscles.
Copulutory bulb 57-65um (x-61jm) long. 33-
39um (х=3брт) wide, with accessory sac. Pros-
tate duct reservoirs parallel. Cirrus (based on 8
adult specimens except measurements of the
width of the distal introvert based on two inverted
specimens ex QMGL 18624 and 18625) 38-47
(x-43um) long in total. Shaft, cone-shaped,
curved, thin-walled; proximal opening 15-19pm
(x=16um) wide with narrow nm. Introvert 3-4jim
(х=3рт) wide at base, both sides 6-8um
(x=7pm) long (i.e. introvert about 2.5 times
longer than width of base), distal opening 3-4um
(x-4pm) with narrow, near symmetrical swell-
ing.

Hasts. Cherax c.f. bicarinatus (Gray, 1845); C.
c.f. guinquicarinatus (Gray, 1845); C. ren-
мітапих (Smith, 1912): Parastacidie.

Locality. Deep R. and Inlet R, WA,

ETYMOLOGY

Greek, stenotes, narrow referring to the narrow-
ness of the cirrus introvert.

415

REMARKS

Evidently quite close to the type species C.
pista, but with a characteristically narrow cirrus.
The seminal Vesicle and copulatory bulb show a
distinct semi-spiral arrangement of muscles. The
tentacles of all the specimens we observed had
extremely rugose crenulations independent of
fixation, compared to the other species (Fig. 1D),

DISCUSSION

For over 100 years Craspedella spenceri hus
been the only temnecephalan species recognised
with papillate posterior dorsal ridges. Other spe-
cies have possibly been confused with it (c.g.,
Cannon & Jennings, 1987), The small size and
delicate nature of these worms, and their cryptic
habitat in the branchial chamber, has evidently
ensured the richness of the fauna was overlooked.

The distinctive facies of this group of genera
with their well developed papillate ridges and
crenulate tentacles presumably relate to their hab-
itat, the branchial chamber. The relationship of
form, function and habitat are the subject of con-
tinuing detailed studies by one of us (KBS) on С.
pedum from the 'redclaw' Cherax
quadricarinatus.

We have recognised the Craspedellinae and
include in it 3 genera compnsing a total of 11
species of temnocephalans, all from the branchial
chamber of crayfish and characterised by posses-
sion of crenulate tentacles and one or more trans-
verse dorsal papillate ridges, The morphological
differences between this group and the remainder
of the Temnocephalidae are insufficient in our
opinion, to justify the re-erection of the family
Craspedellidae created hy Baer (1931), In this we
concur with Bresslau & Reisinger (1933), but the
similar facies of so many taxa indicates a sub-
family rank is appropriate,

As with many small worms, fixation Ip provide
consistent form for comparison 1s difficult. Cold
fixation usually ensures considerable distortion.
The hot water treatment was effective in produc-
ing uniform fixation of extended worms suitable

FIG. 10. Light photomicrographs: A, Craspedella spenceri, Longitudinal section through centre of specimen
QMGL18490 showing weak pharynx and 3 prominent lamellae. Scale = 200m. B, longitudinal section through
anterior end of Hepracraspedella peratus gen. & sp. nov. showing large, elongate excretory ampullae, Scale =
100pm. C, copulatory bulb of live Craspedella pedum sp. nov. showing ‘reduced’ ejaculatory sac contained
within the musculature of the bulb. Scale = 50m, D, copulatory bulb from wholemount of Craspedella shorti
зр. nov. Note absence of ejaculatory sac and baccate pattern formed by the prostate duct reservoirs. Scale =
Sum. E, vagina of Craspedella spenceri cleared in de Faure's medium to reveal shape of vaginal cavity. Scale
= 50р. Е, vagina of Craspedella simulator sp. nov, cleared in de Faure's medium to reveal shape of vaginal
cavity and prominent lateral pocket, Scale = 50m,

416

for wholemounts, although it could be considered
only ‘adequate’ for preparation of specimens for
sectioning, Similarly the quality of the SEM fix-
ation for this study was only adequate and this
prompted the study of Sewell & Cannon (1995)
lo determine an optimum fixation regime for the
SEM of Craspedella pedum.

In the wild, C. spenceri and C. simulator oc-
curred together in the branchial chamber of 2
species of Cherax. Co-occurrence was alsa oh-
served with Z stenota and Z. diemara on С.
tenuimanus, It is of ecological interest that differ-
ent yet closely related species of Craspedellinae
co-inhabit the branchial chamber microhabitat of
a single crayfish host. In particular, questions are
raised as to how such a niche is partitioned.

The strong association of the Craspedellinae
with Cherax spp. crayfish raises issues of host
phylogeny, taxonomy and biogeography ee. the
occurrence of Zygopella spp. on Cherax of West-
em Australia and not Craspedella spp. which are
confined to Cherax of eastern Australia. These
questions can hopefully be addressed after anal-
yses of the distribution of many potential new
species of temnocephalan species collected from
Australian Cherax spp. in the course of fieldwork
associated with the present study. Evidence for à
high diversity of temn alan species associ-
ated with Euastacus was presented by Cannon &
Sewell (1994).

It is perhaps not surprising that distinctive
Heptacraspedella occurs on the crayfish Euasta-
cus. The two crayfish genera Cherax and Euasta-
cus are distinct and show considerable
differences in habitat and life style. Cherax are
commonly found in slower, lowland or coastal
streams, often (but not always) with open canopy
above: Euastacus prefers fast, and often high
country, streams with a closed forest canopy. In
examination of many species of Euastacus from
Cape York to the Grampians this was the only
occasion on which gill temnocephalans (with the
exception of Didymorchis spp. which were not
included in this study) were detected. Most often
Ewastacus specimens were collected by hand
from the edges of streams and though many were
mature and were host to external temnocephalans
the crayfish were usually small. If large speci-
mens of Euastacus can be obtained from traps
other species of gill dwelling temnocephalans
may be found.

Our species descnptions rely mòre heavily on
descriptions of the cirrus than previous recent
hlications on Australian Temnocephalida (e.g.,
сктап, 1967; Cannon, 1991, 1993). The cirrus

t

MEMOIRS OF THE QUEENSLAND MUSEUM

introvert is a reliable character to separate species
because it is the cirrus structure which is first
formed in juveniles and its dimensions, unlike
those of the cirrus shaft, remain constant as the
worms age (unpublished observations), The use
de Faure's medium allows more accurate detail
of the morphology of the cirrus than has been
available in the past, We did not base the separa-
tion of species on the dimensions and arrange-
ment of the introvert spines, as these characters
could not always reliably be determined, particu-
larly from specimens fixed prior to placement in
de Faure's medium. Furthermore, the introvert
swelling was often difficult to observe in speci-
mens which were placed live into de Faurc's
medium as it a to clear more than for
specimens fixed previously, Moreover, when the
cimus was everted, the swelling was obscured.
Therefore, we excluded the introvert swelling
from our measurements of the cirrus.

The original description of Craspedella
spenceri Haswell, 1893 was based largely on the
lamellate, papillate frills of the species. We were
able to clearly recognise this species with large
lamellations and prominent lobes tipped with pa-
pillae, which we have called C. spenceri, These
features were observed even on small juvenile
worms. However, the cirrus figured by Haswell
(1893) does not conform to that which we ob-
served for this species. The goblet shape of the
shaft and the narrow width of the introvert sug-
gest that this cirrus belongs instead to C. simula-
tor which we observed to co-inhabit the branchial
chamber of the crayfish Cherax destructor over a
wide geographical area. Furthermore, Haswell
(1893) apparently figured the cirrus erroneous] y
as he drew the teeth of the inverted introvert as
pointing proximally not distally as we observed
for all the species we studied. Similarly, we did
not observe the ejaculatory sac to be discrete as
figured by Haswell (1893). In all species of
Craspedellinae observed to possess an ejacula-
tory suc, it is ‘reduced’ i.e. contained within the
copulatory bulb musculature. In the shadow of
this uncertainty, we chose to retain the name C.
spenceri to describe the worm with large Jamel-
lations. However, additional exceptional circum-
stances exist in that the crayfish host recorded by
Haswell (1893) cannot be reliably identified, and
the localities he provided are vague and wide
ranging. Therefore, we erect a neotype for C.
spenceri from what is in our opinion the host most
likely to have been collected by Haswell (1893)
(Le. Cherax destructor), from a locality approxi-
mately at the middle of the geographical range of

CRAYFISH ECTOSYMBIONTS

the crayfish (John Short, personal communica-
tion).

Our field collecting has not been exhaustive,
We predict that with further field examination of
crayfish hosts, the branchial chamber habitat of
Australian crayfish will reveal a greater diversity
of the Craspedellinae. Moreover, there is evi-
dence that the group also occurs outside Aus-
tralia. A undescnbed, putative species of
Craspedella with a cirrus clearly different to that
of any of the species described in the present
study, collected from the crayfish Cherax com-
munis Holthuis, 1949 from Papua New Guinea,
is held in the private collection of Dr Bons Joffe
of the Zoological Institute, St. Petersburg, Russia
(В, Joffe, personal communication),

ACKNOWLEDGEMENTS

The contribution of KBS to this study forms
part of a PhD within the Department of Parasitol-
ogy and Anatomical Sciences, the University of
Queensland.

First, we wish to thank the Australian Biologi-
cal Resources Study for a grant (87/5909) to
LRGC which funded the study, We thank mu-
seum staff of the state museums of New South
Wales, Victoria and Westem Australia, who as-
sisted greatly in appraising collections and re-
cords of crayfish to assist in locating suitable
collecting sites, most notably Dr Penny Baerents,
Australian Museum, Dr Gary Poore, Museum of
Victoria and Dr Gary Morgan, Western Austra-
lian Museum, We are indebted to Dr Don Horn-
ing from the McLeay Museum, University of
Sydney for his attempts to locate specimens or
data pertinent to Haswell's collections. Dr Pierre
Horwitz (Curtin University) and Dr Brenden
Knott (University of WA) provided valuable as-
sistance with WA crayfish localities and habitats
and Dr Louis Evans (Edith Cowan University ))
provided data of Western Australian tem-
nocephalans. Mr Wolfgang Zeidler, South Aus-
tralian Museum and Dr Тап Beveridge, University
of Melbourne kindly provided data and worms
respectively. Mr John Short of the Crustacean
Section of the Queensland Museum (QM) was
extremely helpful in the identification of crayfish
and with literature on the hosts. Furthermore, we
would like to thank Christine Lee and Zeinah
Khalil (QM) for their intelligent and dedicated
attention to specimen preparation in the labora-
tory. Dr Boris Joffe of the Zoological Institute, St.
Petersburg, Russia, enlightened us to the value of
de Faure's preparations of the cirrus and provided

417

invaluable expert advice. We thank officers of the
National Parks and Forestry in all states for assis-
tance with locating crayfish habitats.

LITERATURE CITED

ALDERMAN, D.J], & POGLASE, J.L. 1988. Patho-
fens, parasiies and comevensals. Рр. 167-212. In
Holdich, С.М. & Lowery, R.S. (eds.) "Freshwater
Crayfish Biology Management and Exploitation’.
(Croom Helm Ltd.: London).

BAER, J.G. 1931. Etude monographique du groupe des
Temnocéphales. Bulletin biologique de la France
el de la Belgique 65; 1-57.

1953. Zoological results of the Dutch New Guinea
Expedition 1939 No. 4. Temnocéphales,
Zoologische Mededelingen Leiden 32: 119-140,

1961. Classe des Temnocéphales, In Р.Р. Grassé
(ed.) "Traité de Zoologie" (Paris Masson et Cic).
4: 213-241.

BRESSLAU, E. & REISINGER, E. 1933, Turbellarin.
Pp. 52-308, In Kuckenthal, W. & Krumbach, T.
(eds.) ‘Handbook de Zoologie". Bd. 2., Н.І. (Wal-
ter de Gruyter und Co,; Beriin-Leipzig.)

CANNON, 1986, ‘Turbellaria of the World - A guide
to Families and Genera’. (Queensland Museum:
Brisbane.) 136 pp.

CANNON, L.R.G, 1991, Temnocephalan symbioms of
the freshwater crayfish Cherax quadricariratus
om northern Australia. Hydrobiologia 227: 341-

7.

1993. New temnocephalans (Platyhelminthes):
ectosymbionts of freshwater crabs and shrimps.
Memoirs of the Queensland Museum 33: 17-40.

CANNON, L.R.G. & JENNINGS, J.B. 1987. Occur-
rence and nutritional] relationships of four
ectosymbiotes of the freshwater crayfishes
Cherax dispar Riek and Cherax punctatus Clark
(Crustacea: Decapoda) in Queensland. Australian
Journal of Marine and Freshwater Research 38:
419-427,

CANNON, L.R,G, & SEWELL, К.В, 1994, Symbionts
and biodiversity. Memoirs of the Queensland Mu-
seum 33: 36-40.

CANNON, L.R.G. & WATSON, N.A. (in press). The
postero-lateral glands of Temmocephala minor
Haswell, 1988 (Platyhelminthes, Tem-
nocephalida). Australian Journal of Zoology.

DALLWITZ, M.J. & PAINE, Т.А. 1986. User's guide
to the DELTA system: a general system for pro-
cessing taxonomic deseriptions. CSIRO, Division
of Entomology, Report 13: 1-106.

HASWELL, W.A, 1893. A monograph of the Tem-
nocephaleae. Proceedings of the Linnean Society
of New South Wales (Macleay Memorial Vol-
ume) pp. 94-152.

HICKMAN, У.У. 1967. Tasmanian Temnocephalidea.
Pupers and Proceedings af the Royal Society of
Tasmania 101: 227-250,

418

HYMAN, L.H. 1951. The Invertebrates. Il:
Platyhelminthes and Rhynchocoela. McGraw-
Hill Book Co. Inc. New York. 550 pp.

JENNINGS, J.B. 1988. Nutrition and respiration in
symbiotic Turbellaria. Pp. 1-13. In Ax, P., Ehlers,
U. & Sopott-Ehlers, B. "Free-living and Symbiotic
Plathelminthes'. (Gustav Fischer: Stuttgart).

JENNINGS, J.B., CANNON, L.R.G. & HICK, AJ.
1992. The nature and origin of the epidermal
scales of Notodactylus handschini - an unusual
temnocephalid turbellarian ectosymbiotic on
crayfish from northern Queensland. Biological
Bulletin 182: 117-128.

JONES, T.C. & LESTER, R.J.G. 1992. The life history
and biology of Diceratocephala boschmai
(Platyhelminthes; Temnocephala), an ectosymbi-
ont on the redclaw crayfish Cherax
quadricarinatus. Hydrobiologia 248: 193-199.

MATJASIC, J. 1990. Monography of the family
Scutariellidae (Turbellaria, Temnocephalidea),
Academia Scientiarum et Artium Slovenica Clas-
sis IV: Historia Naturalis 28: Znan-
stvenoraziskovalni center SAZU, Bioloski Institut
Jovana Hadzija 9 (Ljubljana: Slovenska
Academija Znanosti in Umetnosti), 167 pp.

MORRISSY, N.M. & CASSELLS, G. 1992, Spread of
the introduced yabbie Cherax albidus Clark, 1936
in Western Australia. Fisheries Research Report
Fisheries Department of Western Austraha 92:
1-27.

MEMOIRS OF THE QUEENSLAND MUSEUM

ROHDE, K. 1987a. Ultrastructural studies of epidermis,
sense receptors and sperm of Craspedella sp, and
Didymorchis sp. (Platyhelminthes, Rhabdocoela).
Zoologica Scripta 16: 289-295.

1987b. Ultrastructure of flame cells and pro-
tnephridial capillaries of Craspedella and
Didymorchis (Platyhelminthes, Rhabdocoela).
Zoomorphology 106: 346-351.

SEWELL, К.В. & CANNON, L.R.G. 1995. A scanning
electron microscope study of Craspedella sp.
from the branchial chamber of redclaw crayfish,
Cherax quadricarinatus, from Queensland, Aus-
tralia. Hydrobiologia 305: 151-158.

SEWELL, K.B. & WHITTINGTON, I.D. 1995. A light
microscope study of the attachment organs and
their role in locomotion of Craspedella sp.
(Platyhelminthes: Rhabdocoela: Tem-
nocephalidae), an ectosymbiont from the bran-
chial chamber of Cherax quadricarinatus
(Crustacea: Parastacidae) in Queensland, Aus-
tralia. Journal of Natural History. 29:1121-1141.

WATSON, N.A., ROHDE, K. & SEWELL, K.B. 1995.
(In press). Ultrastructure of spermiogenesis and
spermatozoa of Decadidymus gulosus, Tem-
nocephala dendyi, T. minar, Craspedella sp., C.
spenceri and Diceratocephala boschmai
(Platyhelminthes, Temnocephalida, Tem-
nocephalidae), with emphasis on the intercentrio-
lar body and zone of differentiation. Invertebrate
Reproduction and Development.27:131-143.

A PERMINERALISED CUPULATE FRUCTIFICATION FROM QUEENSLAND
H. TREVOR CLIFFORD

Clifford.H.T. 1995 12 01: A permineralized cupulate fructification from Queensland. Mem-
oirs of the Queensland Museum 38(2):419-427. Brisbane ISSN 0079-8835,

Cupules containing 2-8 ovules are described as Dawesia cupulata gen. et sp. nov., a taxon
of uncertain affinities. The cupules and single dispersed ovules are embedded in per-
mineralised peat boulders of Jurassic age which occur surficially at Miles but are incorporated
in sediments of the Kumbarilla Beds at Chinchilla.[ ] Jurassic, fructification, Dawesia

cupulata.

H. Trevor Clifford, Queensland Museum, PO Box 3300, South Brisbane, Queensland, 4101,

Australia; 1 August 1995,

Permineralised peat known to the lapidiarists of
southeastern Queensland as "forest-floor" is rich
in plant material amongst which, leaves, stems
and ovules are conspicuous. The preservation of
tissues in the peat is poor, except for that of the
wood and the ovules, the cell structure of whose
integuments is discernible. Abundant im-
pressions of a Cladophlebis sp. are also present

on some surfaces. Although most ovules are sol-
itary they sometimes occur in clusters which are
enclosed in cupules (Figs 1,2). The earliest
known collection of a cupule is that of E.O Marks,
who early this century, found a specimen in the
Chinchilla district. A photograph of the cupule,
labelled "Gymnospermous seed capsule" has
been published by Hill et al. (1966).

FIG.1. Dawesia cupulata gen. et sp. nov., holotype, QMF32157, rosette of ovules, exposed on the surface of a
boulder of "forest floor"; collected at Miles: oc, ovule cavity; cw, cupule wall; int, integument.

420

FIG.2, Dawesia cupulata gen. et sp. nov. A-B, Cupules
of paratypes exposed on the surfaces of two "forest
floor" boulders collected from along Rocky Creek
near Chinchilla. A, QMF32185. B, QMF32184. Oc,
ovule cavity; int, integument; cw, cupule wall; cb,
cupule base; o, ovule.

ABBREVIATIONS USED: QM, Queensland Mu-
seum; GSQ, Queensland Geological Survey.

STRATIGRAPHY

The source of most "forest floor" specimens is
uncertain because all have been collected as sur-
ficial boulders from hillsides or stream beds.
However, at least some of those boulders which
show little evidence of abrasion and have been
collected from along the bed of Rocky Creek,
Chinchilla almost certainly derive from a lens of
"forest floor", incorporated in a single sandstone
bed which is exposed in the low cliffs forming the
right-bank of the Creek (26'34'S, 150'36' E)
100-200m upstream from the collecting site.

The strata which are exposed in the cliffs are
members of the Kumbarilla Beds a series of fine
to coarse grained sediments and thin bands of
coal, for the most part, laid down in stream,
deltaic and lake environments (Reiser, 1971).

MEMOIRS OF THE QUEENSLAND MUSEUM

Lenses of permineralized peat are also present.
The Beds are about 600m thick and dip gently to
the south. They range in age from Late Middle
Jurassic to earliest Cretaceous (Day et al., 1983)
or to Upper Cretaceous (Exon, 1976).

The only in situ "forest floor" encountered in
this study was a lens about 2m long and about
15cm thick which being incorporated in a sand-
stone bed that occurs in the upper portion of the
Kumbarilla Beds is close to the Jurassic-Creta-
ceous boundry. None-the-less the Cladophlebis
leaf impressions associated with the cupules de-
rive from a typically Jurassic taxon (J. Rigby,
pers. comm.) thereby confirming the pre-Creta-
ceous age of the "forest floor". As the ovules in
the permineralized peat are not compressed it
must have been silicified before being deeply
buried. The source of silification is not known but
from an examination of thin sections of "forest
floor" it is evident that at least two phases of
silicification have occurred (A. Cook, pers.
comm. ).

FIG.3. А-В, Integument of Dawesia cupulata gen. et sp.
nov. A, QMF31911, upper portion of integument of
central ovule in Fig.10C viewed in reflected light. B,
QMF32160, portion of an integument viewed in trans-
mitted light. oz, outer zone; iz, inner zone; oc, ovule
cavity.

PERMINERALISED CUPULATE FRUCTIFICATION

421

FIG. 4. Dawesia cupulata gen. et sp. nov. A-D, QMF32161, transverse sections of integument, viewed in transmitted
light. A, Integument and ovule cavity. B, Inner zone of integument and ovule cavity. C, Isodiametric cells of
integument. D, elongated cells of integument close to ovule cavity: m, investing matrix: oz, outer zone; iz, inner
zone; oc, ovule cavity filled with chalcedony; ec, elongated cells; ic, isodiametric cells.

SYSTEMATIC PALAEOBOTANY

Division INCERTAE SEDIS
Form Order DAWESIALES ord. nov.
Form Family DAWESIACEAE fam. nov.
Dawesia gen. nov.

DIAGNOSIS
As for the only presently known species.

TYPE SPECIES
Dawesia cupulata sp. nov.

DIAGNOSIS

Ovuliferous organs consisting of cupules, each
bearing 2-8 turbinate ovules on its inner surface,
micropyle not extended and the integument mem-
branous throughout, lacking vascular tissue and

resin bodies; cupules pedicellate and possibly
occuring in pairs.

ETYMOLOGY
For Graham Wallace Dawes, the collector of
QMF32157. Latin cupulata, like a cup.

Dawesia cupulata sp.nov.
Figs 1-10

DIAGNOSIS

Cupule hemispherical, c. 16mm in diameter, the
outer surface ribbed; ovules turbinate, 8-9mm
long and 7mm in diameter.

MATERIAL EXAMINED
HOLOTYPE: QMF32157, Miles, Queensland 26?35'S,
150°16'Е.

FIG,S, Dawesia cupulata gen. et sp. pow, A, Recanstruc-
tion of a pair of cupules. B, Diagrammatic longitudinal
section of cupule pair illustrated in Fig 5A. Though
illustrated as occuring in pairs the cupules may be
solitary. C, cupule; cw, cupule wall; o, ovule; pe.ped-
icel; pd, peduncle; or, ornamental rib.

PARATYPES: QMF32156, QMF31911, GSQF!41;
Chinchilla, Queensland 26°134'S,) 50°36" E.

DESCRIPTION

Ovule morphology. Mature ovules, as
recognised by their having been shed from the
cupule, are turbinate and 8-9mm long, with max-
imum width c.7mm one third of the distance from
the broad chalazal to the narrow micropylar end.
The radial symmetry of the ovules may be mod-
ified by compression resulting from contact with
adjacent ovules. The ovules arise from the basal
region of the inner surface of the cupule and they
lack micropylar beaks.

The integument is unvascularised and has no
internal glands. It is about Imm thick and is
usually differentiated into two zones which are
separated by an irregular boundary. The outer and
usually narrower of the two zones, when present,
is light- to pale-brown whereas the inner is brown
to dark-brown. The zones are evident in both
transverse (Fig. 10A-B) and longitudinal (Fig.
10C) sections and are clearly discernable in both
transmitted (Fig. 4A) and reflected light (Fig.

MEMOIRS OF THE QUEENSLAND MUSEUM

10A-C). Although the boundary between the
zones is quite definite it has no regular pattern and
is not associated with any constant differences in
cell-type, It is apparently an artifact that arose
during the process of mineralization. Support for
this interpretation of the origin of the zonation is
afforded by the inconstancy of the widths of the
two zones and the complete absence from some
integuments, ог parts thereof, of the pale outer
zone (Fig. 3B).

The cells of the integument are either straight-
walled and isodiametric with diameters of 25-
35um (Fig. 4C) or are elongated with slightly
curved walls (Fig. 4B,D). The elongated cells,
which may be up to three times longer than broad
with diameters similar to those of the isodiametric
cells, oceur towards the inner side or the basc of
the integument. The long axes of the elongated
cells have no consistent orientation but are mostly
disposed obliquely or at right angles to the long
axis of the ovule, The cells lining thé micropylar
canal do not differ markedly from the adjacent
cells of the integument.

The shapes of the epidermal cells of the integ-
ument are unclear and no stomates have been
observed. Furthermore, no megaspores, mega-
spore walls or gametophytes have been observed.
The ovule cavities are filled completely with
chalcedony.

Cupules, The ovules are borne in pedicellate
cupules. The pedicels are possibly paired and are
attached laterally close to the bases of the cupules
(Figs 5A-B). As seen in vertical section the walls
of the cupule completely embrace the ovules and
are thinner on their sides than bases. Each cupule
contains up to eight avules which arise near to the
base fram its inner surface (Fig. 5B), A fully
mature 6-8 ovuled cupule has a diameter of about
16mm. The upper surface of the cupule is ribbed
(Figs 5A,10A,B).

The epidermal cells of the outer surface of the
cupule illustrated tn Fig. 2B are of two types. The
majority are square to rectangular in outline with
slightly curved anticlinal walls. As seen from
above the surfaces of the epidermal cells are
pitted or finely grooved. The remaining cells are
arranged in nngs of 5-6 and disposed around
apertures which presumably lead to sunken
stomates (Fig. 6).

The above description of the cupule morphol-
ogy derives from three sources.

l. Partially exposed ovules, The beautiful ro-
sette of ovules which first directed attention to
Dawesia cupulata is exposed on the surface of a
boulder (Fig. 1). The cupule which is here re-

PERMINERALISED CUPULATE FRUCTIFICATION

423

FIG. 6. Dawesia cupulata gen, et sp. nov. Epidermal cells as revealed in an electron micrograph of latex mould
taken from the central area of the outer surface of the cupule illustrated in Fig. 2B: sp, stomatal pore.

vealed in transverse section is represented only
by fragments of its wall. There is no clear indica-
tion of a central axis at the centre of the rosette
suggesting the cupule is not peltate, as 1s con-
firmed by longitudinal sections. The two other
clusters of ovules (Fig. 2) are revealed through
the eroded bases of their cupules whose slight
assymmetry is ascribed to the lateral attachment
of their pedicels.

2. Parallel sections. The outlines of a partial
cupule (Figs 7,8) and its enclosed ovules are
exposed on the four surfaces of two adjacent
slabs, cut from the same boulder as that with the
rosette on its surface.

Using à common margin of the slabs as a refer-
ence line and allowing both for the differences in
slab thickness and the width of the saw-cut a
section of the cupule was constructed as it would
appear at right angles to the exposed surfaces
(Fig. 9). The dispositions of the ovules in this
reconstructed section confirm they were borne in
a cupule. The lack of continuity of the right hand
wall ofthe cupule and the separation of the ovules
suggests that considerable decomposition had
taken place before preservation was achieved,

3. Single sections. Three additional cupule out-
lines are available, Using ovule shape as the
determining criterion two of the sections are re-
garded as transverse and the third as longitudinal.

(a) Transverse sections. Because the outlines of
the ovules are circular to elliptical with widths
near to the maximum observed it is assumed both
cupules have been cut across transversely and
near to their bases (Figs 1OA-B,11A-B).

The cupule walls are thin (Figs 10B,11B) and
their outer surfaces are ornamented with small
projections which are interpreteted as superficial
ridges seen in section.

(b) Longitudinal section. Because the outlines
of its ovules are turbinate it is assumed the cupule
has been cut longitudinally an opinion supported
by the outline of the central of the three ovules
whose integument is uniformly wide except
where interrupted by the micropylar canal (Figs
10C,I 1C). As expected where ovules are ar-
ranged in a circle the micropylar canals of the
adjacent ovules are not visible on the exposed
surface but can be seen through their integuments
by focusing down into the almost transparent
investing matrix.

424

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 7 Dawesia cupulata gen. et sp. nov. A-D, sections of an incomplete cupule of as exposed on the four planes
resulting from the cutting of two parallel slabs from the boulder that includes the holotype. A-B, QMF32159a.
C-D, QMF32159b: pe, pedicel; int, integument; oc, ovule cavity; cw, cupule wall.

The cupule wall completely embraces the
ovules and is quite thin except at the base where
it is attached to the pedicel. Unfortunately no
tissues are preserved in this area but it appears
that at least part of the apparent thickness of the
cupule base may result from separation of the

outer epidermis from the surface of the cupule. In
the right-hand side of this space a small rock
fragment has been incorporated.

In this plane of section no surface ornament is

present as would follow if the projections seen in
the tranverse sections are those of superficial

PERMINERALISED CUPULATE FRUCTIFICATION

FIG. 8. A-D. As for Fig. 7. A, Pedicel of cupule. B-D,
transverse sections of cupule.

ornamental ridges radiating from the base of the
cupule.

Neither of the elliptical structures close to the
apex of the ovule on the right-hand side of the
cupule is regarded as associated with that struc-
ture.

The two cupules of which one is exposed in
longitudinal section (Fig. 10C) and the other in
transverse section (Fig. 10B) occur in close prox-
imity on the facing surfaces produced by a single
saw-cut. Though no organic connection between
the two cupules has been established it has been
assumed their close association indicates that the
cupules occur in pairs. The manner of arrange-
ment of these pairs into infructescences is not
known.

5mm

FIG. 9. Dawesia cupulata gen. et sp. nov., longitudinal
section of a cupule reconstructed from the four tran-
sects marked on the sections illustrated in Fig. 8.

425

DISCUSSION

The cupules of Dawesia cupulata are of partic-
ular interest for being permineralised and un-
crushed their gross morphology can be
reconstructed with reasonable certainty. None-
the-less, due to their poor state of preservation
and lack of associated foliage the taxonomic af-
finities of the genus are unclear. The protection
of ovules within a cupule is a feature that has
evolved on several occasions and so is of little
value for defining higher level taxa. Therefore, in

FIG. 10. Dawesia cupulata gen. et sp. nov. A-C, sec-
tions exposed on the surfaces of slabs cut from the
boulder that includes the holotype. A, QMF32186,
transverse section of a 2-ovulate cupule. B,
QMF32156, transverse section of a >3-ovulate cu-
pule.C, QMF31911, longitudinal section of >3-ovu-
late cupule with the central ovule cut in the median

376

5mm

FIG. 11, Dawesia cupulara gen. et sp. nov. As for Fig.
10. cw, cupule wall; int, integument; mc, micropylar
canal; ос, ovule cavity; or, ornamental rib; pe,pedicel,

order to determine the taxonomic affinities of
Dawesia particular attention must be paid to the
structure of its ovules, In particular the vas-
cularisation, presence of sclerenchymatous bands
and the occurence of secretory tissue are import-
ant diagnostic features.

Although there is some differentiation of cells
within the integument the cells are all parenchy-
matous and secretory tissue is lacking. It is as-
sumed that the integumentary cells of Dawesia
cupulata are primarilly thin walled and that this
condition does not result from the destruction,
during fossilisation, of otherwise thick cell walls
thereby leaving only their middle lamellae. The
destruction of such thick walled cells has been
reported by Srivistava (1946) to have occurred in
ihe integuments of some Carnoconites (Form
Genus for the ovules of Pentoxylon) specimens

MEMOIRS OF THE QUEENSLAND MUSEUM

he investigated. The single integument of Dawe-
sia ovules and its lack of differentiation into threc
layers, an outer and inner composed of paren-
chyma with a fibrous middle layer, reduces con-
siderably the taxa to which the genus may be
related.

Thus the integument being single rather than
double disqualifies Dawesia from membership of
the Cordaitales, Gnetales and Taxales (Coulter &
Chamberlain, 1917) as well as the primitive an-
giosperms (Denffer et al., 1980). Furthermore,
lack of differentiation of the integument excludes
Dawesia from membership of the Pinophyta,
Cycadophyta, Ginkgophyta, Benneititales
(Coulter & Chamberlain, 1917), Caytoniales and
Pentoxylales (Sporne 1974).

Attention is therefore drawn to the Mesozaic
"seed ferns” , Peltaspermales, Corystospermales
and Petriellales as possible relatives of Dawesia
forall Ihese Orders have undifferentiated, uniteg-
mic ovules. Furthermore, ovulate fructifications
of both Peltaspermales (Holmes, 1982) and
Corystospermales (Holmes & Ash, 1979: Holmes
1932 & 1987; Playford et al., 1982: Retallack.
[980; Shirley, 1898) have been reported from
Australia. All reports are from Triassic rocks
which are much older than the Kumbarilla Beds
at Chinchilla. Solely on the basis of its ovule
structure Dawesia has previously been assigned
tentatively to the Peltaspermales by Clifford and
Carney (1994). However,the cupulate infructesc-
ence of Dawesia makes it unlikely that the genus
is à pellasperm, the ovules of which are borne on
the lower surface of umbrella-like discs. The
Corystospermales are also unlikely to be relatives
of Dawesia because although their ovules are
borne in cupules these are uniovulate and the
apices of the ovules are curved and bifid.

The cupules of Perriella like those of Dawesia
are embedded in permineralised peat but are
much better preserved notwithstanding their
Triassic age. Although Petriella has multiovulate
cupules these differ markedly from those of
Dawesia in being much smaller, and the ovules
are triangular instead of circular in transyerse
section with the integument forming а distinct
tube about the micropyle (Taylor et al., 1994),
Accordingly, the two genera cannot be regarded
as closely related on the basis of their cupüle and
ovule morphologies.

Therefore, until further evidence is available, it
is appropriate that Dawesia be placed in a unique
Order of uncertain affinity.

PERMINERALISED CUPULATE FRUCTIFICATION

ACKNOWLEDGEMENTS

Ithank Department of Earth Sciences, The Uni-
versity of Queensland for preparing sections; the
Australian Museum for hospitality; Dora Aitken
for the scanning electron micrographs; Mary
Dettmann for the photographs presented in Fig.4;
Mary Wade for several kindnesses during the
preparation of the paper and Natalie Camilleri for
unstinting assistance in preparing the diagrams;
Graham Davies and Grace Lithgow provided
specimens and guidance in the field; Mike Pole
read and offered useful comments upon the
manuscript.

LITERATURE CITED

CLIFFORD, H.T. & CARNEY, L.G. 1994. A non-de-
structive technique for determining the shape in
situ of permineralised seeds, Memoirs of the
Queensland Museum 35(1):23-25.

COULTER, J.M. & CHAMBERLAIN, C.J. 1917. Mor-
phology of Gymnospermns. (University of Chi-
cago Press:Chicago). 1-xi, 1-466.

DAY, R.W., WHITAKER, W.G., MURRAY, C.G..
WILSON, LH. & GRIMES, K.G. 1983. Queens-
land Geology. Geological Survey of Queensland,
Publication 383.

DENFFER, D. VON, SCHUMACHER, W,
MAGDEFRAU, K, & EHRENDORFER, F.
1980. Strasburger's Textbook of Botany. trans. P.
Bell & D. Coombe. (Longmans Group Limited:
London). i-xvi, 1-877.

EXON, N.F. 1976. Geology of the Surat Basin in
Queensland. Australian Bureau of Mineral Re-
sources Geology and Geophysics. Bulletin 166.

HILL, D., PLAYFORD, G., & WOODS, J.T. 1966.
Jurassic Fossils of Queensland. Queensland Pal-
acontographical Society, Brisbane.

427

HOLMES, W.B.K.1982. The Middle Triassic flora
from Benolong, near Dubbo, central-western New
South Wales. Alcheringa 6:1-33.

1987. New corystospermum ovulate fructifications
from the Middle Triassic of eastern Australia.
Alcheringa 11:165-173.

HOLMES, W.B.K. & ASH, S.R. 1979. An Early
Triassic megafossil flora from Lome Basin, New
South Wales. Proceedings of the Linnean Society
of New South Wales 103:47-70.

PLAYFORD, G., RIGBY, ЈЕ. & ARCHIBALD, D.C.
1982. A Middle Triassic Flora from the
Moolayembar Formation, Bowen Basin, Queens-
land. Geologial Survey of Queensland Publication
380.

REISER,R.F. 1971. 1-250,000 Geological Series - Ex-
planatory Notes. Chinchilla, Queensland. Sheet
SG/56-9 International Index. (Bureau Mineral Re-
sources, Geology and Geophysics: Canberra).

RETALLACK, G. 1980, Late Carboniferous to Middle
Triassic megafossil floras from the Sydney Basin
, in "A Guide to the Sydney Basin" (C.Herbert and
R.Helby eds). Geological Survey of New South
Wales Bulletin 26:384-430.

SHIRLEY, J. 1898. Additions to the fossil flora of
Queensland. Geological Survey of Queensland,
Bulletin 7 (Publication 128):9-25.

SPORNE, K.R. 1974. The Morpholgy of the Gymno-
sperms. The Structure and Evolution of Primitive
Seed-plants, (Hutchinson University Library:
London).

SRIVASTAVA, В.В. 1946. Silicified Plant-remains
from the Rajmahal Series of India. Proceedings of
the National Academy of Sciences 15:185-211,10
plates.

TAYLOR, T.N., DELFUEGO, G.M. & TAYLOR,
E.M. 1994. Permineralised seed fern cupules from
the Triassic of Antarctica: implications for cupule
and carpel evolution. American Journal of Botany
81(6): 666-677.

MICROCHIROPTERAN BAT MORTALITY IN A
HARP TRAP DUE TO GREEN TREE ANT
DECOPHYLLASMARAGDINA ATTACK. Memoirs of the
Oweensland Museum 38/2); 428. 1995. . Microchirpoteran bat
mortality in harp traps has been previously reported in Viclo-
ria (Schulz & Meggs, 1986; Lumsden, 1989; Wallis &
Lumsden, 1993) and the Northern. Тегпїпгү (Undervoud,
19941. In Victorian examples, bats were atlacked and partially
caten by Bush Rats Кома fuscipes, while im the cse Fram the
Northern Territory, mortality was due to Green Tree Ant
Oecophylla smaragdina attack. This note re another case
of О, smarügdine attack оп а harp trap which resulted im the
mortality of 23 Little Bent-wing Bals, Miniopterus australis.

During a fauna survey of coastal wetlands in Trinity Inlet,
Cairns, a harp trap was set on a walking track sirared on the
boundary of a dune ridge community (Meialeauca leucadendra
overstorey 1e Am, with mixed Асасие crisicargs, Alyria
траса, Exocarpus davifolia, Terminiin muelleri, Cuponieop-
348 аплот ех, Pandanus sp mid storey) and a Ceriaps
fogal dominated mangrove community, The trap was placed
between (wo birge M. fewcademdra and, as the author Was
aware of Ihe previous case of Green Tree am attack, care was
laken nol to rest the trap against, or tie Ihe skle arx hor-ropes
pedo the surroum&ng vegetation The trap was set al dusk, and
afer a penod wf seonlighning and ultrasonic bar detection, the
trap was ehecked for the last lime а! 2230 hours, No Bats were
present in the trap al the time of the fast check, despite a large
mater of Individuals being observed Flying overhead and
along the track. Inaction, 50 Green Tree Ants Were observed
en the Ira.

‘The rrap was checked ar approximately 0600 hours the
next Maoming and 23 trapped M. australis were discoverei
dead m the trap. An extremely large number of О, smarageing
(approximate estimate 1-200060 individuals) were present
on We map, partsculimdy within the folds and spaces of the
canvas capture bag and plastic dividing flaps. All bats were
covered with large nuinbers of ants, many of which were also
dead. In remnovme the canvas holding bay and then the bats,
the author was attacked and bitten, and fumes excreted by the
unts cuused gagging and retching, Access to the trap by the
Geeen Tree Ams was trom the ground. The trap had heen set
in similar and slightly more dense vegetation on two previous
mecastons Without attack,

The Green Tree Ant Qecophyila smaragdima(Formicklae)
15 a member of a group known as weaver ants due to their
ability to construct arboreal community nests of woven leaves.
‘This odd world genus is widespread throughout tropical Asiz
Africa and Australia, and forms large decentralised colonies
of upto 500,000 individuals (workers, larvae workers, queens
and queen larvae) representing many hundreds of nests in irees
covering areas over I000m* (Holldobler & Wilson, 1990).
Green Tree Ants are renowned for thelr highly evolved com-
municalion and co-operative social systems which include
alarm responses, defence and food collection systems and
recruitment and exploration strategies, all developed 10 can-
trol and maintain their large home range (Hólldobler, 1933).
Short and long-distance recruitment is achieved hy а combi-
nation of rectal gland odour trails, regurgitation of liquid Gop
contents, sternal gland secretions, mandibular alacm phere-
mones, body jerking displays und tactile stimulation. The
composition and identity of these chemical signals are colony-
specific (Holldobler & Wilson, 1990),

From the knowledge of Green Tree Ant biology und
inierochiopreran bat feeding parterns, à scenario forthe canse
of the attack and subsequent death of the bats in the harp trap
can be hypothesised, Recruitment strategies allow Green Tree
Ani nesimaies іо explore and occupy а new space in à very
short space of lime. Worker ants exploring this territory:
penndically touch their abdomen Tips to The new substrate

MEMOIRS OF THE QUEENSLAND MUSEUM

leaving draps of brown Muid thal act as colony-sgecific trail
markers, Previous studies involving the placemert of pot-
plants tio territorial areas indicated that werkers actively
enter and search newly opened terrain (HOlldobler & Wilsan,
1994). It is suggested that the placement of the harp trap onta
ibe track represented. new temitory thal was rapidly explored
and occupied during the course of the evening, before any bats
were trapped.

Microchiropteran bats generally demonstrate bimodal
feeding activity patterns, with post-dusk and pre-dawn activ-
ity peaks (Taylor et #1,,1987). 1и the above case, the har irap
failed to capture any bats during the first peoa of feeding,
despite a high level of observed activity. Therefore the bats
subsequently trapped were possibly captured during the pre-
dawn feeding period. Between the final trap checking and
dawn, large numbers Green Tree Ants explored and occupied
the harp trap, utilising the trap-Jegs us am access рш. It as
hypenhesised thar when the bars started stoking and Falling
into the hürp-trap holding bag, this disturbance caused rapid
and aggressive defence responses in the ams. Green Tree Ants
йге highly responsive to substrate vibratinns (C. Hill, pers.
come.), The bats were probably killed hy а combination cf
ant-bites and secretions from the Green. Tree Ams. Under-
wood (1994) incorrectly reported that the bats had been 'ap-
paresily killed by the repented stings inflicted һу the ants”.
Cireen Tree Ants do not possess abdominal ‘stings’ bus rather
use their mandibles to bite and sprays of formic acid from dheir
glands in defensive or attacking responses.

When harp trapping in tropical environments where Green
Tree Ants are present, Itis suggested that a number of pescau-
tyans are laken іо oyercome the possibility of ants accessing
and eccupying the trap, including: minimising the contact of
the {гир on surroundiig Vegetation, no tang the stabilising
Езу торе io surrounding vegetation, using surface insect
repellent on the irap legs or portiones of the ігар that are
potential access points for rhe qns and placing the tray kegs in
small containers of water.

Acknowledgements

Thanks to Dr Chris Hill (Zoology Department, James
Cook University), Trapping was conducted under Ihe terms
and conditions of Queensiand Department of Environment
and Heritage permit number T-0) 182.

Literulure Cited

Ноот. B. 1983. Tertitoeval behaviour in the Green Trec
Ant (Oecophylli тагои). Biowopica 143k 241-
25

Hölldobler, B. & Wilson, E.O. 1590. The Ants. Belknap Press,
Springer-Verlag, Berlin,

Lomsden, L. F. 1989. Predation by a Bush Ка! Rotts мау
of hats coptive in a harp trap. Macroderma 5* 41-42.

Schulz, M. & Megs, R. 1986, Predation on harp trap captives:
whodunit? Macroderma 2: 67-68.

Taylor, К), O'Neill, M.G. & Reardon, T, 1987, Tasmanian
buts: identification, distribution and natural history.
Papers and proceedings of the Royal Society of Tasma-
nia 121: 109-119,

Underwood, G, 1994, Note on the death of bats trapped in 2
harp trap in tropical Australia. Australasian Bar Society
Newsletter 3: 16

Wallis, R. & Lumsden, L.F: 1993 Another predation of hats
captive in a harp trap, Australasian Bal Society News-
letter 1:11

А. S. Кип, Australian Centre for Tropical! Freshwater Re-

search, James Cook University, Townsville, Queensland

4811, Australia; 10 June 1995.

GASTROPODS FROM THE UKALUNDA BEDS AND DOUGLAS CREEK, EARLY
DEVONIAN, NORTH QUEENSLAND

ALEX G. COOK

Cook, A. G, 1995 12 01: Gastropods from the Ukalunda Beds and Douglas Creek, Early
Devonian, north Queensland. Memoirs of the Queensland Museum. 38(2): 429-435, Bris-

bane, ISSN 0079-8835,

Four gastropods are recognised from the Ukalunda Beds and related sediments Irom Douglas
Creek, Early Devonina north Queensland. This endemic fauna comprises Tropidodiscus
foliatus sp. nov., Trochonema (Eunema) sp, Kitikamispira ukalundensis sp. nov., and
Homyzyga camilleriae gen. et sp. nov. Hornyzvga gen. nov. is erected for pal-
aeozygopleurids with coarse nbbing and a characteristic vertical subsutural surface on the
upper whorl face. [ ] Gastropods, Devonian, Emsian, Ukalunda Beds, Queensland.

Alex С. Cook, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101,

Australia; 1 August 1995.

Gastropods are subordinate components. of
many Devonian fossil assemblages in north
Queensland. Diverse, and highly endemic gastro-
pod faunas are known from the Middle Devonian
of the Broken River and Burdekin Provinces
(Heidecker, 1959; Cook, 1993b). This paper doc-
urnents the gastropod fauna collected from the
Ukalunda Beds and the siliciclastic sediments
associated with the Douglas Creek Limestone.
Brachiopod, coral, and conodont faunas have
been documented from the Ukalunda Beds (Hill
et al., 1967; Jell & Hill, 1969; Brock & Talent,
1993). Parfrey (1989) recorded a number of
brachiopods, a single taxon of bivalve and an
unidentified gastropod from outcrops of the
Ukalunda Beds near Mount Coolon. Studies of
the faunas of the Douglas Creek Limestone and
related siliciclastic units to the immediate south
have concentrated on the coralline, brachiopod
and conodont faunas (Hill. 1939; Jones, 1941;
Phillip & Pedder, 1967; Jell & Hill, 1970; Brock
& Talent, 1993). A detailed record of the previous
palaeontological work was provided by Brock &
Talent (1993), who also discussed evidence for
an Emsian age for both the Ukalunda beds and the
Douglas Creek Limestone and related siltstones.

Recent stratigraphic mapping by Bennedick
(1993) in the Pyramid area informally docu-
mented some of the molluscan taxa, namely
Nuculana sp., Parralellodon sp., Paracyclas sp.,
Leptodesma sp., and gastropod indet. Study of
Bennedick's material forms part of this work.

The four taxa described below are endemic to
north Queensland, but there are generic-level
links to faunas from New South Wales, northeast
Japan and other old world realm faunules,

LOCALITIES AND MATERIALS

Material was collected by the Bureau of Min-
eral Resources (now AGSO), Macquarie Univer-
sity, Jarrod Bennedick of James Cook University,
Townsville, and staff of the Queensland Mu-
seum. Details of BMR localities have been re-
corded by Brock and Talent (1989). James Cook
University localities are prefixed JCUL and are
detailed in the Appendix along with Queensland
Museum localities (QML). Material described
herein is registered with the Commonwealth Pal-
aeontological Collections (CPC), Queensland
Museum (QMF) and James Cook University
(JCUF).

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA
Class GASTROPODA
Family BELLEROPHONTIDAE McCoy, 1851
Subfamily TROPIDODISCINAE Knight, 1956
Tropidodiscus Meek & Worthen, 1866

TYPE SPECIES

Tropidodiscus curvilineatus (Conrad) from the
Lower Devonian of New York, by original des-
ignation.

Tropidodiscus foliatus sp. nov
Fig, IC-H

MATERIAL EXAMINED
HOLOTYPE: QMF32176 from QML1008.
PARATYPES: QMF32362-4 from QML1005. All mate-
rial from the Ukalunda Beds, Pyramid Station and
Mary Creek, Old Hidden Valley Station.

430

DIAGNOSIS
Carina well developed; growth lines coarse,
well-spaced, foliaceous.

DESCRIPTION

Medium sized, isostrophic, doubly phan-
eromphalous, up to 12mm wide and 16mm in
diameter, bearing a prominent dorsal carina
which bears a deep v-shaped sinus. Whorl profile
rounded from base of dorsal crest to a weak
shoulder, then more strongly rounded into the
umbilicus. Ornament consists of prominent,
widely spaced foliaceous coarse growth lines,
which are gently curved across the whorl profile
and strongly inflected adapically at the dorsal
crest, but this latter feature varies from early to
later parts of the final whorl. Umbilicus deep;
aperture expanded. The holotype is 5mm wide
and 11 тт in diameter.

REMARKS

Prominent dorsal angulation, and foliaceous
growth ornament suggest affinity to Temnodis-
cus, but that genus is disyunctly- coiled.
Tropidodiscus cultricarinatus Linsley, 1968 and
T. vesculilineatus Linsley, 1968 from the Middle
Devonian Anderdon Limestone, north America,
both have the dorsal angulation, but lack the
foliaceous growth ornament. The type species T.
curvilineatus (Conrad), (see Knight, 1941) from
the Lower Devonian of New York lacks the an-
gulation and the foliaceous growth lines.
Tropidodiscus centrifugalis (Chapman) of
Tassell (1977) from the lower Devonian of Vic-
toria, does not bear growth ornament as coarse as
in the Ukalunda species. Tropidodiscus (T.)
nakazatensis Kase & Nishida, 1986, from the
Middle Devonian of northeast Japan is close in
size and prominence of the carina, but the oma-
ment is finer and the umbilicus narrower in the
Japanese material,

ETYMOLOGY
Latin foliatus, leaved.

Family TROCHONEMATIDAE Zittel, 1895
Trochonema Salter, 1859
Trochonema (Eunema) Salter, 1859

TYPE SPECIES

Eunema strigillatum Salter from the Middle
Ordovician Black River Group, Quebec, Canada,

MEMOIRS OF THE QUEENSLAND MUSEUM

Trochonema (Eunema) sp.
Fig. 1A,B

MATERIAL EXAMINED
QMF33369-61 from QMLI005, Mary Creek, Old
Hidden Valley,

DESCRIPTION

Many-whorled, high-spired, gradate gastro-
pod, greater than 35mm high and 22mm wide
with an apical angle of approximately 35°. Suture
canaliculate in two of the specimens, particularly
in later whorls, with an angulation at its abaxial
margin. Whorl profile dominated by a pair of
widely spaced midwhorl carinae. A fourth carina
js present on the lower whorl profile, but it is only
known from the final whorl. Growth lines are
fine, numerous, opisthocline on the upper whorl
surface, a broad. weak sinus is present between
the two midwhorl carina. Base and aperture un-
known.

REMARKS

The canaliculate suture, the high-spired form
and the broad weak sinus ally the material to the
subgenus. T. (E.) strigillatum (Salter), from the
Ordovician Black River Group, as figured by
Knight (1941), has stronger growth lines, and is
disjunct in the final whorl. Widening of the canal
in the final whorl of this material may be precur-
sory to a disjunct whorl. The absence of material
defining the apertural and basal features preclude
a specific assignment for this material.

Family PALAEOTROCHIDAE Knight, 1956
Kitikamispira Kase & Nishida, 1988

TYPE SPECIES

Kitikamispira kaneoki, from the Middle
Devonian (Eifelian) Nakazato Formation,
Kitikami Mountains, northeast Japan, by original
designation.

DIAGNOSIS
See Kase & Nishida (1988).

REMARKS

Kase & Nishida (1988) remarked on the simi-
larities between Burdikinia Knight, 1937 and
Kirikamispira. Burdikinia burdekinensis
(Etheridge) is a substantially different form, hav-
ing only the surface spines and large size as
similarities. Kase & Nishida (1988) state that the
type species, К, kaneoki, possesses only one row
of spines on the upper whorl surface and four on

EARLY DEVONIAN GASTROPODS

FIG. 1. A-B, Trochonema (Eunema) sp. A, latex mold of QMF33359 x 1.8. B, latex mold of QMF33360 x1.8.
Note the widening of canal in last preserved whorl. С-Н, Tropidodiscus foliatus sp. nov. C, holotype QMF32176
apertural view , x 3. D, holotype QMF32176 side view x 2.8. E, latex mold of paratype QMF33364, apertural
view x 2.5. F, latex mold of paratype QMF33367, oblique view x2. G, latex mold of Paratype QMF33364, side
view x 3. H, latex mold of paratype QMF33362, side view x 2.6.

432 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. A-H, Kitikamispira ukalundensis sp. nov. A-C, holotype QMF33358. A, apertural view x 2.7. B, apical
view x 2.7. C, side view showing protoconch x 5. D, E, paratype QMF32179, side and apical views, x 2.8. F,
latex mold of QMF33371 side view x 6.5. G, latex mold of paratype QMF 33354, side view x 2.8. H, latex mold
of paratype QMF33372, basal view x 2.8.

EARLY DEVONIAN GASTROPODS

the base, but their illustrations show one on the
uppermost whorl surface, two on the midwhorl,
and up to three on the lower whorl surface. Ori-
уота Munier-Chalmas, 1876 is phan-
cromphalus, but has a similar, if finer and more
complex arrangement of spinose spiral cords.
Kase & Nishida (1988) placed the genus within
the Palaeotrochidae, apparently on the basis of
the thickened shell and the thickened panetel
region,

Kitikamispira ukalundensis sp. nov,
Fig. 2

1989 Gastropod indet. Parfrey - 20.

MATERIAL EXAMINED

HOLOTYPE: QMF33358, from "loc 76", Mary Creck
Old Hidden Valley, collected J. A, Talent.
PARATYPES; QMF32179,32180 from QML1008,
QMF33353-33357,QMF33371,33372 from
OMLI1O005. OTHER MATERIAL: GSQF13456, from
GSQL2765, JCUF12779 from JCUL810, CPC33678,
fram CL 172, tenatively assigned to this species.

DIAGNOSIS
Small, Kitikamispira with flattened whorl pro-
file between suture and first spiral row of spines.

DESCRIPTION

Small to medium, medium spired, turbiniform,
gastropod up to 19mm high and 19mm wide, with
an apical angle of c.50°. Whorl profile generally
rounded, but flattened in the uppermost whorl
surface. Whorl surface bears 5 rows of spirally
arranged spines, in some specimens upon a weak
thread. There is one row of spines at the edge of
the subsutural shelf, two rows at the midwhorl
periphery, and two on the lower whorl face.
spines are slighlty elongate along the spiral cords,
numerous, with approximately 14 per row per
whorl; some are directed slightly abapically. Su-
tures slightly impressed, situated just below the
midwhorl and the third row of spines, so that the
spines are exposed on the flattened subsutural
shoulder. The protoconch is preserved on the
holotype, it consists of two unornamented volu-
tions, the first vertically coiled and the second
dextral and horizontally coiled; or “deviated
paucispiral" of Knight et al. (1960). Aperture is
rounded; the shell is moderately thick for its size,
and appears thickened in the parietal region but
the apertural region is broken in this material.
Growth lines fine, strongly prosocline upon the
upper whorl surface, nearly orthocline on the

413

midwhorl, and where preserved appear to he pro-
socline on the lower whorl surface.

REMARKS

This species is smaller than the type, and has a
higher spire, It retains a similar arrangment of the
spiral nodes, but the position of the uppermost
row of spines is more adapertural and the whorl
profile from the suture to the uppermost row of
spines is more flattened. The parietal callus, pres-
ent in the type, is missing in this material. The
external mould from Douglas Creek is a very
poorly preserved specimen. probably conspecific
with the Ukalunda material. I have examined
material mentioned by Parfrey (1989) and
Bennedick (1993) which is certainly conspecific.

ETYMOLOGY
For the Ukalunda region.

Family PALAEOZYGOPLUERIDAE
Horny, 1955
Hornyzyga gen. nov

TYPE SPECIES
Hornyzyga camilleriae sp. nov.

DIAGNOSIS

Vertical unornamented surface below the su-
lure, generally rounded midwhborl profile bearing
coarse collabral ribs.

ETYMOLOGY
For Radvan Horny.

REMARKS

Palaeozygopleura Horny seems to encompass
forms with finer collabral ribs, and no shoulder,
such as Loxonema roemeri of Whidborne (1892),
and Palaeozygopleura muoni Tassell 1982.
Linsley (1968) seems to adopta wider view of the
genus by the inclusion of the slightly coarser
ribbed Palaeezygopleura sibleyense Linsley
from the Middle Devonian Anderdon Limestone.
Further latitude given to the importance of size
and rib coarseness within Palaeozygopleura may
accomodate Loxonema scalariaeforme
Holzapfel of Whidborne, 1892 and Loxonema
altacostatum Tassell, 1982 which is anomalous
within Loxonema. Devonozyga Horny. 1955 pos-
sesses a distinct shoulder and thicker ribs.
Homyzyga is separated from both Pal-
aeozygopleura and Devonozyga by the vertical
surface below the suture, and restriction of the
ornament to the wide midwhorl,

434

Fig. 3 A-C. Hornyzyga camilleriae gen. et sp. nov. A,
Latex mold of holotype QMF33385, apertural view x
2.2. B, Latex mold of Paratype QMF33369, side view
x2. C, Latex mold of holotype QMF33385, x 6. Note
the flattened subsutural areas.

Familial placement may be considered prob-
lematic without protoconch material, following
Homy (1955) and Knight et al. (1960), but seems
reasonable given the gross morphological simi-
larities.

MEMOIRS OF THE QUEENSLAND MUSEUM

Hornyzyga camilleriae sp. nov,
Fig. 3 A-C

DIAGNOSIS
As for genus.

MATERIAL EXAMINED

HOLOTYPE: QMF33385 from QML1005.
PARATYPES: QMF33369-70 from L1006, QMF33386
from QML1005

DESCRIPTION

Medium sized, high-spired, many-whorled, up
to 30mm high and 11mm wide, with an apical
angle of c.15*. Sutures adpressed, sutural slope
shallow. Midwhorl surface 1s dominated by thick
collabral ribs, approximately 12 per whorl, which
are variably opisthocline. Below the suture is a
vertical and unornamented surface, but the whorl
profile is convex where it bears the thick ribs.
There is a vertical unornamented area below
these riblets upon the final whorl, and below this
the lower whorl profile is smooth and rounded.
The suture is situated just below the midwhorl
ribs. Aperture is apparently rounded, Growth
lines unknown, Protoconch unknown. The holo-
type is 30mm high and 10mm wide, but lacks a
protoconch. Paratype QMF33369 is crushed, and
QMF33370 is represented by two late whorls.

REMARKS

Loxonema altacostatum Tassell, 1982 from the
Early Devonian of Taemas, New South Wales,
lacks the subsutural surface of this taxon, giving
the Taemas species a more rounded and even
whorl profile. Tassell’s species is better placed
within Palaeozygopleura. Other ribbed
Loxonema include L. roemeri Kayser of
Whidborne (1892) from the Devonian of Devon,
which also lacks the subsutural surface, and tends
to has a more orthocline to prosocline arrange-
ment of closely spaced riblets. L. scalariaeformie
Holzapfel of Whidborne (1892) has more im-
pressed sutures and also lacks the subsutural sur-
face.

ETYMOLOGY
For Natalie Camilleri.

ACKNOWLEDGEMENTS

Ithank Peter Jell & Chris Tassell for comments
on the manuscript, Bob Henderson, Des Strusz
and Susan Parfrey for loan of materials, and Nat-
alie Camilleri for field assistance.

EARLY DEVONIAN GASTROPODS

LITERATURE CITED

BENNEDICK, J. L. 1993. Geology and stratigraphy of
the Pyramid area, north-west Drummond Basin,
north Queensland. (Unpublished B.Sc.(hons.)
thesis, James Cook University of Queensland:
Townsville). 85pp.

BROCK, С. A. & TALENT, J. А, 1993. Emsian (Early
Devonian) brachiopods from the Ukalunda Beds
and Douglas Creek, north Queensland. Memoirs
of the Australasian Association of Pal-
acontologists 15; 225-248,

COOK, А, G. 1993, Fleicherviewia sepiaia, а new
high-spired, septate gastropod from the Devonian
of north Queensland, Journal of Paleontology 67:
816-821.

HEIDECKER, E. 1959. Middle Devonian molluscs
from the Burdekin Formation of North Queens-
land, University of Queensland, Department of
Geology Papers 5(2): 3-11.

HILL, D. 1939. The Middle Devonian rugose corals of
Queensland 1. Douglas Creek and Drummond
Creek, Clermont District. Proceedings of the
Royal Society of Queensland 50: 55-65,

HILL, D., PLAYFORD, G. & WOODS, J, T. 1967.
Devonian fossils of Queensland, (Queensland
Palacontographical Society: Brisbane) 32pp.

HORNY, R. 1955. Palaeozygolpeuridae, nov, fam.
(Gastropoda) from the Devonian of Central Bohe-
mia. Sbornik Ustredniho ustsvu geologickeho
oddil paleontologicky 21: 17-143.

JELL, J. S. & HILL, D. 1969, Devonian Corals from
Ukalunda. Geological Survey of Queensland.
Publication 340: 1-27,

1970. Revision of the coral fauna from the Devonian
Douglas Creek Limestone, Clermont, central
Queensland, Proceedings of the Royal Society of
Queensland 81: 93-120,

JONES, О. А. 1941. The Devonian Tabulata of Douglas
and Drummond Creeks, Clermont, Queensland.
Fre of the Royal Society of Queensland
53: 41-60.

LINSLEY, R. M, 1968, Gastropods ef the Middle
Devonian Anderdon Limestone. Bulletins of
American Palaeontology 54( No. 244): 328-465.

KASE, T & NISHIDA, T. 1986. A Middle Devonian
gastropod faunule from the Kitikumi Mouniatins.

435

northeast Japan. Bulletin of the National Science
Museum, Tokyo. Series C 13(3): 73-89.

1988. Two new Middle Devonian gastropods from
the Nakazato Formation of Kitikami, Northeast
Japan. Saito Ho-on Kai Special Publication (Pro-
fessor Tamio Kotaka Commerorative Volume).
257-261.

KNIGHT, J. В. 1937. Genotype designations and new
names for invalid homonyms among Paleozoic
gastropod genera. Journal of Paleontology
11:709-714.

1941. Paleozoic gastropod genotypes. Geological
Society of America Special Papers. 32. (Geolo-
gical Society of America: Baltimore). 510pp.

KNIGHT, J. B., COX, L. R., KEEN, A. М. BATTEN,
R. L, YOCHELSON, E. L. & ROBERTSON, К.
1960, Systematic Descriptions. Pp.1169-331. In
Moare, R. C. (ed.) Treatise on Invertebrate Pal-
aeontology. Part I Mollusca 1. (Geological Soci-
ety of Amenca and University of Kansas:
Lawrence, Kansas)

PHILIP, С М. & PEDDER, A. E. H. 1967. Stratigraphi-
ral correlation of the principal Devonian lime-
stone sequences of castern Australia. Pp.
1025-1041 In Oswald, D. (ed) International Sym-
posium on the Devonian System. Volume 2 (Al-
berta Society of Petroleum Geologists: Calgary }

PARFREY, S. M. 1989. Early Devonian fossils from
the Ukalunda Beds, north-west o£ Mount Coolon,
central Queensland. Queensland Government
Mining Journal! 80: 20-21.

TASSELL. C, B, 1977, Gastropods from some Early
Devoninn limestones of ihe Walhalla Svnelinor-
ium, Central Victoria, Memoirs of the National
Museum of Victoria 38:23 1-246.

1982, Gastropods from the Early Devonian
"Receptaculites" Limestone, Таста, New South
Wales. Records of the Queen Victona Museum
TT; 1-59,

WHIDBORNE. G. Е. 1889-1892. A monograph of the
Devonran fauna of the South of England, Volume
1, The fauna of the limestone of Lummaton,
Wolborough, Chircomne Bridge and Chudleigh.
Palaeontographical Spciely Моеса, (Pal-
acontographical Society: London.) 344p.

APPENDIX

OML1005 Un-named tributary of Mary Creek, 80m from creek in prominent eroded gullies, 20° 47.7° S, 147°
20.2" E, Old Hidden Valley Station. Ukalunda Beds.Collected A. Cook & N. Camilleri, 1994.

QML1006 Un-named tributary of Mary Creck, 200m NE of L1005, section centred on 20° 47,7 S, 147" 20.37E,
Old Hidden Valley Station. Ukalunda Beds. Collected А. Cook & N, Camilleri, 1994.

QML1008 Small un-named creek 1,5km S of ‘Sugarloaf Dam’, at 20° 55.4", 147

Beds. Collected А, Cook & N. Camilleri, 1994,

d 3.8", Pyramid Station, Ukalunda

GSQL2765 Northem tributary of Boundary Creek, at metric grid GR 009 539, Mt Coolon 1 : 10000 sheet (8355).

Ukalunda Beds. Collected C. Wilkinson, 1988,

JCUL810. Un-named creek, 1.5km west of "The Sugarloaf” at metric grid GR 068 R68, Glendon 1: 100 000

sheet, Pyramid Station. Collected J, Bennedick, 1993,

436

TWO SIGNIFICANT VERTEBRATE FAUNA RE-
CORDS FROM MID-ALTITUDE WET TROPICAL
RAINFOREST, LAMB RANGE STATE FOREST. Mem-
oirs of the Queensland Museum 38(2): 436. 1995:- A fauna
survey of low to mid altitude wet tropical rainforest on the
easter escarpment of the Lamb Range State Forest (17°00"S,
145'40"E) was conducted as part of the Queensland Electric-
ity Commission's Chalumbin-Woree 275 kV transmission
line environmental impact stàtement. Much of the study area
is dominated by vegetation (уре 2a mesophyll vine forest, with
small areas of of types 8, 13c, 131, 14 and 16a (Tracey, 1982;
Kutt etal, 1995), The survey examined 25 potential ndge-top
lower sites ranging in altitude from 300-700m including the
linenr habitat corridor between them. A total of 173 vertebrate
fauna species were recorded (Kutt et al., 1995) including
twleve of conservation significance: Two of these are consid-
ered to be noteworthy and are reported below.

Northern Beltong Bet/ongia Iropica

Locality: ITOT 20" S, 145740 20" E, Altitude: 620m, Date
10 November 1994, Нараг. паре top (spur), leading to
higher altitude forest, Vegetation: Type 13c vine forest with
emergent Eucalyptus grandis and midstorey Acacia
melanoxylon, A, aulacocarpa (Tracey, 1982), Record rype:
hair sample in canine faecal pellet. Number of individuals
recorded: unknown, likely to be single, Conservation status:
Endangered (Schedule 1, Commonwealth Endangered Spe-
cies Protection Act 1992),

The Northern Bettong is thought to be restricted to a
narrow hand of tall open forest (typically Casuarina torulosa
forest and medium Eucalyptus acmenoides, E. phueotricha,
E. intermedia woodland/open forest) on the western edge of
the Wet ‘Tropics World Heritage Area, running from Mt
Windsor to Ravenshoe (Winter et al., 1991), The site where
the Northern Betiong hair sample was collected represents
atypical habitat for this species, though it his been historically
recorded from closed forest communities (K, Vernes, pers.
comm). It is likely that the hair sample originated ex-situ, as
canine predators utilise large home ranges (e.g. nv. 21km2 for
Dingos in south-east Australian forests, av, 30km? in Kakadu
(Corbett, 1995), with vagrant individuals foraging over even
larger distances, Nuinerous old forestry racks alse traverse
the region, which would facilitate rapid movement of large
mobile predators, The closest known populations occur 10 km
west (straight-line distance), adjacent to Davies Creek Na-
tional Park. However, given past records of B. rropica using
rainforest habitat and the occurrence of potentially suitable
open forest and rainforest communities in the region between
where the hair sample was recovered and Davies Creek, there
is a possibility that individuals or populations may exist in
other areas in the Lamh Range State Forest.

Flute-nosed Bat Murina Jlorium
Locality: 16°59'20"S, 145° 3T 00"E, Altitude: 640m,
Date: 21 October 1994, Habitat: ridge-top, along access track
lo transmission line clearing, Vegetation: Mesophyll vina
forest type 2a, with fringing (3с vine forest (with emergent
Eucalyptus grandis, midstorey Acacia melanoxylon, A. au-

MEMOIRS OF THE QUEENSLAND MUSEUM

lacorarpa) on surrounding ridge lines (Tracey, 1982). Record
гуре? harp-trap, Number of individuals recorded: single male,
forearm 35,5mm, Conservation status’ Vulnerable (Queens-
land Nature Conservation [Wildlife] Regulation, 1994), Sym»
patric species trapped: Eastern Horseshoe Bal Rhinolophus
megaphyllus.

The Flute-nosed Bat was once considered Australia's
‘rarest’ mammal by virtue of a single record from clouded
upland (1120 m) rainforest (Richards ef al. 1983). More recent
records include specimens from upland rainforest (2 1000m
asl) near Ravenshoe , lowland rainforest (<250m) at Rowville
and Gap Creek, Cedar Bay, NEQ (H. Spencer, pers. comm.)
and from specimens of uncertain taxonomic status from Iron
Range (Van Dyck, 1991).

Published knowledge of the bats biology and habitat is
limited (Richards et al., 1983; museum records), The capture
теропец here represents the first mid-altitude record for the
species and the first trom the Lamb Range State Forest and
surrounding region,

Acknowledgements

I thank Marc Hero, Hugh Spencer, Karl Yernes, (James
Cook University, Townsville) and Trevor Hunt (Queensland
Electricity Commission). Barbara Triggs (c/- Dead Finish,
Genoa, Vic, 3891) identified the Northern Bettong hair sam-
ple. Trapping was conducted under QDEH permit to take no.
T-01 181 and the QDPI forestry permit to collect по, 75 1/2.

Literature Cited

Corbett, L. 1995. The Dingo in Australia and Asia, (UNSW
Press: Sydney)

Ingram, GJ. & Raven, R.J. 199]. An atas of Queensland's
frogs, reptiles, hirds and mammals. (Queensland Mu-
séum Brisbane),

Kutt, A., Skull, 5., Burnett, S. & Kemp, J, 1995, Chalumbin
to Woree 274 kV transmission line environmental im-
pact assessment: flora and fauna, Components 3 and 4,
Unpublished ACTFR report no. 95/04 10 the Queens-
land Electricity Commission.

Richards, 6.C., Hall, L.H., Helman, Р.М. & Churchill, S.K.
1982, First discovery of a species of the rare Tube-
nosed insectivorous bat (Murina) in Australia, Ausira-
lian Mammology 5(2): 149-151.

Tracey, J.G. 1982. The vegetation of the humid tropical region
of North Queensland. (CSIRO: Atherton, Queensland).

Winter, J.W., Atherton, R.G., Bell, F.C. & Pahl, 1.1. 199].
Distribution of selected north-eastern Australian
rainforest mammals, Chp. 10in The Rainforest Legacy:
volume 2. G. Werren & P. Kershaw (eds). (АНС,
AGPS: Canberra).

Van Dyck, S.M. 1991. The status of mammals. Pp. 394-453
In Ingram, GJ. & Raven, RJ. (eds) An atlas of
Queensland's frogs, reptiles, birds and mammals.
(Queensland Museum: Brisbane)

A.S. Kuti, Australian Centre for Tropical Freshwater Re»
search & SB. Burnett, Zoology Department, James
Cook University, Townsville, Queensland 4811, Aus-
tralia; 10 June 1995.

THE MAYTOWN OCHRE SOURCE
BRUNO DAVID, ALAN WATCHMAN, ROSEMARY GOODALL & ERIC CLAYTON

David, B,, Watchman, A., Goodall, К. & Clayton, E. 1995 12 01: The Maytown ochre source,
Memoirs af the Queensland Museum 38(2): 441-445. Brisbane, ISSN 0079-8835,

The mincralogy and elemental fingerprint of a well-defined ochre source near Maytown,
north Queensland are described. Characteristics of the ochre were investigated via petro-
graphic, XRD, SEM/EDXA, FTIR and PIXE/PIGME analyses, Potential archaeological
implications of this source are briefly explored. [ ] Ochre, sourcing, rock art, north
Queensland, Australian prehistory,

Bruno David, Department of Anthropology and Sociology, University of Queensland,
Queensland 4072, Australia; Alan Watchman, Data-Roche Watchman Inc,, 1631 Rue Eden,
Ancienne-Loretle, Quebec, Canada; Rosemary Goodall, Department of Chemistry, Queens-
land University of Technology, 2 George St, Brisbane 4000, Australia; Eric Clayton,
Australian Nuclear Science and Technology Organisation, PMBI, Menai, New South Wales

2234, Australia; 15 June 1995.

Earth pigments (ochres) were, and in many
places continue to be, commonly used by Aborig-
inal people to paint, draw, stencil or print on a
variety of surfaces (e.g., wooden objects, cave
walls, bodies). Ochres were sometimes obtained
from distant places, at times measunng in the
thousands of kilometres. This was especially thc
case with ochres of high repute, be it for their
particularly high physical qualities or for their
special Dreaming significance (e.g., Peterson &
Lampert, 1985; Sagona, 1994). In this paper, we
differentiate between a source, being a location
of readily available raw material (not necessarily
exploited), and a guarry, being a demonstrably
exploited source.

Once characterised, well-defined ochre
sources offer archaeologists the potential of pro-
venancing ochre excavated from archaeological
deposits, Once archaeological specimens are pro-
venanced, prehistoric trade orexchange networks
and systems of inter-regional interaction can be
reconstructed. For such a program to succeed,
however, elemental and/or mineralogical
‘fingerprints’ of specific sources need to be first
established. To date, the characteristics of poten-
tial sources of ochre have been almost totally
ignored (but see David et al, 1993; Sagona,
1994). This has resulted in adearth of information
fundamental to the sourcing of archacological
specimens. This paper thus aims 10 initiate a
program of ‘fingerprinting’ ochre sources in
northern Australia. So far, our efforts have con-
centrated on ochrous pebbles from creek and
river beds, The boundaries of such sources tend
to ће ill-defined, although in north Queensland
well-deflned ochre sources and quarries are €x-
tremely rare. In this paper, we report one occur-

rence of a well-defined and highly localised ochre
source from north Queensland, here referred to as
the Maytown Source.

THE SOURCE

The Maytown source is located in the bed of
the Palmer River, immediately west of the now
abandoned town of Maytown, north Queensland
(Limestone Creek 1:50,000 map sheet, edition
1-ААЗ, Grid Reference AC892287). The source
is entirely submerged under water during the wet
season, but totally exposed during the dry. It is an
extremely localised and well detined outcrop of
weak red to dark reddish gray pigment (dry
Munsell 10R 4/2 to 2.5 YR 4/2), measuring c.30
x 30 x Lím (Fig. 1). The source does not show
any évidence of previous extraction, although
prehistoric quarrying activity would be extremely
difficult to detect given the friable nature of the
ochre and the annual flooding of the outcrop in
the Palmer River bed. No other ochrous outcrops
have yet been noted nearby.

The ochre appears to he homogeneous to the
naked eye, A large piece of ochre (с.20 x 10 x
Sem) and a few smaller pieces were detached by
us from the block outcrop for mineralogical and
Pomena) analyses. The results are presented

low,

X-RAY DIFFRACTION (XRD)

A flake broken from a larger piece of the
Maytown ochre source was pulyensed and finely
2round using an agate pestle and mortar. An XRD
pattern (Laval University spectrum #3569) was
obtained using a Siemens diffractometer
equipped with a Copper anode (1.54184Ко X-
rays) operated at 40kV and 20mA. Two theta scan

442

FIG. 1. The Maytown Source,

range was from 5 to 65 degrees at 10 degrees two

theta/minute and a time count of 1.2 seconds.
The spectrum of the pigment is predominantly
quartz with minor muscovite (Fig. 2).

SCANNING ELECTRON MICROSCOPE ENERGY
DISPERSIVE X-RAY ANALYSIS (SEM/EDXA)

SEM observations and analyses were made on two
sub-sample flakes, mounted carbon discs and coated
with Au-Pd, using a JEOL JSM-840A SEM with a
germanium window and equipped with a Tracor
Northem energy dispersive X-ray analytical system.
Operating conditions were 15kV and 100UA with a
working distance of 15mm.

A relatively uniform composition was mea-
sured across the broken pigment samples. Quali-
tative assessment of elements present include Si,
O, AI, Fe and K (Fig. 3). There may also be a very
slight trace of carbon-bearing mineral, but the
very low peak could also reflect a slightly higher
background. Secondary electron images of the
fractured surfaces (Fig. 4) show a sub-parrallel
arrangement of platy crystals, typical of a schist.
The visual microscopic observations and the ele-
mental compositions, when combined, indicate
that the pigment is a ferruginised quartz-mica
schist.

MEMOIRS OF THE QUEENSLAND MUSEUM

FOURIER TRANSFORM A INPRARED ANALYSIS
(FTIR)

The infrared spectrum was recorded using a
Perkin Elmer FTIR 2000 with a MTEC pho-
toacoustic detector on a solid piece of ochre. The
spectrum (Fig. 5) indicates a mica species with
bands typical of muscovite or sericite. Small
bands in the OH stretching region of the spectrum
indicate small amounts of a clay mineral. A small

Maytown pigment source

Muscovite

FIG. 2. XRD spectrum.

MAYTOWN OCHRE SOURCE

u

| |
| asa e cL a

VS + 255 10 240
MAYTOWN TRAOK

FIG. 3. SEM/EDXA spectrum.

band at 1430cm:! indicates a carbonate in the

sample.

PIXE/PIGME

The PIXE/PIGME results were recorded on the
2.5 Mev van der Graff accelerator facility at
Lucas Heights. A pinhole filter was used to re-
duce the effects of high Fe concentrations. Seven
samples were analysed in two separate runs (four

443

samples in 1990 and three in 1994). The 1994
samples (1-3 in Table 1) are those used for infra-
red analysis. Raw results were calibrated against
geological standards and calculated using the
PIXAN package (Clayton, 1986). Quantitative
assessment of 28 major and trace elements was
made, and the results of 15 elements are given in
Table 1. The presence of Ca, Ti and Na in minor
amounts (1-4%) appears to be typical of ochres
from this region (David et al., 1993). Fe is present
in high amounts, as is to be expected for a red
coloured material. Si and Al are also major ele-
ments, K is present as a major component; this is
consistant with a muscovite-mica type mineral-
ogy.
Multivariate (Average Linkeage Cluster) anal-
ysis undertaken on nearly 100 red ochre samples
from northern Australia have linked the seven
Maytown source samples presented here into a
single cluster, separating out from all the other
(non-Maytown source) samples. The implication
is that, on elemental characteristics alone, the
Maytown source is quite distinctive from the
other known ochre sources of northern Australia.

FIG. 4. SEM image. Scale= jim.

TABLE 1, Element concentrations determined by PIXE/PIGME
analysis. All concentrations are mg/kg unless noted as weight

percent.

sr [no [mso |123.0 (633 |702

The results of these investigations will be pre-
sented elsewhere.

The Maytown ochre source, a ferruginised
quartz-muscovite schist, is the first to be system-
atically described from north Queensland. Lim-
ited by a lack of comparative material, it is
therefore too early to determine with any author-
ity whether or not local cave paintings were made
with this material. Nevertheless, preliminary

l ñ * |
\ | | /

' \ || j
j Т.
\ || ү | ki |
А \ I

MB |

>
P nn

a

—>
E

Agi

ва

Фа мю аю Ios dos oo Vw а бю юу,

FIG. 5, Fourier Transform Infrared Photoacoustic
(FTIR-PAS) Spectrum,

[No || |2 J|] ga 5 je [| |
Е pen [mun [p30 [i909 "ees i6i64 110055
[Na _ |4505.0 |3258.0 |4234.0 |4430.8 |3680.7 |4387.0 [4194.4 |
Акаа js js po pa ра ра d
sim) [327 290 [335 jas |395 fasa (375 |

11100 |
тіж) [05 os jos jos joe jos jos |
Ma. eoo 760 fasso [ps jos ia 2s |

230 — 6.6 |
c. fso [wo lso [s loo [ss |o: |
Iz |700 (sto |szo [s24 Моо (1004 fass |
me s [iso pons [nes pe [nez тз

z uos зао пто [132 loo Joo [oo —

MEMOIRS OF THE QUEENSLAND MUSEUM

mineralogical investigations of paintings
from the Blue Figures, Cockatoo,
Quinkans, Longtom and Mushroom Rock
artsites near Laura (to the immediate north
of the Maytown Source) have revealed
similar, although not identical, mineralog-
ical and SEM/EDXA features to the May-
town pigment source (Watchman et al.,
1993). This does not prove that pigment

кое) |за las lao ja: b3 s [sa | samples from the Maytown Source were
[ca looo [noo [1100 |a laoo [ooo [soo | used in Laura paintings, but it does sug-

gest that similar geological material in the
Lower Proterozoic Hodgkinson Forma-
tion (de Keyser & Lucas, 1968) was used

rms |53 | [or 37 tag tas Kee Uy
| painting pigment. It also high-
TH 150 fiso | TENES nm lights the need for a detailed mapping of

| potential pigment sources across north
Queensland. It is only under such a re-
search program that definitive statements
on the provenance of archaeological oc-
| hres will be warranted.

ACKNOWLEDGEMENTS

We thank Mr. and Mrs. Wilson for allowing us
to collect samples of the Maytown ochre; the
Department of Anthropology and Sociology, the
University of Queensland and the School of
Chemistry (especially J. Bartley, G. Kimber & P.
Fredricks), Queensland University of Technol-
ogy for support, and the Australian Institute of
Nuclear Science and Engineering (AINSE) for
two separate grants (to David and Goodall) to
undertake PIXE/PIGME analyses at the ANSTO
laboratories at Lucas Heights.

LITERATURE CITED

CLAYTON, E, 1986, PIXAN: the Lucas Heights PIXE
analysis computer package. Australian Atomic
oe Report Ѕепеѕ AAEC/M 113:

DAVID, B., CLAYTON, E.& WATCHMAN, A, 1993,
Initial results of PLXE analysis on northem Aus-
tralian ochres. Australian Archaeology 36: 50-57.

DE KEYSER, F. & LUCAS, K.G, 1968. ‘Geology of
the Hodgkinson and Laura Basins, North
Queensland'. (Bureau of Mineral Resources Ge-
ology and Geophysics Bulletin 84: Canberra).

PETERSON, N. & LAMPERT, R. 1985. A central
Australian ochre mine. Records of the Australian
Museum 37(1); 1-9.

SAGONA, A. (ed) 1994. ‘Brising the Red Earth:
Ochre Mining and Ritual in Aboriginal
Tasmania’. (Melbourne University Press: Mel-
bourne).

MAYTOWN OCHRE SOURCE 445

WATCHMAN, A., SIROIS, J. & COLE, N. 1993. *Archaeometry: Current Australasian Studies'
Mineralogical examination of Aboriginal rock- (Department of Prehistory, Research School of
painting pigments near Laura, north Queensland. Pacific Studies, Australian National University:
Pp. 141-150. In Fankhauser, B. & Bird, R. (eds.), Canberra).

DEGRADED RIPARIAN HABITAT AT CHARTERS
TOWERS AND ITS POTENTIAL TO DECREASE
LOCAL DIVERSITY AND/ OR CAUSE EXTINC-
TIONS. Memoirs of the Queensland Museum 38(2); 446,
[995.— The imperzance of riparian zones as pabura! wildlife
corridors, as refuges during dry seasons aad as centres of
increased species diversity and abundance has been well
documented (Redford & de Fonesca, 1986; Bennett, 1990;
Williams, 1994). In morthem Australia, npárian zones form
narrow corridors of mesic streamside habitat extending into
drier inland woodlands (Williams, 1994), and allow species
otherwise unable to survive in arid and semi-arid habitat ¢e.g.
aquatic, semi-aquatic, coastal and wener forest species, mi-
gratory species) to expand into these areas (Land Conserva-
lion Council, 1987), Features of прапап vegetation that
coniribule to Ihe observed high diversity and abundance of
fauna include its Florsie and sumructural diversity and hence,
the higher availability of resources (Bennett, 1990; Williams,
1994). Riparian habitats are therefore important sites for local
and regional species conservation. If riparan habilat (s de-
graded or altered, there is a potential for a decrease in species
diversity and abundance, and possibly for shon and long term
species extinctions.

A shor terrestrial flora and faura survey of riparian habitat
was conducted in June 1994 on the Burdekin River near the
Charters Towers Weir by the Australian Centre for Tropical
Freshwater Research, James Cook University of North
Queensland, The study area eoenpriseif The river impoundment
and riparian zone to the bigh Ievee bank, to а poirt Skm
upstream nf the current weir. Veriebrate fauna was surveyed
using à combination of Elliot and cage trapping, spotlighting,
active searching and bird census, Vegetation wad surveyed by
foot iraverse. Secondary sources for Mora and fauna records
were also exanined. A polal of E bird, 12 mammal (four
introduced’, seven reptile ard one impoduced amphibian spe-
cies were Quring the peesent survey. Acklitional
sources (Queensland Museum records; White, unpubl, data)
list а further four native mamenals, six reptiles and four bird
species. A total of 99 vascular plants were recorded, 35% of
which were exotic. Three major vegetation communities were
identified: Melaleuca leucadendra, М, arpenta and M,
viminalis river bed and bank community, tall (up 25m) closed
to open Euzlvprue tereticarmis woodland on the first levee
bank; гай to medium open mixed species woodland (E. res-
sellaris, E, eryrheophloin, E. pinna, E dolichocarpa, E,
drepanophyila} on the highest levee bank.

Exotic species dominated the understorey in all vegetation
types surveyed, Of particular coneern 1% infestation of Rubber
Vine Crypéostegia grondiflora, which threatens the long-term
viability of the two woodland communities. Open canopy
observed in the E. 1ereticarmis commanity is an artefact o£ this
exotic species reaching the canopy and killing mature trees:
Recruitment of trees in the mid and understorey was evident,
bur these Were also being smothered Extensive grazing of
cattle along the river bank has also degraded the native vege-
tation in the riparian zone, As a result, ground cover has been
substantially reduced (in some cases 15-20%), exotic invasion
enhanced, soil erosion increased and recruitment of naive
species decreased.

Though the bird communily was diverse and species rich,
24% of these were waterbirds, present in high numbers diae to
the impoundment, Excluding waterbirds, 47% of all species
recorded were ‘uncammon’ («5 observed). A number of
typical пралат and woodland species that forage in ihe-shrub
and роі layers Were observed to be of reduced abundance

MEMOIRS OF THE QUEENSLAND MUSEUM

пе absent (e.g. meanhizids, sylviids, muscicapids, cuculiis.
malurids). Mediam 10 large macropeds (6 spp.) and Feral
species (4 spp.) were Ihe dominant element terrestrial
(иий. No small ground-dwelling mammals were recorded.
either by live trapping or by secondary sources, This is à
typical m thr L arid and semi-and Queensland
(Уап Dyck, 1991), though confirmation of the total 3hsence
of small terrestrial fauna im the study area requires further
survey. Both observed patterns are a reflection of extremely
poor condstion and quality of the ground and shrub flora, and
overall vegelation communily structure. Exotic weed inva-
sions, excessive grazing, unrestricted bank access by catile
and river bank erosion have all combined to create a riparian
habitat that is degraded and unlikely to act аз а refuge for fauna
orasacentre of increased biotic diversity and abundance. Two
large vertehrate species recorded have the potential to become
extinct in the study area within the near future. A nesting pair
of White-bellied Sea-cagles and a Greater Glider Perauroides
volans were recorded in the Queensland Blue Gum E. rer-
eiiéornis woodland. As outlined above, this community is
heavily infested with Rubber Vine, and has already regressed
from 2 closed woodland to an open woodland as a result of
the loss of mature trees, Loss of the dominant overstorey
would result in the loss of nesting, denning and breeding
resources for these two species, Therefore the conservation
status of these two species in the local area is considered Ko he
‘endangered’ -species at serious risk of disappearing from the
wild stare within 10-20 years if present land use and other
cuts factors continue 10 operate (Ingram & Raven, 1991).

A number of limitations may have influenced many of the
patterns observed including a relatively short survey, season
and prevailing drought conditions, Riparian arcas аге import-
апі sites [ог local and regional species diversity and conser-
vation, especially im sub-optimal conditions, and degradation
мін such sanes creates the potential for а reduction in
biodiversity and local species extinctions. Degraded ripsnan
habitan and vegetation such às this is prevalent throughout the
Burdekin- Haughton catchment and the careful management
and rehabilitation of these zones should be a priority Гог
conservation within tis region.

Literature Cited

Bennett, А.Р. 1990. Habisi corridors. Their role in wildlife
management and conservation. Arthur Rylah Institute
for Environmental Research, Department of Conserva-
tian and Environment, Victoria.

Ingram, G.J, & Raven, RJ, (eds) 1991, Am atlas of
Queensland's frogs, reptiles, birds and mammals
(Queensland Museum Brisbane).

Land Conservation Council. 1987. Report on the Mallee area
feview, LOC, Melbourne, Victoria

Redford, К. & de Fonesca, С. 1985. The role of gallery forests
in the zoageagraphy of the Cerrado s non-volant marn-
mals fauna. Biotropica, 18:126-135.

Van Dyck, S., M. 1991, The status of mammals. 349-353. In,
Ingram, G,J, & Raven, R,J.(eds), An atlas of
Queensland's frogs, reptiles, birds and mammals
(Queensland Museum: Brisbane).

Williams, S. 1994. The importance of riparian habitats to
vertebrate assemblages in North Queensland woo-
lands. Memoirs of the Queensland Museum, 35(1): 248,

Alex Kurt & Steve Skull, Australian Centre for Tropical Fresh-

water Research, James Cook University, Townsville, Queens.

land, 38) | Australia; (U June 1995.

NEW RECORDS OF CRYPTOPODIA (CRUSTACEA: DECAPODA:
PARTHENOPIDAE) FROM AUSTRALIA

РЈЕ. DAVIE & Р.А. TURNER

Davie, P.J.F. & Turner, Р.А. 1995 12 01: New records of Crypropodia (Crustacea: Decay à
Parthenopidae) from Australia. Memoirs of the Queensland Museum 38(2): 447-462. Bris-
bane. ISSN 0079-8835,

Six species of Cryptopodia are recorded from northern Australian waters and an identifica-
tion key is provided. Three, C. angulata Н. Milne Edwards & Lucas, 1841, С, dorsalis White
& Adams, 1847, and C. pan Laurie, 1906, are new to the Australian fauna and represent large
range extensions. The known range of C. queenslandi Rathbun, 1918, is extended, and
allometric growth is noted for this species. Dorsal and ventral photographs, and figures of
the male first gonopods, are provided for all six species. Crustacea, Brachyura,
Parthenopidae, Cryptopodia, Indo-West Pacific, Australia.

P.J.F. Davie & Р.А. Turner, Crustacea Section, Queensland Museum, P.O, Box 3300, South

Brisbane, Queensland. 4101 Australia; 20 September 1995,

Collections on which the current study is based
were largely provided by two surveys undertaken
by the Commonwealth Scientific and Industrial
Research Organisation (CSIRO), off North West
Shelf, north-western Australia aboard the R.V.
Soela, and in the Gulf of Carpentaria, using the
К.У. Southern Surveyor. These have been supple-
mented by additional material in the Queensland
and Northern Territory Museum collections. The
North West Shelf was investigated using a beam
trawl and epibenthic sledge, and the results of the
analysis of the structure of the decapod commu-
nity have been reported by Ward & Rainer
(1988).

There has been relatively little work on the
Indo-West Pacific Parthenopidae, with Flipse
(1930) still being the single most important refer-
ence. Miers (1879), Haswell (1879), and Camp-
bell & Stephenson (1970), have made the most
significant contributions to our knowledge of the
Australian fauna. Very little is known of the rich
group of tropical parthenopid species.

Abbreviations used in the text are: c.b., cara-
pace breadth; c.l., carapace length; G1, first male
gonopod; NHM, The Natural History Muscum,
London; NTM, Northern Territory Museum,
Darwin; QLD, Queensland, Australia; QM,
Queensland Museum, Brisbane. Measurements
given in the text are of the carapace breadth
(measured at the widest point) followed by
length.

SYSTEMATICS

KEY TO AUSTRALIAN SPECIES OF
CRYPTOPODIA

1, Carapace with cardio-intestinal region relatively
flat, and laterally demarcated by long, narrow,
very deep grooves so as to appear lyre-shaped

14 FS oF As C. dorsalis White & Adams, 1847
Cardio-intestinal region separated by broad shal-

low depressions; cardiac region more-or-less ele-

Mam as o daga ME ep a gmt opt oo 2

2. Carapace with margins strongly serrated and spi-
fos s eoe. LONE ea ule: stabs ite he he
‚С, angulata H. Milne Edwards & Lucas, 1841
Carapace sometimes with anterolateral margins
moderately spinous, but posterior margin at
mostcrenellated.......-..-....4.-

3.Third maxilliped noticeably swollen, with conspic-
uous, broad, flattened granules on inner margin
of ischium which may extend over the entire
outer surface. . .. .. .. C. pan Laune, FR
Third muxilliped notswollen .........

4. Carapace surface relatively smooth except for
slightly g granular crests; shallow gastric depres-
sion centrally; presence of closed fissures radiat-
ing in trom margins visible dorsally ......

v emu. i C. queenslandi Rathbun, 1918
Carapace surface more-or-less coarsely granu-
lated and pitted; deep gastric depression cen-
trally; marginal closed fissures not obvious
doisally «^. 5. Lo us vL I o utut ac ж.н 5

5, Carapace with margins of rostrum subparallel!
over proximal half; branchial, cardiac, and gas-
trc regions strongly inflated ‚,,,,,,,,,

©» җе, C. fistulosa Chiong & Ng, 1994

448 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Male first gonopods of Cryptopodia species, showing entire view and magnified view of apex. A, B, C.
pan (QMW18473). C, D, C. queenslandi (QMW 18981). Е, Е, C. angulata (QMW18306). С, С! (rotated view).
H, C. fistulosa (QMW18980). I, J, C. dorsalis (QMW18291). Scale line: A, C, E, G-0.05mm. D, B, F, Н,
J=0.0imm; I20.10mm.

NEW RECORDS OF CRYPTOPODIA 449

FIG. 2. Cryptopodia angulata H. Milne Edwards & Lucas, 1841 (QMW18299). A, dorsal view. B, ventral view.
Scale line in mm.

450

Carapace with margins of rostrum tapering from
the base; branchial, cardiac, and gastric regions
not strongly inflated

se à o» à

Cryptopodia angulata
Н. Milne Edwards & Lucas, 1841
(Figs 1E, F; 2A, B)

Cryptopodia angulata Н. Milne Edwards & Lucas,
1841: 481, pl. 28, figs 16-19; Alcock, 1895; 282;
Chopra, 1935: 473; Chhapgar, 1957: 415, pl. 4;
Ahmad et al., 1972: 15 (listed); Tirmizi, 1980: 107
(listed); Tirmizi & Kazmi, 1983: 369 (listed); 1991:
211-213; Banu & Nurul Hudu, 1989: 646-647.

Cryptopodia angulata var, cippifer Alcock 1895: 283;
Flipse, 1930: 62, 82.

MATERIAL EXAMINED

CSIRO, F.R.V. Southern Surveyor, Gulf of Carpenta-
па: QMW 18305, & (34,2 x 20.2mm), 13 02°5,
139?22.2'E, Stn 36, 5&m, 24.11.1991. QMW18306, d
(21.9 x 14.1 mm), 1325.6 S, 138'36.0' E, Stn 34, 54m,
24.11,199]. QMW18298, ? (38.2 x 22.2mm),
14'00,7'S, 139'11.6'E, Stn 35, 59m, 28.11.1990,
QMW18299, d (37.1 x 22.8mm), ? (54.6 x 30,2mm),
1328.9'S, 139°11.9'Е, Stn 34, 57m. 28.11.1990,
QMW17334, d (37.8mm c.b., rostrum damaged),
1427,45, 138°11,9°E, Stn 20, 52m, 25,11,1990,
OTHER MATERIAL: МТМ Cr0902, 29 (37.3 x 22.1,
51.5 x 28.2mm), Arafura Sea, 1258.0 S, 132°10.08,
Stn HL, 81-82, 27m, 19,10,81, R.V, Gemini.

REMARKS

The present study greatly extends the pre-
viously known range of C, angulara, Specimens
collected from the Gulf of Carpentaria, apart from
8 few minor differences, correspond closely with
the descriptions of Milne Edwards & Lucas
(1841) and Alcock (1895), The triangular gastric
depression, noted by Alcock (1895) as being
‘very deep’, appears to be variable in depth: in
our specimens it is moderately deep, but Chopra
(1935) found it to vary from more or less shallow
to quite deep. On our specimens the border of this
depression had, in most cases, a tubercle at each
branchial angle and, less frequently, two small
tubercles side by side on the anterior angle of the
depression. No tubercles were present on the
summit of the cardiac region. Chopra (1935) and
Tirmizi & Kazmi (1991) found this tuberculation
to be variable, with one small individual exam-
ined by Chopra bearing small spines instead of
tubercles, However, none of the specimens exam-
ined by Chopra (1935), Tirmizi & Kazmi (1991),
nor ourselves had ‘large, erect definitely-placed

MEMOIRS OF THE QUEENSLAND MUSEUM

spines’, as described by Alcock (1895), for C.
angulata var. cippifer.

Alcock described the rostrum as ending in a
sharp point; the rostra of the specimens detailed
hercin are dome-shaped and end in a relatively
blunt point. A dome-shaped rostrum was found
in a male specimen by Tirmizi & Kazmi (1991),

Alcock (1895) pointed to the presence of spines
on the meri of the ambulatory legs. This spination
is present on our specimens but only on the first
and last pair of legs. Tirmizi & Kazmi (1991)
described an identical pattern of spination to that
in our specimens.

Granulation on the carpus of the chelipeds was
variable among the specimens examined by
Chopra (1935), but showed little variation among
the specimens of Tirmizi & Kazmi (1991). Most
of Chopra's (1935) specimens were either
smooth or slightly granular, as described for C.
angulata var. cippifer, by Alcock (1895).
Alcock's (1895) comparison of C. angulata and
its variety suggests that there was no granulation
on the carpi of the chelipeds of his specimens of
C. angulata. The diagram of Tirmizi & Kazmi
(1991) shows more granular carpi than those of
either Chopra's (1935), or our specimens. Banu
& Nurul Huda (1988) reported the presence of a
granular carpus for a specimen collected off Pe-
nang Island, Malaysia, whereas a number of spec-
imens collected by the same authors from the
Chittagong Coast bordering Bangladesh pos-
sessed smooth carpi. There rs only а small amount
of carpal granulation among our specimens.
Granulation is more evident on the ventral sur-
face of the carpus. On two of our specimens, a
male and a female, there is a small, sharp, sub-
distal, median spine on the upper surface of the
carpus. This replaces a small tubercle in the other
specimens. No mention of this character is made
by other authors for C. angulata.

The status of Cryptopodia angulata var. cippi-
Jer Alcock, 1895, is still uncertain, The other
character used by Alcock (1895) to separate C.
angulata var. cippifer from C. angulata, is the
presence of prominent spines surrounding the
central depression. Evidence from Chopra
(1935), Tirmizi & Kazmi (1991), and this study,
suggests a degree of intraspecific variation that
may encompass most of the characteristics de-
scribed by Alcock (1895) for C. angidlata var.
cippifer. It is probable that C. angulata var, сіррі-
Jer is a junior synonym of C. angulata, but exam-
ination of Alcock's type specimens along with a
large series of specimens from across the range

NEW RECORDS OF CRYPTOPODIA 451

FIG. 3. Cryptopodia dorsalis White & Adams, 1847 (QMW18297). A, dorsal view. B, ventral view. Scale line
in mm.

452 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Cryptopodia pan Laurie, 1906 (QMW 18461). A, dorsal view. B, ventral view. Scale line in mm.

NEW RECORDS OF CRYPTOPODIA

of occurrence of С, angulata is required for final
certainty.

DISTRIBUTION

C. angulata: Type locality unknown; Karachi;
west coast of India; Malabar Coast, Orissa Coast,
Sandheads at mouth of Hugli River, India; Cey-
lon; Malaysia; and now northern Australia, from
the Arafura Sea to the Gulf of Carpentaria. C.
angulata var. cippifer is only known from
Karachi, Pakistan (type locality), Bathymetric
range: 52 to 59m.

Cryptopodia dorsalis
White & Adams, 1847 (in White, 1847)
(Figs 1 1, J; ЗА, B)

Cryptopodia dorsalis White & Adams, 1847, in White,
1847a: 125 [nomen nudum]; 1847b: 84; White,
1847c: 205; Adams & White, 1848: 30, pl. 6, fig. 5;
Flipse, 1930: 63, 82,

MATERIAL EXAMINED

CSIRO, R.V. Soela, North West Shelf: QMW 18290,
3 (62.3 x 35.3mm), 19'29.6' S, 118°52.2'E, Stn
OSDO3BT, 40m, 25.10.1983. QMW1829I, 4 (64.5 x
35.8mm), 19°58.6'5, 117'49.4' E, Stn 03D09BT, 43m,
26.06.1983. QMW18292, g (17.1 x 114mm),
19'56.8'S, 117°53.5'E, Stn 03B02BT, 44m,
25.06.1983. QMW18293, 2 (16.2 x 10.6mm),
19'45.7'S, 117'52.0'E, Stn 01ВІЗВТ, 54m,
20,02,1983. QMW 18294, д (20.8mm c.l; lateral mar-
gin broken), 19°03.6'S, 119°03.4'E, Sm OSBI2BT,
82m, 23.10.1983. QMW18295, d (25.1 x 15.5mm),
19'59,2'S, 117'03.6'E, Stn O4BISBT, 52m,
05.09.1983, QMW18296, 3 (11.4 x 8.0mm),
19'30.8'S, 118°49.1°E, Stn 038075, 37-38m,
28.06.1983, OMW 18478, d (11.3 x 7,8mm), ? (16,0
x 11,6mm), 19730.6' S, 11849. 4' E, Stn 03BO7BT, 37-
38m, 28.06.1983. QMW 18552, d (14.7 mmc... lateral
margin damaged), 19'30.8'S, 118749.3'E, Stn
O4B07BT, 38-39:m, 30,08,1983, QMWI8289, 2d
(62.3 x 35.8, 22 4 x 16.4mm), 1955.5 S, 117'55.5' E,
Stn O2BÜ3BT, 42m, 22.04.1983. QMW 18288, 4 (18.0
x 11.5mm), 19°54.6'S, 117'56'E, Stn 01BO3S, 44m,
18,02.1983, QMW18287, 9 (73.6mm c.b., rostrum
broken), 19'55.9'S, 117'55.5'E, Stn 03BO3BT, 42-
43m, 26.06.1983.

OTHER MATERIAL: QMW18297, 3 (61.1 x 35.3mm),
Arafura Sea, 12°15.6°S, 129'15"E, Stn 508, 29m,
17,11,1989, Bureau Rural Resources, QMW 12771, d
(76.3 x 42.4mm), N. of Cape Bowling Green,
19'08.9'S, 147'23.3'E, 09.05.1985, С, Jones, Qld.
Fisheries Service.

REMARKS
The authorship of this species has been attrib-
uted to Adams & White, 1847, by Flipse (1930)

453

in his major revision of the Parthenopidae; how-
ever the first valid citation in White (1847b) puts
the authorship as White & Adams (the very first
mention of the name occurred in White (1847a)
but was a nomen nudum). This species was listed
as new in four separate publications, and there ts
some difficulty in being certain which of the two
descriptions that appeared in 1847 has nomencla-
tural priority. Apparently the first to appear was
the article in the 1847 Proceedings of the Zoolog-
ical Society of London, This volume, according
to the date stamp of the British Museum Library,
was not available until July, 1848, but according
to Sclater (1893) the journal was published in
separate shects prior to binding, and White's ar-
licle on pp, 84-86, appears with sheet cixxv which
was delivered from the printers to the Zoological
Society on 20 July 1847, White (1847c), as cited
here, was date stamped "47.9.22.5" by the British
Museum Library and therefore was available
only later in September, 1847. There could be
some dispute as to whether the earlier published
work was distributed, and thus available, prior to
the second that appeared in September, but this
will probably never be known, and thus I choose
to cite the publications in order of their known
printing dates. Cryptopodia dorsalis 1s also de-
scribed in Adams & White's 1848 report of the
“Voyage of the Samarang’, and in this, authorship
is reversed and given as Adams & White.

The distribution of C; dorsalis suggests that il
is common throughout the waters of northern
Australia and the Indonesian Archipelago, There
is little doubtas to the correct identification of this
species, given the presence of two deep, lyre-
shaped grooves which border the lateral edges of
the cardiac region, a character unique to C. dor-
salis.

DISTRIBUTION

Sulu Sea (tvpe locality); northem Australia,
from the North West Shelf, Arafura Sea, and
north of Cape Bowling Green (Townsville).
Bathymetric range: 29 to 82m,

Cryptopodia pan Laurie, 1906
(Figs ІА, B; 4A, B)

Cryptopodia pan Laurie, 1906; 392, fig. 4, pl. 1, fig. 6;
Rathbun, 1911; 259; Flipse, 1930: 63, 78. 82; Tan
& Richer de Forges, 1993- 131, figs 6E, F.

MATERIAL EXAMINED

CSIRO, К.У. Soela, North West Shelf: OMW18474,
29 (10.1x 7.9; 10.6 x 8.0mm), 20°00.2’S, 117°00.5'E,
Sin OLBI7BT, 53m, 22.02.1983. QMW18476, 4

454 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. Cryptopodia queenslandi Rathbun, 1918 (QMW18309). A, dorsal view. B, ventral view. Scale line in
mm.

NEW RECORDS OF CRYPTOPODIA 455

FIG. 6. Cryptopodia queenslandi Rathbun, 1918, spotted variant (QMW 18981): A, dorsal view. B, ventral view.
Scale line in mm.

456

(14.2mm c.b., rostrum damaged), 19'03.6'S,
119'00.6'E, Stn OLBIIBT, 81m, 13.02.1983.
QMW18464, 4 (23.8mm c.b., rostrum damaged),
19'57,9'S, 117 49.3 E, Stn O3DOTBT, 40m,
26.06.1983. QMWIS46S, 3 (15.5 x 11.7mm).
19*03.2'S, 119°02'E, Stn 06BI2TN, 78-80m,
11.12.1982. QMWI18466, © (14.6 x 1l, 1mm),
19'29.4'S, 118°51.5'E, Stn OSDOTBT, 40m,
25.10.1983. QMWIS8467, 3 (22,1 x 15,1mm),
19'04.6'S, 118'57.9' E, Stn 03BIOBT, 81-821,
30.06.1983. QMWIEB468, 4 (15.1 x 11.0mm),
19'30'S, 11852" E, Stn 050055, 36-37m, 25.10.1983.
QMWIS8469, 24 (22.3 x 15.3; 10.2 x 8,1тт),
20*00.4'S, 117*00.4'E, Stn 04B17BT, 52m.
04,09,1983. QMWI8470, 9 (14.8 x I0, 8mm),
19'05.3'S, 118'54.0'E, Stn O3B04BT, 82m,
29.06.1983. QMWI8471, © (12,7 x 10.7mm),
18'59.1'S, 118'45.9'E, Sin OGBOSTN, 84m.
07,12.1982, OMWIR4T72, I juv. (7,1 x 6.6mm),
19'42.0'S, 11757 E, Stn 018155, 36m, 20.02.1983.
QMW18473, ¢ (23.4 x 16.6mm), 19'24.8'S,
118'57,6'E, Stn O6BO8TN, 47-48m, 08.12.1982.
QMW18475, 9 (13. 1mm c.L, lateral margin dam-
aged), 1959,1'S. 117°51,6'Е, Stn OJBOIBT, 42m,
27.08.1983. OMW 18463, 2 (carapace damaged),
19'56.7'S, 117'53.6' E, Stn OSBOZRT, 40m,
26,10,1983, QMW 18462, 8 (21,0 x 14.2mm),
19*04.9'S, 118'50.6'E, Stn üSBOSBT, 81m,
30.10.1983. QMW 18451, 3 (13.8 x 10.2mm), 2 (17.1
x 12.7mm), 19'05'S, 118750.5 E, Stn OGBÜSBT, 83-
Ват, 29.06.1983,

REMARKS

This study, and that of Tan & Richer de Forges
(1993), greatly extends the known distribution of
this species. Ourspecimens agree closely with the
description of Laurie (1906). In many specimens,
the exposed surface of the ischium of the third
maxilliped is completely covered in large, squa-
mous granules, and in a few specimens, the gran-
ules are more or less coalesed over the outer
two-thirds. This differs from Laurie" s (1906) type
description, where he said that only the inner third
of the ischium is covered in such granules,

DISTRIBUTION

Sn Lanka (type locality); northern Australia,
from the North West Shelf; New Caledonia,
Bathymetric range: 28 to 84m.

Cryptopodia queenslandi Rathbun, 1918
(Figs 1C, D: 5A, B; бА, B, 7A, В)

Cryptopodia fornicate. Haswell, 1879: 454, 1882: 37
[not C. formica (Fabricius, 1781)].

Cryptopodia Queenslandi Rathbun 1918: 26, pl. 12.

Cryptepodiàa fornicate var, Queenslandi: Flipse, 1930:
65-66, 78, 82.

MEMOIRS OF THE QUEENSLAND MUSEUM

MATERIAL EXAMINED ——
CSIRO, R.Y. Soela, North West Shelf: QMW 18322,
9 (22.8 x 15.7mm), 19'29.8'S, 118°52.3'E, Stn
05008ВТ, 37-38m, 25.10.1983. QMW 18323, 4 (17.1
x 11.2mm), 19"29,4' S, 11852. T E, Stn O2BOSBT, 38-
39m, 26.04.1983. QMW 18324, 3d (16.0 x 10.9 -24.3
x 15.7mm), 3? (16.8 x 11.0 - 30,6 x 20,1mm),
19'30.8'S, 118749, 1 E, Stn O4BO7BT, 36-39,
30.08.1983. QMW18325, d (24.5 x 15.5mm),
19°28.4°S, 118'55.2' E, Stn 04B09RT, 39m,
31.08.1983. OMW18326, а (17.4 x 11.4тт), 29
(18.1 x 11,7; 13.0 x 87mm), 19'30.6'S, 11849 4'E,
Stn O3B07BT, 37-38m, 28,06.1983. QMW 18327, 43
(22.5 x 14.7 - 37.6 x 22.7mm), 29 (37,3 x 23,6; 40.8
x 24.6mm), 19'29.4'5, 118524" E, Sm O5D02BT, 37-
38m, 24.10.1983. QMW 18328, 26 (33.5 x 20.6; 33.5
x 20.3mm), 2? (22.5 x 14.7; 20.7 x 13.7mm),
19'20.5'S, 118'52.2' E, Stn OSDOIBT, 37m,
24,10,1983. QMW18329, 24 (16.4 x 11.3; 13.3 х
12.4mm), 2? (16.8x 11.1; 20.2x 12.5mm), 19/28.1'8,
118 55.2"E, Sin (03BO9BT, 38-40m, 28.06.1983.
QMWI8330, ? (11.7 x &.1mm), 19730.9'S,
118'48.7'E, Stn 02В07ВТ, 39-40m, 26.04.1983.
QMW18331, 42 (8 4mm c.l., lateral margin damaged
; 102 x 7.2mm ; 8. Imm c.l., lateral margin damaged;
10.8 x 7.6mm), 19'28.6'S, 118'55'E, Stn 02B095,
38-40m, 26.04. 1982. OMW 18332, 8 (7.5mm c.l., lat-
eral margin damaged), 1955,5'S, 117°55.5'Е. Stn
O2B03BT, 42m, 22.04.1983. QMW 18333, 29 (11.4x
7.9mm ; 11.2 x 7.9mm), 19°29,9°S, 118'52'E, Sin
028085, 38-39m. 26.04.1983. QMW 18334, © (cara-
pace damaged), 1958.3 S, 11749.4' E, Stn 030015,
43m, 25.06.1983. QMW18335, 3 juveniles (3,1 x 2.6
- 3.1 x 4.9mm), 19°29.3'5, 118'52.6 E. Stn
OIBO8RevS, 36m, 15.02.1983. QMW18336, 1 juve-
nile (5.1 x 4,Smm), 19'58,9'S, LI7'51,7 E, Stn
048015. 42m, 27.08.1983. QMW18337, © (B2 x
6.0mm), 19°58.1'5, 117°49.2°E, Stn 020078, 40m,
26.06.1983, QMW 18338, 2 juveniles (4.9 x 4,0, 3.3 x
3.0mm), 19'20'S, 118'53,5'E, Sin ÜIBOSS, 40m,
12,02.1983. QMW18312, 3d. (7.8 x 6.0 - 264 x
16,7mn:), 59 (10.0x 7,1 -24.3 x 15.1 mm), 19°30,9"S,
118°48,7'Е, Stn 02BO7BT, 39-40m, 26.04.1983.
QMW18313, 9 (32.0 x 20.6mm), 19'53.1'5,
118'03.9"E, Stn O6BOIS, 36-38m, 05.12.1982.
QMW18314, 24 (20.2mm c-l., lateral margin dam-
aged; carapace damaged), 19°29.9'S, 11852 .0' E, Sta
050018, 37m, 24.10.1983, QMW18315, 3 (23.1 x
15.1mm), 2Ẹ (26.9 x 17.4; 20.7 x 13.4mm), 1930.9" S,
118749.2'E, Stn OSBOTBT, 33-39m, 25.10.1983.
QMWIS316, 2 (16.5 x 11.90), 19'56.7'S,
117°53.6°E, Sin O5BO2BT, 30m, 26.10.1983.
QMWI18317, 3 (372 x 229mm). 29 (21.2 x 15.0;
22,4 x 14,4mm), 19°29,4'5, 118 52,4" E, Sin
üSDOSBT, 38m, 25.10.1983. ОМА 18318, g (25.5 x
16.7mm), 1929.6 S, 118°52,2'Е, Stn O4B08BT, 38-
39m, 30,08,1983, QMW18319, 2 (16.1 x 10.9mm),
19756,8'5, 117'53,5' E, Stn 03B02BT, 44m,
25.06.1983. QM W | 8320, 9 (14.0mm c.l, lateral mar-
gin damaged), 19°55.6'S, 117 56.0 E, Sm O4BOGBT,
3-44. 26.08.1983. QMWI8321, d (17.5 x

NEW RECORDS OF CRYPTOPODIA

breadth/length ratio

5 ia (5 20 25 30 35 au
carapace breadth (mm)

breadth/length ratio

35

2ü 25 50
carapace breadth (mm)

FIG. 7, Allometric growth equations for C. queenslandi. A, males:
г2=0.82, n-27. B, females: у=0.007х+1.37, 12-0.63,
n=37. Dashed lines represent the 95% confidence interval for the true

у=0.1х+1.32,

mean of y,

11.3mm), 19°30.8°S, 118°49.1°E, Stn 03B07S, 37-
38m, 28.06.1983,

CSIRO, R.V. Southern Surveyor, Gulf of Carpentaria:
QMW18307, 6 (35.8 x 21.5mm), 13'40.1'S,
136'45.6'E, Stn 30, 22m, 23.11.1991. QMW18309,
28 (45.0 x 25,1; 40.3 x 23.4mm), 10'52.6'S,
136.12.1'E, Stn 12, 42m, 21.11.1991. QMW18308, 9
(21.3 x 15.0mm), 10°57.6'5, 140"23' E, Stn 58, 54m,
29.11.1991. QMW18310, 9 (36.7 x 22.8mm),
11°50.8°S, 136'33,9'E, Stn 4, 33m, 18.11.1991.
QMWI8311, 2? (40.7 x 25.2; 26.9 x 17.7mm),
13°02.8'S, 141727.7 E, Stn 45, 21m, 26.11.1991.
QMW18300, d (26.2mm c.l, lateral margin dam-
aged), 11'58.5'S, 140'41.4"E, Stn 63, 53m,
04,12,1991, QMW 18301, d. (21.9mm c.l., lateral mar-
gin damaged), 1033.4'S, 138'42.6'E, Stn 83, 53m,
09.12.1991, OMW18302, d (40.0 x 23.1mm),
11°09.2'S, 139'41.8'E, Stn 82, 55m, 08.12.1991.
QMWI8303, 9? (37.6 x 23.4mm), 15'00.9'S,
140^ 12'E, Stn 43, 48m, 30.11.1990. QMW18304, 2d
(30,8 x 19,6mm; 21 4mm c.l, lateral margin damaged),
29 (30.3 x 18.4; 27.2 x 17.7mm), 16'01.8'S,
140°11.9'E, Stn 41, 31m, 29.11,1990. QMW 7316, 2
(37,1 x 21.0mm), 14'27.5'S, 13842" E, Stn 97, 52m,

457

12.12.1991. QM unreg., d (22.8 x
14.7mm), 143'08.6'S, 11°49.9°E, Sin
007, 22m, 13.1.93.

OTHER MATERIAL: QMW1495, d
(48.8 x 28,5mm), Green Island,
Moreton Bay, 27°26’S, 153'14'E.
QMWIE98I, d (22.8 x 14.6mm),
Shelburne Bay, 11'51.9'S,
143'0R.9'E, Stn 7, 22m, 13.01.1993,
CSIRO, F.R.V. Southern Surveyor.

REMARKS

Flipse (1930) suggested that C.

queenslandi is a variety of C.
Jornicata. However, there are dif-
ferences between these two species
which warrant C. queenslandi
being recognised as a separate spe-
cies from C. fornicata. There are
significant differences in the malc
G1. In C. fornicata, the apex of the
G] is produced into two nearly
symmetrical lobes (Dai & Yang,
1991: fig. 90), whereas in C.
queenslandi, these lobes are highly
asymmetrical, with one being
much smaller (Fig. 1C, D). Also, in
C. fornicata, the G1 tapers more
rapidly than in C. queenslandi, In
these respects, the G1 of our spec-
imens are like those of Campbell &
Stephenson (1970), collected from
Moreton Bay, Queensland. The
chelipeds of C. fornicata are con-
siderably less than twice the length
of the carapace (Alcock, 1895), whereas in C.
queenslandi they are twice the length (Rathbun,
1918). On average, C. fornicata appears to be a
larger species, with the specimens examined by
Dai & Yang (1991) being 3446 larger than thc
largest crabs (females and males) examined in
this paper. Considering the large amount of ma-
terial available for our study this seems signifi-
cant. Rathbun's (1918) adult male holotype of C.
queenslandi was 28% smaller than that of Dai &
Yang's (1991) specimen. The only specimens of
C. queenslandi that approach the size of Dai &
Yang's (1991) specimen are those described by
Flipse (1930) from Indonesia (89% for males and
95% for females); and it would be desirable in a
generic revision to check the identity of his spec-
imens,

The degree of concavity of the posterior margin
seems unreliable for separating the two species.
Rathbun (1918) described the posterior rim of C.
queenslandi as concave in its middle two-fifths;

45 50

40 ag

458 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. Cryptopodia fistulosa Chiong & Ng, 1994 (paratype, QMW 18980): A, dorsal view. B, ventral view. Scale
line in mm.

NEW RECORDS OF CRYPTOPODIA

FIG. 9. Cryptopodia spatulifrons Miers, 1879, holotype
(NHM 1858.172), male first gonopod (left), showing
entire view, and magnified ventral and dorsal
views.(Taken from Chiong & Ng (1994)),

whereas Flipse (1930) said it was straight or very
weakly concave. However, some records have
also described or figured specimens of C.
fornicata as having à convex posterior rim that
forms a continouus smooth edge with the antero-
lateral margin (eg. Sakai, 1976: 292, text-fig 163,
Dai & Yang, 1991).

Flipse (1930) described C. fornicata as being
1.5 times as wide as long, and C. queenslandi as
being twice as wide as long. Dai & Yang (1991)
described C. fornicata as being 1.6-1.8 times as
wide as long, while the specimen examined by

459

Rathbun (1918), a male, had a ratio of 1.8. Wc
have plotted length against breadth for the speci-
mens in this study (Fig. 7A, B); and therc is
distinct allometric growth with the length/breadth
ratio ranging from 1.35 in the smallest specimens,
to c. 1.8 in the largest. This 1s discussed further
later. Therefore we disagree with Flipse (1930)
that the simple breadth/length ratio is useful in
separating the two species.

C. queenslandi normally has no obvious dorsal
patterning, but an unusual specimen from the
Gulf of Carpentaria (б. 22.8 mm c.b.,
QMW18981), bears numerous spots over the en-
tire dorsal surface (Fig. 6). Morphologically it
cannot be separated. Dr P.K.L. Ng has informed
us that juvenile C. fornicata have a striking colour
pattern that is absent in adults, however while
there were many smaller specimens in the present
series, only the individual above showed the dis-
tinctive pattern described.

C. queenslandi appears to be restricted to wa-
ters around northern Australia and possibly Indo-
nesia, whereas C. fornicata, as reported by Sakai
(1976), is found in Sagami Bay, Japan; China
Sea; the Phillippines; the Gulf of Thailand; Sin-
gapore; and westward to the Gulf of Mattaban;
Andaman Sea; Sri Lanka; Palk Straits; and the
Persian Gulf. We believe Haswell (1880) incor-
rectly identified C, fornicata from Port Denison,
north Queensland, and that his specimens were
most likely C. queenslandi.

Allometric growth: Allometric growth has been
reported in the Parthenopidae previously by Gore
& Scotto (1983). As we had an abundance of
specimens of C. queenslandi we did a simple plot
of length/breadth ratios. Fig. 7A, B shows the
linear relationship between carapace length and
breadth for male and female C. queenslandi, re-
spectively. Both diagrams display statistically
significant allometric growth for both sexes
(220.821 (males) and 1220.63 (females),
р<0.05). The correlation coefficients for both
sexes are not significantly different (р>0.05).
However, males broaden to a significantly greater
extent than females with increasing size (p«0.05).

DISTRIBUTION

Cape Gloucester, Bowen, Queensland (type lo-
cality); Java Sea; North West Shelf; Arnhem Bay
(Northern Territory); Torres Strait, Shelburne
Bay and Palm Island (north Queensland);
Moreton Bay (southeast Queensland); Woody
Head (northern New South Wales). Bathymetric
range: 21-55m.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 10. Cryptopodia spatulifrons Miers, 1879 (Holotype d, NHM1858.172, 48.8 mm c.b.): A, dorsal view. B,
ventral view.

NEW RECORDS OF CRYPTOPODIA

Cryptopodia fistulosa Chiong & Ng, 1994
(Figs 1G, Н; ЕА, B)

Cryptopodia spatulifrons- Miers, | 884: 203-204 (spec-
imen from Thursday Island) [not С, spatulifrons
Miers, 1879].

Cryptopodia fistulosa Chiong & Ng, 1994: 952-957,
figs 1A, 2А, ЗА, 4A, 5A, C, D, О, Н.

MATERIAL EXAMINED

CSIRO, R.V. SOELA, NORTH WEST SHELF:
QMWI18995, 9 (23.8 x 16.1mm), 1955.28,
117°56,0°E, Sth OSBO3BT, 40m, 26.10.1983.
QMW18980, d (33,3 x 21.3mm), 19°28.4°S,
118755.2"E, Sin 04B09BT, 39m, 31.08.1983.
QMW 18994, 5 (16,1 x 11.3mm), 20'00.2'S,
117°00.5'Е, Stn 04B 17S, 52m, 04.09.1983,

REMARKS

This recently described species is very similar
in general appearance to C. spatulifrons Miers,
1879. The two species can be separated using the
key provided in the present paper, but Chiong &
Ng (1994) should be consulted for a full list of
characters by which they differ.

DISTRIBUTION

Northern Australia, from Shark Bay, W.A. to
Torres Straight, northern Queensland. Bathymet-
ric range: 5-52m.

Cryptopodia spatulifrons Miers, 1879
(Figs 9A-C, 10A, B)

Crypropodia spatulifrons Miers, 1879; 26, pL5, fig. 10;
Haswell, 1879: 454; 1882: 37; Ortmann 1894; 48;
Flipse, 1930: 63, 78, 82; Chiong & Ng, 1994; 950-
952, figs 1B, 2B. 3B, 4B, 5B, E, F, I, J.

REMARKS

This endemic Australian species is only known
with certainty from two specimens, and no new
material has been examined as part of this study.
The species was redescribed by Chiong & Ng
(1994).

DISTRIBUTION

Shark's Bay, Western Australia (type locality);
North West Shelf; Prince of Wales Channel, Tor-
res Strait (Miers, 1884); Questionably from Port
Jackson (Haswell, 1880). Bathymetric range:
13m (Miers, 1884).

161

ACKNOWLEDGEMENTS

Dr Peter Ng of the Zoology Department, Na-
tional University of Singapore, is gratefully
thanked for sending us photographs of the holo-
type of C. spatulifrons, and for helpful discus-
sions on the manuscript, We are also indebted to
Paul Clark of the Natural History Museum, Lon-
don, for researching the dates of publication of
White's papers describing C. dorsalis.

LITERATURE CITED

ADAMS, A, & WHITE, A. 1848. Crustacea, In Adams,
A. (Ed.) "The zoology of the voyage of H.M.S
Samarang; under the command of Captain Sir
Edward Belcher, C.B. F.R.A.S., F.G.S., durin
the years 1843-1846". Part 1, pp. 1-32, pls 1
(Reeve, Benham. and Reeve: London).

AHMAD, M-F., S.M. SAYED, KARIM, M.S., NIAZI,
TJAWAID & M. MASIHUZ ZAMAN, 1973,
Marine fauna supplement, Records of the Zoolog-
ical Survey (Pakistan): IV (1 & 2): i-iv, 1-46,

ALCOCK, A. 1895. Materials for a Carcinological
Fauna of India, No. I. The Brachyura Oxy-
rhyneha. Journal of the Asiatic Society of Bengal
64 (P1.2, No.2): 157-291.

BANU, Q, & NURUL HUDA, K.M. 1989, New record
of Cryptopodia angulata Milne Edwards & Lucas
(Parthenopidae: Decapoda: Crustacea) from Ma-
laysian waters. Journal of the Bombay Natural
History Society 85: 646-647.

CAMPBELL, B.M, & STEPHENSON, W. 1970. The
sublittoral Brachyura (Crustacea: Decapoda) of
Moreton Bay. Memoirs of the Queensland Mu-
seum 15(4): 235-301.

CHIONG, W.L, & NG, P.K.L. 1994, The identily of
Cryptopodia spatulifrons Miers, 1879, and de-
scription nf a new species, Cryptopodia fistulosa
(Crustacea: Decapoda: Brachyura:
Parthenopidae) from Australia. Raffles Bulletin of
Zoology 42(4); 949-959,

CHHAPGAR, B.F. 1957, On the marine crabs
(Decapoda: Brachyura) of Bombay State. Part 1.
Journal of the Bombay Natural History Society
54(2): 399-439,

CHOPRA, B. 1935. Further Notes on Crustacea
Decapoda in the Indian Museum, VIII. On the
Decapod Crustacea collected by the Bengal Pilot
Service off the Mouth of the River Hughli;
Brachygnatha (Oxyrhyncha and Brachyrhyncha).
Reeords of the Indian Museum 37(4): 463-514.

DAI, A. & YANG, S. 1991. ‘Crabs of the China Seas’
(China Ocean Press: Beijing),

FABRICIUS, J.C. 1781. Species Insectorum exhibentes
conim Differentias, specificas, Synonyma au-
ctorum, Loca natalia, Metamorphosis adiectis Ob-
servationibus, Descriptionibus. i-viii, 1-552 рр.

FLIPSE, H.J. 1930. Die Decapoda Brachyura der
Siboga-Expedition. VI. Oxyrhyncha:

462

Parthenopidae. Siboga-Expeditie, Leiden Livr.
112, Monogr. 39c: 1-96. —

GORE, В.Н. & SCOTTO, L.E., 1983. Studies on deca-
pod Crustacea from the Indian River region of
Florida. xxv, Carapacial and abdominal allometry
in five species of subtropical parthenopid crabs
(Brachyura, Parthenopidae). Crustaceana 44: 1-
22

HASWELL, W.A. 1879. On the Australian Brachyura
- Oxyrhyncha. Proceedings of the Linnean Society
of New South Wales 4: 431-58.

1882, Catalogue of the Australian Stalk and Sessile-
Eyed Crustacea (Sydney): 1-323.

LAURIE, R.D. 1906, Reporton the Brachyura collected
by Professor Herdman, at Ceylon, in 1902, In
W.A. Herdman, Report to the Government of
Ceylon on the Pearl Oyster Fisheries of the Gulf
of Manaar. Part V. Supplementary Report, 40:
349-432,

MIERS, E.J. 1879. Descriptions of new or little known
species of Maioid Crustacea (Oxyrhyncha) in the
collection of the British Muscum. Annals and
Magazine of Natural History (5), 4: 1-28.

1884. Crustacea. In ‘Report on the Zoological col-
lections made in the Indo-west Pacific Ocean
during the Voyage of the H.M.S, "Alert", 1881",
London.

MILNE EDWARDS, H. & LUCAS, H. 1841. Descrip-
tion des Crustacés nouveau ou peu connus
conservés dans la collection du Museum
d'Histoire Naturelle. Archives du Muséum
d'Histoire Naturelle. Paris П: 461-73..

ORTMANN, А. 1894, Crustaceen. In К. Semon,
Zoologische Forshungreisen in Australien und
dem Malayischen Archipel. Denkschriften der
Medizinisch-Naturwissenschaftlichen Gesell-
schaft zu Jena 8(1): 1-80.

RATHBUN, M.J. 1911. Reports of the Percy Sladen
Trust Expedition to the Indian Ocean in 1905,
under the Leadership of Mr. J, Stanley Gardiner.
Vol. Ш. No. XI. Marine Brachyura. Transactions
of the. Linnaean Society. London Ser. 2. Zool.,
14(2): 191-261,

1918, Report on the spider crabs obtained by the
F.L.S. Endeavour on the coasts of Queensland,
New South Wales, Victoria, South Australia, and

MEMOIRS OF THE QUEENSLAND MUSEUM

Tasmania. Chapter 1. In *Hiological Results of
the Fishing Experiments carried out by the F.I.S.
Endeavour, 1909-14", (Commonwealth of Aus-
tralia, Department of Trade and Customs),

SAKAI, T. 1976. Crabs of Japan and the Adjacent Seas
(3 vols.) (Kodansha: Tokyo).

SCLATER, P.L. 1893. List of the dates of receipt from
the printers of the sheets of Society's
‘Proceedings’ from 1831 to 1859 inclusive, Pro-
ceedings of the Zoological Society of London.
1893: 435-440.

TAN, C.G.S. & RICHER DE FORGES, B. 1993. On
the systematics and ecology of two species of
mimetic crabs belonging to the family
Leucosiidae (Crustacea: Decapoda: Brachyura).
Raffles Bulletin of Zoology 41; 119-132.

TIRMIZI, N.M. 1980. Marine Crustacea (Decapoda and
Stomatopoda) of Pakistan. Proceedings 1st Paki-
stan Congress of Zoology: 97-114.

TIRMIZI, М.М. & KAZMI, О.В. 1983. Carcinological
studies in Pakistan, with remarks on species in
common to the Red Sea and the Meditteranean.
Bulletin of the Institute of Oceanography and
Filet (9) Marine Science in the Red Sca: 347-
380.

TIRMIZI, N.M. & KAZMI, Q.B. 1991. Marine fauna
of Pakistan: 4. Crustacea: Brachyura (Dromiacea,
Archaeobrachyura, Oxystomata, Oxyrhyncha).
Publication I, BCCI Foundation Chair, Institute of
Marine Sciences, University of Karachi. 245 pp.

WARD, T.J. & RAINER, S.F. 1988. Decapod crusta-
ceans of the North West Shelf of north-western
Australia. Australian Journal of Marine and Fresh-
water Research 39: 751-765.

WHITE, A. 1847a [April]. List of species in the collec-
tions of the British Museum. (Edward Newman:
London).

1847b [July]. Short descriptions of some new spe-
cies of Crustacea in the collection of the British
Museum. Proceedings of the Zoological Society
of London 15; 84-86,

1847c [September]. Short descriptions of some new
species of Crustacea in the collection of the Brit-
ish Museum. In, Proceedings of Learned Socie-
ties. Zoological Society. Annals and Magazine of
Natural History 20(132): 205-207.

A NEW SPIDER GENUS (ARANEAE: AMAUROBIOIDEA: AMPHINECTIDAE)
FROM THE WET TROPICS OF AUSTRALIA

VALERIE TODD DAVIES

Davies, V.T. 1995 12 01: A new spider genus (Araneae: Amaurobioidea: Amphinectidae)
from the wet tropics of Australia. Mernoirs of rhe Queenslarid Museum: 38(2):463-469.

Brisbane. ISSN 0079-8835.

The distribution of 9 species of Kababina gen.nov. in the rainforests of the wet tropics region
of northern Queensland indicates local species richness and endemism of spiders there. The
new species are К. alta, K. aquilonia, K. colemani, К. covacevichae, К, formartine, К,
inferna, K. isley, K. superna and K. yungaburra. Kababina is provisionally placed in the
Amphinectidae, [ ] Kababina, wet tropics, Amphinectidae,

Valerie Tadd Davies, Queensland Museum, PO Box 3300, South Brisbane, Queensland

4101, Australia; 21 June 1995.

Many Australian amaurobioid spiders are
recognised by their unexceptional appearance;
straw-coloured carapace (often with two darker
longitudinal bands), abdominal pattern of light
spots in a vague chevron pattern, 3 claws and a
cribellum. This new genus belongs in a large
group of undescribed species with the following
further characters; cephalothorax highest in the
foveal region, posterior eye row straight or
slightly recurved; anterior median eyes reduced.
Two retromarginal arid two promarginal chelic-
eral teeth; prolateral filamentous seta at base of
fang, longer than other setae, Labium about as
long as wide; sternum slightly longer than wide,
pointed posteriorly. Legs 4123; notched trochan-
ters. Feathery hairs on legs. Tarsal trichobothria
increasing in length distally; bothrium collari-
form. Tarsal organ slit-like, broadening distally.
Epigynum with medial fossa (opening) wider
than long; spermathecae posterior or lateral to
fossa. Male palp with rounded tegulum; course of
sperm duct showing clearly. Membraneous con-
ductor; embolus with or without basal embolic
apophyses, arising ventrally; without median
apophysis. Tibial apophysis branched. Cribellum
(two fields) present or absent in fernales, absent
in males; proximal calamistrum with one row of
setae. Colulus large when present. Anterior spin-
nerets largest; two major ampullate spigots of
unequal size on antcrior spinnercts.

MATERIALS AND METHODS

The spiders have been collected from
rainforests in the Wet Tropics region of northeast-
ern Queensland between latitudes 15'48'-
17'16'S. Collection methods include litter
sieving followed hy heat exlraction in funnels,

pitfall collection, pyrethrum spraying of fallen
logs and hand collecting from under logs and at
night.

Notation of spines follows Platnick & Shadab
(1975). All material is lodged in the Queensland
Museum (QM). Measurements are in
millimetres.

ABBREVIATIONS USED

CL, carapace length; CW, carapace width; AL,
abdomen length; AW, abdomen width. Eyes:
AME, anterior median; ALE, anterior lateral;
PME, posterior median; PLE, posterior lateral,
Spinnerets: ALS, anterior; PMS, median; PLS,
posterior; 1, left; г, nght.

Collectors: DC, D. Cook; EWQM, Earth-
watch/Queensland Museum; GBM, G.B. Mon-
teith; СТ, G. Thompson, HJ, Н. Janetzki; JC, J,
Covacevich; NC, N. Clyde Coleman; RJR, R J.
Raven; VED, V.E. Davies.

SYSTEMATICS
Kababina gen. nov,

TYPE SPECIES
К. айа sp. ney,

DIAGNOSIS

Three-clawed cribellate. Viewed from top,
eye-rows slightly recurved; AME reduced
(Fig.1B). Stemum pointed posteriorly; endite
longer than wide (Fig. 1С). Feathery hairs present,
Small medial epigynal fossa. Tegulum with pro-
lateral ridge; semicircular embolus and conduc-
tor, arising ventro-retrolaterally on tegulum;
without median apophysis. Male palpal tibia not
much longer than wide; single long seta prolateral

464

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. I. A-l, Kababina alta. A, 9 cephalothorax, lateral. B, ? eyes and chelicerac. C, cephalothorax, ventral. D,
abdomen, E-G, epigynum, ventral, ventral (cleared), dorsal. H, 1, д palp.

to ventral portion of tibial apophysis (Fig. 11).
Paracribellar spigots present on 2 PMS.

DESCRIPTION

Carapace straw-coloured to brown. Abdomen
with pattern of pale spots (Fig.ID). Carapace
highest in foveal region (Fig. lA). Clypeus x 2
AME. Labium slightly longer than wide; endite
almost twice as long as wide. Chelicera with two
retromarginal and two promarginal teeth. Pro-
lateral seta at base of fang longer than other setae.
Legs 1423; trochanters notched. Row of tarsal
trichobothria increasing in length distally; bothr-
ium collariform (Fig. 5A); trichobothria on
cymbium. Tarsal organ (Fig. 5B) slit-like broad-
ening distally. Two cribellar fields; proximal
calamistrum іп 9. Large colulus in d. Anterior
spinnerets largest. Epigynum with wide insemi-
nation ducts running forwards, turning back at
swollen glandular area, then narrowing to enter
pear-shaped spermathecae, d palp with branched
tibial apophysis, a blunt ventral portion and two
long retrolateral branches. Course of sperm duct
visible postero-retrolaterally. Tracheal system
simple.

REMARKS

Forster & Wilton (1973) placed most of New
Zealand's amaurobioids which have a medial
epigynal fossa in the Amphinectidae.
Amphinecta, itself is characterised by having un-
reduced AME, 8-9 promarginal teeth on the che-
licera, preening combs on metatarsi III and IV, a
median apophysis on d palp, and by lacking
feathery hairs, None of these characters is found
in Kababina. However Maniho, placed in the
Amphinectidae, has reduced AME, feathery hairs
and is without preening combs and also shares
with Kababina a general similarity of structure in
epigyna and d palps, though Kababina lacks
epigynal teeth and median apophysis.

Kababina is provisionally placed im the
Amphinecudae.

ETYMOLOGY

Kababina - the Aboriginal word for rainforest,
Kuku-Yalanji language, Bloomfield area of
northeastern Queensland.

NEW SPIDER GENUS

TABLE 1. Palp and leg measurements of Kababina alta
sp. nov.

Femur | Patella |Tibia [Metat. |Tarsus | TOTAL |
apog юз fos |. оу |7 |

Kababina alta sp.nov.
(Figs 1, 5)

MATERIAL EXAMINED

HOLOTYPE: QMS22807 2, Mt Bellenden Ker Range,
17°16'S, 145°51°E, 1054m, under logs, 17.x-
5.xi.1981, EWQM .

PARATYPES: QMS22808 ð, same data as holotype, ;
QMS22809 9; QMS22812 9 epigynum. QMS22810
9, South Peak summit Mt Bartle Frere, 1620m, 6-
8.x1,1981, EWQM.

DIAGNOSIS
Epigynal fossa much wider than long. Male
embolus with notch; anterior retrolateral branch

of tibial apophysis lightly sclerotised and about
half length of posterior branch.

DESCRIPTION

Female: CL 2.3, CW 1.6, AL 2.4, AW 1,5,
Ratio of AME:ALE:PME:PLE is 4:6:6:6. Leg
measurements (Table 1). Notation of spines.
Femora: I, D110, P011, R001; II, D110, P011,
R011; m, D110, P001, R011; IV, 0110, POOL,
R001. Patellae: T, D101; H, D101; m, D101; IV,
D101. Tibiae: I, PO11, V120; II, P011, V220,
R001; m, D101, POLI, V110, К011; IV, DIMI,
P011, V111, RO11. Metatarsi: several spines with
whorl of 4 distally. Epigynum (Fig. IE-G) with
elliptical fossa, ratio of width: length, 1:0.2. Spin-
nerets (Fig. 5C-G): ALS with two major ampull-
ate spigots, one larger than the other; about 24
piriform spigots. PMS with large anterior spigot
(?cylindrical) and large posterior spigot (?minor
ampullate), several paracribellar spigots and
some aciniform spigots. PLS with several spigots,
one larger than rest (?cylindrical).

Females varied in length, 4.1-5.0.

Male: CL 2.0, CW 1.6, AL 2.0, AW 1.2. Ratio
of AME:ALE:PME:PLE is 4:6:7:6, Notation of
Spines. Femora: I, 0011, PO11, КОО1; П, D111,
P011, ROI I; m, D111, P011, R001; IV, D110,
P001, R101, Patellae: I, D100; II, D101; m,
D101; IV, 0101. Tibiae: I, PO11, V220, КОТІ; И,
DOOL, POLI. V220, RO11; m, D101, P011, V110,

FIG. 2. A-C, Kababina inferna. D, K. isley. E, К. yangaburra. ^, B, epigynum, ventral, dorsal. C-E, palps,

466

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. A-D, Kababina covacevichae; E,F, К. colemani; О.Н, К. formartine; IK, К. superna. A-C, I-K, epigyna,
ventral, ventral (cleared), dorsal; D, d palp; E,F;G,H, epigyna, ventral, dorsal.

R011; IV, D101, P111, V111, ROII. Metatarsi:
several spines, whorl of 4-5 distally.

Male palp (Fig. IH,I): Long femur, broad tibia;
ratio of tibial length: width is 1:0.7. Tibial apoph-
ysis with blunt ventral branch and two retrolateral
branches. The anterior branch short and lightly
sclerotised, the posterior long, curved and heavily
sclerotised. Embolus with sub-distal notch.

ETYMOLOGY
Latin, altus, high, referring to the altitude.

Kababina inferna sp. nov.
(Fig. 2 A-C)

MATERIAL EXAMINED

HOLOTYPE: QMS22813 2, Mt Bellender Ker, 500m,
litter, 1-7 xi, 1981, EWQM.

PARATYPES: QMS22814 d, same data as holotype ;
QMS22815 49 548.

DIAGNOSIS

Epigynal fossa much wider than long, ratio
1:015. Embolus notched; anterior retrolateral
branch of tibial apophysis sclerotised, almost as
long as posterior.

DESCRIPTION

Female: CL 1.9, CW 1.3, AL 2.2, AW 1.5,
Colour, eye size and leg spination similar to К.
alta. Epigynum (Fig. 2 A,B) Females varied in
length, 3.5-4.2.

Male: CL 1.9, CW 1.4, AL 1.9, AW 1.2; 5 palp
(Fig. 2 C), broad tibia, ratio length: width. 1:0.9.
Tibial apophysis with blunt ventral portion and 2
sclerotised retroventral branches; the anterior
branch bent back before ending bluntly, the pos-
terior longer, curving and pointed. Embolus with
sub-distal notch. Large colulus; ALS with 2
major ampullate spigots, one larger than the
other, 18-19 piriform spigots. PMS with a few
spigots, undefined. PLS with several spigots with
long thin fusules (aciniform), no large spigots
seen. Males varied in length, 3.2-3.8.

ETYMOLOGY
Latin, infernus, lower, referring to the altitude.

Kababina isley sp. nov.
(Fig. 2D)

MATERIAL EXAMINED

HOLOTYPE: QMS22824 d , Upper Isley Ck, 17° 03'S,
145° 41°E, 750m; 29-30.xi. 1993. DC, GBM, HJ.
PARATYPE: QMS22825 d , same data.

NEW SPIDER GENUS

FIG. 4. A,B, Kababina aquilonia, б palp. A, holotype;
В, (Mt Spurgeon).

DIAGNOSIS
Notched embolus. Anterior retrolateral branch
of tibial apophysis bifurcate.

DESCRIPTION

Male: C1 1.8, CW 1.3, AL 1.7, AW 1.0. Colour,
eye size, leg spination similar to К. alta, Male
palp (Fig. 2D), embolus notched, Tibia length:
width, 1.08; anterior retrolateral branch of tibial
apophysis bifurcate; posterior narrows abruptly,
pointed. Paratype б same size. Female unknown.

ETYMOLOGY
For Isley Creek.

Kababina yungaburra sp. nov.
(Fig. 2E)

MATERIAL EXAMINED

HOLOTYPE: QMS22823 d, Yungaburra, 17°16'S,
145° 35'E, 700m, pitfall trap, 8-14.xii. 1988, GBM,
GT.

DIAGNOSIS
Notched embolus. Anterior retrolateral branch
of tibial apophysis pointed, posterior bifurcate.

467

DESCRIPTION

Male: CL 1.9, CW 1.5, AL 1.8, AW 1.2. Male
palp (Fig. 2E). Embolus notched. Tibia length:
width, 1:0.8. Tibial apophysis with blunt ventral
portion, 2 sclerotised retrolateral branches, ante-
rior pointed, posterior narrow, slightly longer,
forked at tip. Female unknown.

ETYMOLOGY
For the town Yungaburra.

Kababina covacevichae sp. nov,
(Fig. 3 A-D)

MATERIAL EXAMINED

HoLoryPE: QMS22816 9, Shiptons Flat, 15° 48'S,

145* 45'Е, under rotting log, 16-21.xi. 1975, VED, JC.
PARATYPE: QMS22817 д, same data.

DIAGNOSIS

Epigynal fossa wider than long, 1:0.4; glandu-
lar area of insemination duct aligned parallel to
body axis. Embolus unnotched.

DESCRIPTION

Female: CL 2.0, CW 1.2, AL 2.6, AW 1.8.
Coloration, eyes, leg spination similar to K. alta.
Small epigynal fossa (Fig.3A-C).

Male: CL 2.0, CW 1.5, AL 2.0, AW 1.3. Male
palp (Fig. 3D), embolus unnotched. Broad tibia,
length: width, 1:0.9. Tibial apophysis with blunt
ventral portion and 2 sclerotised retrolateral
branches; anterior ending bluntly, posterior
curved, pointed.

ETYMOLOGY
For Jeanette Covacevich.

Kababina superna sp. nov.
(Fig. 3 I-K)

MATERIAL EXAMINED
HOLOTYPE: QMS22818 9, Mt Lewis, 16" 34 S, 145°
16°E, 1 100m, 7.xi. 1975, VED, RJR.

DIAGNOSIS

Epigynal fossa wider than long, 1:0.3; glandu-
lar area of insemination duct aligned transversely
to axis of body.

DESCRIPTION

Female: CL 2.2, CW 1.5, AL 2.7, AW 1.6.
Epigynal fossa width: length, 1:0.3 (Fig. 3 I-K)
Male unknown.

468 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. A-G, 9 Kababina alta. A, bothrium. B, tarsal organ. C, spinneret field. D, ALS (1). E, PLS (r). F, PMS
(r). G, PMS, paracribellar spigots.

NEW SPIDER GENUS

ETYMOLOGY
Latin- supernus - high, referring to the altitude,

Kababina colemani sp. nov.

(Fig. 3 E,F)

MATERIAL EXAMINED
HOLOTYPE: QMS22821 9, Black Mountain, 16° 39'S,
145" 29' E, 30.1. 1972, NC.

DIAGNOSIS
Glandular area of insemination duct aligned
parallel to axis of body.

DESCRIPTION

Female: CL 1.8, CW 1.4, AL 2.2, AW 1.3.
Epigynal fossa width: length, 1:0.3 (Fig. 3E,F).
Male unknown.

ETYMOLOGY
For the late Clyde Coleman, a north Queens-
land naturalist.

Kababina formatine sp. nov.

(Fig. 3 G,H)

MATERIAL EXAMINED

HOLOTYPE: QMS22822 9, Mt Formartine South, 16°
44'S, 145° 3T'E, 700m, pitfall trap, 23-24.xi. 1990,
GBM, GT.

DIAGNOSIS
Epigynal fossa a little wider than long.

DESCRIPTION

Female: CL 2.1, CW 1.5, AL 2.4, AW 1.7.
Epigynal fossa almost circular, width: length,
1:0.8 (Fig. 3G,H). Male unkown.

ETYMOLOGY
For Mt. Formartine.

Kababina aquilonia sp .nov.
(Fig. 4A, B)

MATERIAL EXAMINED

HOLOTYPE: QMS22819 d, Spear Creek, nr Mt
Molloy, 16° 41'S, 145" 20' E, litter, 3-10.xi. 1975, RJR,
VED.

PARATYPE: (MS 22826 d , same data.
OTHER MATERIAL: QMS22820 5, Mt Spurgeon, 16°
28'S, 145° ITE, 1100m, pitfall trap, 11-21.х.1991,

DIAGNOSIS
Unnotched embolus.

469

DESCRIPTION

Male: CL 2.3. CW 1.7, AL2.3, AW 1.5. д palp
(Fig. 4A). Embolus unnotched. Tibial length:
width, 1:0.9. Tibial apophysis with blunt ventral
portion, anterior retrolateral branch lightly
sclerotised ending bluntly; posterior longer, ta-
pering to point.

Paratype 5 smaller, 3.8, Male from Mt Spur-
geon larger, (5.6) and may not be conspecific; the
posterior retrolateral branch of tibial apophysis
(Fig. 4B) is less tapered than the holotype.

Female unknown.

REMARKS

Males of the species can be divided into those
which have a small sub-distal notch (K. alta, K.
inferna, K. isley and K. yungaburra) and those
without the notch, the more northerly species (K.
covacevichae, K. aquilonia). Similarly females
divide on the shape of the epigynal fossa; it is
much wider than long in the more southern spe-
cies (K. alta, K. inferna) and only a little wider
than long in the remainder. The dividing line
appears to be at about the latitude of Cairns
(16°55'8).

ETYMOLOGY
Latin, aquilonius, northern.

ACKNOWLEDGEMENTS

I thank the Council of the ABRS for funding
rainforest surveys during which some of this ma-
terial was collected and for financial support of
illustrator, Christine Lambkin. I am indebted to
the Wet Tropics Management Authority for fund-
ing SEM technician, Kylie Stumkat. I acknowl-
edge help given by "Earthwatch" and the Council
for Field Research, Boston, Mass. USA for sup-
porting the expedition to Mt Bellenden Ker. Fi-
nally thanks to the staff of the Queensland
Museum, particularly Caroline Davis for her help
in preparation of this paper.

LITERATURE CITED

FORSTER, R.R. & WILTON, C.L. 1973, The spiders
of New Zealand. Part IV. Otago Museum Bullenn
4: 1-184.

PLATNICK, N.I. & SHADAB, M.U. 1975. A revision
of the spider genus Gnaphosa (Araneae:
Gnaphosidae) in America. Bulletin of the Ameri-
can Museum of Natural History 155; 1-66,

476

BRUSH-FOOTED TRAPDOOK (OZICRYPTA SP.)
BURROWS AND GECKOS AT WANDO VALE, NEQ.
Memoirs af the Queensland Museum 38(2):470. 1995;- Main
(1993) saw burrow openings reinforced and raised above the
surface and tightly fitting, thick plug-doors primarily as adap-
tations against flooding. Raven (1994) discounts this as а
major force in the evolution of barychelid burrow structure.

Ozicrypta sp. (Mygalomorphae: Barychelidae) were dug
from their burrows and the structures photographed and drawn
(Fig. 1) and rejectamenra collected. QMS30177 was from the
steep, stony loarn bank of a small, dry gully, 1 m from gully
floor; the others (QMS30179, QMS30182) were under logs in
a broad, shallow gully floor in sandy loam (Fig. 2). All were
suhject to short-duration flooding of unknown frequency.

Main burrows were unlined, vertical and open, openings
unadorned and flush with the surface; hurrow diameters were
с, 15mm; depths 100- 150mm. Spiders were each found im the
bottom third of the burrow in a parchment-silk-lined side-
chamber with the outward-opening, side-hinged door a sitk-
lined plug of earth. The spiders held the doors with their front
legs. The burrow of QMS30179 (Fig. 1B) had three sides
chambers (i, near opening, unlined, with grasshopper remains.
it, (11mm diameter, 25mm long) also near opening, silk-lined
with damaged door; iti, (18mm diameter, 28mm long! silk-
lined, lid intact, occupied, in bottom third of burrow). These
burrows differ from the lidded and silk-lined burrows of many
Ozicrypta which are often buried in litter (Raven, 1994) but
most resemble those of Zaphorame covacevichae, Trittame
and some /diommata (Raven, 1994),

A Burrow-plug gecko, Diplodacrylus conspicillatus. was
found at the bottom of the burrow of QMS30177 (Fig. ТА),
below the trapdoor with tail curled above its head plugging
the burrow, This posture is described in Ehmann (1992). Its
use of spider burrows as a daytime shelter has been reported
(Pianka & Pianka, 1976; Wilson & Knowles, 1988; Greer,
1989; Ehmann, 1992). None of these authors mention the
spider being present, Although Greer (1989) believed all
geckos to be opportunistic arthropod feeders, Pianka & Pianka
(1976) cite termites as the pecko's sole diet and record another
long-bodied termite specialist, Rhynchoedura ornata, also
sheltering in spider burrows. No gecko remains are known in
The rejectamenta of Ozierypia spp. An association between
spider and gecko is possible though the degree of mutual
benefit is unknown. D, conspicillatus leaves the burrow in the
first three hours of the evening to forage for termites (Planka
& Pianka, 1976). The position of the side-chamber door above

FIG. 1. Bucrows of (4) OMS30177; (B) QMS30179
(1-11, see text),

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2, Locality of QMS30179,

the boltom would allow the spider access to the surface when
the gecko is in. Nightly gecko traffic might also explain the
absence of outer tra rs and silk in the main burrows. It is
unknown whether Ozicrypta awaits prey at the mouth of its
underground side-chamber (Main's "pitfall capture"), at the
burrow opening or leaves the burrow entirely. As the grass-
hopper remains are the spiders rejectementa, the spider must
hunt at or beyond the burrow opening. Both size and behavi-
our of grasshoppers makes them unlikely candidates for pitfall
capture.

Material Examined

Ozierypta sp.

QMS30177 penultimate d, Wando Vale sin, MEQ,
19*27.22'8, 144'45.67' E, open bloodwood forest on hills,
17.1X.1995, P.Lawless, A.Cook, C McHenry, QMS30179
immature, Wando Vale stn, 19°32.96'S, 144"40.03'E, open
woodland, Narrow-leaf Ironbark, sandy loam, 11, V11.1995,
P.Lawless, A.Cook, N.Cumilleri. QMS30182 penultimate d ,
same data. Orthopteroid rejectamenta:- pronotum: 7.5mm
long, 6.5mm high; forewings: 28mm long, Smm high; tibia:
12mm long and femur (part).

Literature Cited

Еһтапа, Н. 1992, Encyclopedia of Australian Animals: Rep-
tiles (Collins, Angus & Robertson: Sydney).

Greer, A.E. 1989, The biology and evolution of Australian
lizards. (Surrey Beatty & Sons: Sydney),

Main, B. Y, 1993, From flood avoidance to foraging: adaptive
shifts in trapdoor spider behaviour. Memoirs of the
Queensland Museum 33(2): 599-606.

Pianka, Е.К, & Pianka, H.D. 1976. Comparative ecology of
twelve species of nocturnal lizards (Gekkonidae) in the
Western Australian desert, Copeia 1976(1); 125-142.

Raven, R.J. 1994, Mygalomorph spiders of the Barychelidae
in Australia and the western Pacific. Memoirs of the
Queensland Museum 35(2): 291-706.

Wilson, S.K. & Knowles, D. 1988, Australia's reptiles. (Col-
lins; Sydney),

Philip B. Lawless, Queensland Museum, P.O,Box 3300, Sowth
Brisbane, Queensland, 4101, Australia; October, 1995.

HYBRIDIZATION BETWEEN THE GREAT AND SPOTTED BOWERBIRD
CHLAMYDERA NUCHALIS AND C. MACULATA : AN AUTHENTICATED HYBRID
BOWERBIRD (PTILONORHYNCHIDAE).

CLIFFORD B. FRITH AND DAWN W. FRITH

Frith, С.В. & Frith, D.W. 1995 12 01: Hybndization between the Great and Spotted
Bowerbird Chlamydera nuchalis and C, maculata: an puthenticated hybrid bowerbird
(Ptilonorhynchidac). Memoirs af the Queensland Museum 38(2): 471-476. Brisbane. ISSN
0079-8835,

Evidence of hybridization in the wild between the Grent and Spotted Bowerbirds Chlamydera
nuchalis and С. maculata (Prilonorhynchidac) is described and illustrated from a Queensland
Museum specimen. Another Queensland Museum specimen possibly represents such a
hybrid and is also detailed. { ] Bowerbirds, hybrid.

Clifford B. Frith & Dawn W. Frith, ‘Prionodura’, Р.О, Box 581, Malanda, Queensland 4885,
Australia; | Augusr 1995.

The Australo-Papuan Ptilonorhynchidae (bow- Two Australian monogamous bowerbirds, the
erhirds) consists of 19 species (Sibley & Monroe, Green Ailuroedus crassirostri« and Spotted Cal-
1990; Frith, 1991; Frith et al., 1995а) and was bird A, melanotis, were reported lo have once
long considered closely related to birds of para- hybridized in captivity (Iredale,1950), bul no
dise, римин, (Gould, 1869; Ѕћагре,1&91- specimen apparently exists to confirm the event,

98: 01,1873; Gilliard, 1969: Cooper & In the New Guinea genus Amblyornis, the |
mde re jn ate 1976). Recent studies’ 9Sfitakei4/ sübalakis ahid/MaépreonrwoBiwer- icti m ипи

n p that the two groups are, how) іта: элә гё sbaldecabut угш tbe Owen imd иЗ ume

" Het disparate among раѕкег- "BtanleycRanec.-SéhaddeaibiMaKcag (193) rc ported ^" Ausi

en thes (Eti 35 “Erith & Frith, 1990, 1994 grege Sh ee жЕр e nione Ужа D 25064. Ип.

cies ДЕБ ШЕ; Sibley & Monroe, 1990; Marie ter ih Ee. bet he liy ul LC

Schede © Kasmiesskaostaal. 1993; Christidis & Schodde > the iN омет Suh dey Messe -

mvincraz, L992) -ARthey | Mere supposedly related, hee 19 ус zd. mort o
о

ие ray 4 "

Ae Wer epee Ae

etri wis

d was diedywamausylitiwakanticipated that hybrids migh ШУ. A um эз, e Р um Amblyarnix пурӣ
wilt the edoerurPbalwiiendowerbird species because d bagi W
psali oceny besween!polygynous taxa withinz g qu
of svmpstrpareidisseidaest Mayr, 1941, 1962; Fuller, 1979). Д }
4 Bi ve*tmid&everalybrid bowerbirds have been reported, * * ds in cenital east
cn heir qeurmioüeamhentiéated. A supposed hybrid be-" hu di ipn Quesne! mrg к
!. tween the Satin Prilonorhynchus violuceus and erat! po p violacei aad (тия у
Regent Bowerbird Sericulus chrysocephalus was" ent Bower e cul: chryaucenhus har wes
yr nie RPresented by à single adult male in the irides ү үлү py a quim md m i the inde
cent blue plumage of the former species but with “P ~= © Раде с pores Est «йл
extensive SU wing patches and tipping to (d or Rer pw ер. panes x oer up
saray feathers a "y expressing Regent Bower ir a кар 3 y Натат b "e iie i “arm (0

ange nique specimen, known as pula
cour ans ы 15 ЕНА EET Ts is tics o the

Ter ea пол Верес заан) MESE found wA
n war сейген ЕРТ es. 1867; Gould, 1869; Elliot, sso dE x Jr p ic Ca Te

пиез] Bas Sharpes 1891-98; Iredale, 1950) was the! interacted. (sec :

TET roesubjectot eantrosersy concerning its authenticity" E E gmi- 190552 Habit
3 cacal сакои, 1966»; and its subsequent disappear- gatta by Acacias и
y ботилае Icàvessodni г doubt about its origins (Mar-77* әд к ОШ: г Fs an unen
X bees ghalik M52; Gilliard, 1969). Hybridization must Ass nee j Larissa l'unciotar
уй: agvs leceencededreetpossihility, however. betweent® SE Осе ur cr E located. Сту
ес ies vdyeteqeedtédoselg related (Frith & Frith, 1993) "6 T uda Ж toas And bae of both
ELLO ортара made d et al., 19955). phrfingzapirs mode

we

hk

E ара, Lita) Tix
search foran are

лү =

ar - Soule

472 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. A, anadult male Great Bowerbird (centre) displays his crown to the male hybrid Great x Spotted Bowerbird
here described (rear) as an adult male Spotted Bowerbird (foreground) looks on. B, dorsal & C, ventral plumage
detail of the live hybrid Great x Spotted Bowerbird described here, banded with ABBBS band # 081 71976 when
caught on 11 September 1991. D, ventral & E,F, dorsal appearance of (1. to r.) two Spotted Bowerbird museum
specimens QMO6428 & QMO27263, the hybrid Great x Spotted Bowerbird QMO30059, and three Great
Bowerbirds QMO30058, QMO27527 & QMO?27576.

HYBRID BOWERBIRD

TABLE 1. Measurements of a live hybrid Great x Spotted Bowerbird and those of live
bowerbirds caught at bowers in the same area of north Queensland with the appearance
of typical Great and Spotted Bowerbirds. Measurements in mm.

[Weight [Wing [Tai — [Tas [Bm — пы |
m —

Mem pos п — [me — faa fass — lens |
Range — |(187.219) |(165-175) |(121-134) |(45.1-48:7) |(37.3-39.8) (66.1-68.4)_
ONES CREE es аер

(нуна Бома? [m — Tier — us — lass [srs les |
ГНИ

Docs
Range Joie [aos anos; [oiii p eene
[n= ie Ti СЫН

Great Bowerbird

Spotted Bowerbird

зі ја ja

1994) that is now specimen QMO30059.
Data are from 31 birds at the Bruslee study area.

Bird skin specimens examined are in the
CSIRO Australian National Wildlife Collection
(ANWC) and the Australian (AM), National Mu-
scum of Victoria (MV), Queensland (QM) and
South. Australian (SAM) museums. Capitilized
descriptive words for colours are those of Smithe
(1975) unless we consider them to be inappropri-
ate, in which case we use uncapitalized words.
Colour reference numbers (in parenthesis) are
those of Smithe (1975) closest to that being de-
scribed but not necessarily of the precise colour
observed. Colours considered standard for Great
Bowerbirds in the part of tropical Queensland
concerned (18-23'S and 138-148°E) are taken
from a subadult male (QMO30058) collected at
the same location as the hybrid, and from adult
specimens QMO27375 and QMO27576. Speci-
mens of Spotted Bowerbirds used as standards for
comparison with the Great and hybrid specimens
are QMO6428, QMO10669,0MO10670 and
QMO27263, all in richly-coloured and largely
unworn plumage (Fig. ТА).

RESULTS

At one Great Bowerbird bower (bower 'A" of
Егиһ et al., 1995b), we noted Spotted Bowerbirds
also visiting, displaying and decorating. On 23
September 1989 and during late September 1990
CBF watched and photographed a bird, consid-
ered to be the same individual, that looked the
size of a Great Bowerbird but which had a few
dorsal feathers the rich cinnamon colour of Spot-
ted Bowerbird pl and heavy ventral mark-
ing and barring. This individual associated with

А , Data from three birds at the Bruslee study area and three from Townsville. -
? Data are the average from the two examinations of the live bird (11 Sept 1991 & | Sept

473

both the bower-
owning Great and
several Spotted
Bowerbirds at the
bower to watch and
follow them as they
| decorated or dis-
| played (Fig. 1A).

On 11 September
199] a bird identi-
fied as the same in-
dividual was mist
netted at the same
| Great Bowerbird
bower, photo-
graphed (Fig. 1B,C}
and measured (Ta-
bles 1 & 2). At that
time we thought it
to be hybrid but as
the possibility it was an abnormally plumaged
immature Great Bowerbird remained, we banded
and released it (ABBBS band 081 71976). Its
pink crest was then one third developed, its mouth
yellow and its gape pink-yellow. On 1 September
1994 this individual was again caught at the same
bower. AI this time the bird, now known to be at
least four years old, was fully crested with a
flesh-yellow mouth and plumage characters of
both Great and Spotted Bowerbirds. As we now
had no doubt the bird was hybrid, we collected
and prepared il as a skin for the Queensland
Museum (QMO30059). It had a fully ossified
skull and enlarged (10 x 7mm) gonads.

17

DESCRIPTION OF THE GREAT X SPOTTED HY-
BRID BOWERBIRD SPECIMEN QMO30059

The hybrid differs ventrally from adult-plum-
aged Great Bowerbirds in not having the throat,
breast and flanks uniform buff-grey (Light Drab
119C) but a darker colour, closer to Grayish Horn
(91), because of the influence of heavy mottling
and spotting that develops into barring on the
sides of the chest and continues down the flanks
and onto the thighs. A few of the most heavily
barred feathers to either side of the chest are
washed with pale Cinnamon (39) which is
stronger on the concealed, basal, part of them.
The central abdomen and belly is cream-yellow
(54) as in Greats but it extends further up toward
the chest as in Spotted Bowerbirds, Undertail
coverts and undertail are as in typical Greats but
рее differ little from those of Spotted Bower-

irds,

474

TABLE 2, Measurements of Greal and Spotted Bowerbird skin specimens from north
Queensland localities most near the Bruslee study area in Australian collections and
those of a hybrid individual from these parent species. Measurments in mm.

Oo ————

| Great Bowerbird

E m

[омозтзуо

Is. otted Bowerbird®

! Specimens ате from: 25 km S of Townsville (CSIRO 415093, 415109, 415134),
Guibert River, 74km W of Georgetown (CSIRO 374922, 416269, 41627 9), near back

Esmerelda Homestead (QMO275755, 275765, 275908 [possible hybrid - see text]),
18km 5 of Clermont (CSIRO 431794), 3]km N of Lynd Junction (CSIRO 43141 2)
and 19km NE of Inverliegh Homestead (QMO27515 8).

? Data arc averaged from the 11 Sept 1991 and | Sept 1994 examinations of the live bird

that is now specimen QMO30059,

3 Specimens are from: c.100km NW of Clermont (ОМО 10669, 106702), 67km № nf
Creek crossing, Boulia-Mt. Isa High-
way {0М027263 8), Wernading Homestead (QMO273739? ), near Mt. Isa (CSIRO
66212, 417569 41757 9, SAMB 20505 7), Collulam Homestead ((QMO27303 d ) and

Hughenden (CSIRO 43157 8,43158 $), Waverl

SW Barcaldine (CSIRO 373214).

Unlike adult, subadult and immature Greal
Bowerbird plumages the hybrid's malar, lore and
ear coverts are distinctly mottled with the same
Grayish Hom (91) às the throat and breast. Unlike
Greats but as typical of Spotted Bowerbirds there
is an indistinct but clearly visihle pale off-whitish
malar stripe (only just apparent in Fig. 1C).

The hybrid is dorsally different from typical
Greats of any age. The wings are generally sim-
ilar to those of an adult Great except that the
extensive pale tips to primary covert feathers are
not off-white or whitish but are washed with pale
Cinnamon (123A) as in, but paler than, Spotted
Bowerbirds. The crown feathering is intermedi-
ate between that of the two species. The feathers
are pot simply tipped with pale Buff (124), or
whitish as in Greats, or streaked with rich Raw
Sienna (136) conspicuously edged with dark
grey-brown (119A) as in Spotteds, but are cen-
trally spotted Pale Horn (92) washed to varying
intensity, but stronger nearer their bases, with
pale (39) to darker (123A) Cinnamon (Fig 1D-F).
Crest colour in the hybrid is Rose pink (9) but
may be Magenta (2) or be shot with Purple (1) in
some light as 15 true of both putative parent spe-
cies, Some forwardmost and hindmost crest
feathers are upped silvery-white. Posterior to the

[Mean р
— i ies

[ныне |ro — ies һм Мв [з lesa |
E m e юк Hz 1 _]

MEMOIRS OF THE QUEENSLAND MUSEUM

crest is a discrete
area of con-
trastingly uniform
brownish Glaucous

(79),
Mantle, back and
терю ип из мв юз rump feathers show
characters of both
putative parents but
a je шз э | are more like those
of Spotteds in being

more blackish, al-
most black on the
central back, unlike

the datk grey-
p= [ри _. 12 ha- Jin o] brown (221) of

Great Bowerbird
feathers,
These feathers on
the hybrid do have
the broad pale-grey
tipping that gives
the scalloped ap-
pearance of the
Great's dorsal
Plumage but this is
rendered inconspic-
uous by large cen-
tral feather spotting
of the Spotted Bowerbird type of marking. These
Spots, not as large or symetrical as in the Spotted
but nothing like Great plumage, vary in colour
from a pale almost creamy off-white (92) to, in
few places, the Cinnamon (123A) and rich Mi-
kado Brown (121C) typical of Spotted Bower-
birds (Fig. ID-F). Table 3 comparatively
summarizes characters.

Primary feather colouring and marking is more
like that of the Great than the more brownish and
extensively paler-marked Spotted Bowerbird.
Rump feathers have a conspicuous dark sub-
terminal band typical of Spotteds but not of Great
Bowerbirds, Upper tail colour and markings are
not discernable from those of Great Bowerbirds,
Adult Greats have a clear yellow mouth and
Spotteds a fleshy one, the mouth of the hybnd
being intermediate between them. Measurements
of the hybrid specimen are compared with those
of live (Table 1) and skin Great and Spotted
Bowerhirds (Table 2). |

Mitochondrial DNA study of material from the
trunk of the hybrid specimen confirms that the
female parent was a Great Bowerbird and the
father, therefore, a Spotted Bowerbird

HYBRID BOWERBIRD

475

TABLE 3. Some characters of adult Great and Spotted Bowerbirds in central east Queensland compared with

those of a hybrid between the two of them

| Throat, breast & flanks

| Cream-yellow (54) but
Abdomen, belly & chest healer LI) properat

Pale malar stripe Absent |

Lore & ear coverts

Crown feathering

(Christidis, pers. comm. ; Christidis, Frith & Firth,
unpubl, data).

DISCUSSION

Characters in the hybnd peculiar to the Spotted
Bowerbird are the pale malar stripe, heavily mot-
tled and barred breast and flanks, sienna-streaked
crown and spotted back feathers. Notwithstand-
ing these unequivocal traits, the hybrid is far
larger in all measurements than any Spotted,
being the size of a small Great Bowerbird (Table
1). Peculiar to the Great is dorsal spotting paler
than the rich cinnamon typical of Spotteds. The
hybrid is markedly smaller than most male Great
Bowerbirds, particularly in weight, (Tables 1 &
2). Its weight on 11 September was 168g and on
1 September 1994 172g. Thus its average weight
of 170g is lighter than that of all six Greats
trapped at its specific location (Table 1) and that
of 12 taken (immediately after collected) over
that area of north Queensland (Table 2). Its wing
is smaller than most Great Bowerbirds and its tail
more so (Tables 1 & 2). From its plumage at the
time we are reasonably confident the hybrid bird
was at least two, probably three, years old when
banded on 11 September 1991. Thus it was at
least five years old when collected. These facts
and observations leading to the collection of
specimen QMO30059 clearly indicate it is the
product of hybridization between Great and Spot-
ted Bowerbirds.

[== |с. „ми. шы |

Uniform light drab (119C) |Greyish horn (91) Greaney gow w (54)
Uniform light drab (119C) | Mottled Greyish horn (91) a, | pale cinnamon

|Paleprimarycoverttips — |Off-white — |Very pale cinnamon (123A) T

Central E em
cns ME 8 Tipped pale bu (124) 10 |) tashe P
Ec ст er EN ve

sterminal dark band on rump дет Preset |]

Central back feathering
with broad pale scallopi C—O ===== od
and large central feather 5 potting Present Present.

[Mouth colour «Yellow — — [Yenowishpink [Flesh-pink — jJ

Cream- o o (54) ee (54)

Present — |

Streaked raw sienna (136)
dark grey-brown

le horn

Mention should be made of a bird
(QMO27590) collected by Julian Ford 10km N
of Esmeralda Homestead, north Queensland on
18 September 1986 with unenlarged gonads hut
a fully ossified ‘adult’ skull and identified as C.
nuchalis because it may bc hybrid. This tentative
suggestion is made because the characters now
obvious in the Bruslee hybrid (Fig. 1A) were far
less so three years prior to its collection (Fig. 1A-
C). At that time its back feathers appeared less
black, its whitish primary coverts lacked the cin-
namon wash, and its crest was only one third
developed.

While QMO27590 has the dorsal plumage of a
Great, it has several lower back and upper rump
feathers washed with pale Cinnamon (1234) with
a dark subterminal band as found in Spotted
Bowerbirds, It shows no sign of the pale malar
stripe or heavy darker mottling on ear coverts and
side of neck typical of Spotted Bowerbirds and
present in the Bruslee hybrid, but it is heavily
mottled and barred on the throat, chest and flanks
much like the hybrid. Notwithstanding an ossi-
fied skull, the lack of gonad activity at peak
courting season and only one pink nuchal crest
feather indicates it to be a subadult. Weight and
measurements of Ford's specimen (Table 2) pro-
vide inconclusive evidence of hybridization,
however, and it is therefore treated here as a
subadult C. nuchalis. Should both bowerbird spe-
cies prove to occur where it was collected a
hybrid origin becomes, however, a stronger pos-
sibility. Hybrids may not always be as obviaus as

476

the specimen recorded here, and may more re-
semble one or other of the putative parents. Given
the potential extent of sympatry of bower-main-
taining Spotted and Great Bowerbirds in central
northeastern Queensland (Frith et al., 1995b) it is
possible that occasional hybridization between
the two is more widespread.

ACKNOWLEDGEMENTS

This study would have been impossible without
the kind support, interest, friendship and hospi-
tality of the McCullough family whom we thank
sincerely, We thank Philippa Horton, SAM;
Richard Schodde, lan Mason and John Wombey
CSIRO, ANWC; Walter Boles, AM and Stephen
Van Dyck, Glen Ingram and Wayne Longmore,
QM for access to specimens in their care and
other help. Mary LeCroy provided Smithe (1975)
for this study. Walter Boles, Les Christidis, Rich-
ard Schodde and an anonymous referee provided
constrictive comment on a draft manuscript. Les
Christidis provided evidence of the maternal par-
entage of the hybrid. Dedicated to the memory of
Julian R. Ford who contributed greatly to the
understanding of distributions and radiations of
Australian birds including both putative parents
of the hybrid described herein.

LITERATURE CITED

CHISHOLM, А, Н, 1966, Н.С. rica and
"Rawnsley's Bowerbird". Emu 65: 234-235.
CHRISTIDIS, L. & SCHODDE, R. 1992, Relationships
among the Birds-of-Paradise (Paradisacidac) and
Bowerbirds (Prilonorhynchidac): protein evi-
dae. Australian Journal of Zoology 40: 343-

COOPER, W.T. & FORSHAW, J.M, 1977, The Birds
of Paradise and Bower Birds, (Collins: Sydney).

DIAMOND, J.M. 1972, Avifanua of the Eastern High-
lands of New Guinea. Publications of the Nuttall
Omithological Club, No. 12: 1-438.

DIGGLES, S. 1867. The Omithology of Australia. (The
author: Brisbane).

ELLIOT , D.G. 1873. A Monograph of the
Paradisaeidae, (The author; London).

FRITH, C.B. 199]. Bowerbirds and Birds of Paradise.
In Forshaw,L. (ed.), Enclopedia of Birds.
ines New York).

n Ma g st lon Pire in bowerbirds
d ton) Ene 94: 60-61.

FRITH, СВ, & FRITH, DN. 1990, Nesting biology
and relationships of the Lesser Melampitta
Melampitia lugubris. Emu ©: 65-73,

MEMOIRS OF THE QUEENSLAND MUSEUM

1993, Courtship display of the Tooth-billed Bowerbird
Scenopoeetes dentirostris ands its behavioural
and systematic significance, Emu 93; 129-136,

1994, The nesting biology of Archbold's Bowerbird
Archboldia papuensis and a review of that of other
bowerbirds (Ptilonorhynchidae). Ibis 136: 153-
16)

FRITH, C.B., FRITH, D.W. & McCULLOUGH, M.
1995b. Great and Spotted Bowerbirds
Chlamydera nuchalis and С. maculata
(Prilonorhynchidae) sympatric and interacting at
each others bowers. Australian Bird Watcher 16:
49-57.

FRITH, C.B., GIBBS, D. & TURNER, К, 1995a. The
taxonomic status of populations of Archbold’s
Bowerbird Archbolida papuensis in New Guinea.
Bulletin of the British Ornithologisis' Club 109-
115.

FULLER, E. 1979. Hybridization amongst the
Paradisacidae. Bulletin of the British
Omithologists’ Club 99; 145-152,

GILLIARD, E.T. 1969. Birds of Paradise and Bower
Birds. (Weidenfeld & Nicolson: London).

GOULD, J. 1869, The Birds of Australia, supplement
(The author: London),

IREDALE, Т, 1950. Birds of Paradise and Bower Birds.
(Georgian House: Melbourne).

KUSMIERSKI, R., BORGIA, G., CROZIER, К.Н. &
CHAN, B.H.Y. 1993, Molecular information on
bowerbird phylogeny and the evolution of exag-
gcrated male characteristics, Journal of evolution-
ary Biology 6: 737-752.

MARSHALL, AJ. 1954. Bower-Birds: their displays
and breeding cycles. Oxford University Press,
Oxford.

MAYR, Е. 1941. Listof New Guinea Birds. (American
Museum of Natural History: New York),

1962. Family Paradisaeidae. Pp.181-204. In Mayr,
E. & Greenway Jr., J.C. 1962 (eds), Check-List
of Birds of the World Volume XV. (Museum of
Camparative Zoology: Cambndge Massachu-
sells).

SCHODDE, К, 1976, Evolution in the birds-of-paradise
and bowerbirds, a resynthesis. Proceedings of the
eed lale International Omithological Congress. 1974:

37-

SCHODDE. Я & MCKEAN, J.L. 1973. Distribution,
laxonomy and cvolution of the gardener bower-
birds Ambyornis spp. in eastern New Guinea with
сир of two new subspecies, Emu 73: 51-

SHARPE, R.B. 1891-98. Monograph of the
Paradisaeidae, or Birds of Paradise, and
тезе таш, or Bower Birds, (The author:

ndon)

SIBLEY, C.G. & MONROE, B.L. 1990, Distribution
and Taxonomy of Birds of the World. (Yale Uni-
versity Press; New Haven).

SMITHE, F.B. 1975. Naturalist's Color Guide. (Amer-
ican Museum of Natural History: NewYork).

A REVISION OF DIACARNUS BURTON AND NEGOMBATA DE LAUBENFELS

(DEMOSPONGIAE: LATRUNCULIIDAE) WITH DESCRIPTIONS OF NEW SPECIES

FROM THE WEST CENTRAL PACIFIC AND THE RED SEA
MICHELLE KELLY-BORGES AND JEAN VACELET

Kelly-Borges, M. & Vacelet , J. 1995 12 01: A revision of Diacarnus Burton and Negombata
de Laubenfels (Demospongiae: Latrunculiidae) with descriptions of new species [rom the
west central Pacific and the Red Sea. Memoirs of the Queensland Museum 38(2):477-503,
Brisbane, ISSN 0079-8835,

Species previously assigned to the genus Latrunculia, which possess discate microrhabds as
the microsclere complement, have been re-evaluated and assigned to Diacarnus Burton and
Negombata de Laubenfels, which both contain spinulate rather than discate microrhabd
microscleres. The type species of Diacarnus, Axos spinipoculum Carter, is redescribed, and
seven new species are described: D. bellae, D. erythraeanus, D. levii, D. ardoukobae, D.
bismarckensis, D. tubifera, and D. megaspinorhabdosa, spp. nov. The type species of
Negombata, Latrunculia corticata (Carter), is redescribed and the only other known species,
Latrunculia magnifica Keller, is transferred to Negombara, Diagnostic morphological char-
acters which emphasize a combination of gross morphology, spicule dimensions, microsclere
morphology and disposition, are identified for Diacarnus and Negombata and the affinities
of these and other latrunculiids, Larrunculia and Sigmosceptrella, are compared to éach other
and to other demosponges. New locality and species records reveal a remarkably disjunct
biogeographic distribution for Diacarnus: the. greatest diversity of species is found in
northern Papua New Guinca and in several Micronesian atolls, but the genus extends east
through New Caledonia to Fiji, and south to Port Jackson in south-eastern Australia, Two
species of Diacarnus are also found in the Red Sea. The genus has not been recorded in the
published literature, nor in the extensive unpublished collections known for the Indo-Malay
region. C] Demospongiae, Latrunculiidae, Micronesia, Red Sea.

Michelle Kelly-Borges, Department of Zoology, The Natural History Museum, Cromwell
Road, London, SW? SBD, United Kingdom, and Coral Reef Research Foundation, Р. O.
Box 70, Weno, Chuuk Atoll, Federated States of Micronesia 96942. Jean Vacelet, Centre
d'Océanologie de Marseille, Station Marine d'Endoume, Rue de la Batterie des Lions,

F13007 Marseille, France; 10 June 1995,

Sponges having discorhabds, dicasters or
spinulate acanthomicrorhabds as microscleres
have traditionally been placed within
Latrunculia Bocage, Latrunculiidae Topsent.
Distinctive microscleres of Latrunculia bear four
whorls of spines frequently united in a disc, on à
straight axial rod, which are disposed in an erect
layer above the sponge ectosome. Latrunculia
contains massive sponges with papillae and
sicve-pore areas, with skeletons of monactinal or
rarely diactinal spicules arranged in an irregular
reticulation of poorly defined, somewhat plu-
mose, tracts, Other latrunculiids such as
Podospongia Bocage, Sigmosceptrella Dendy,
and Barboza Dendy, contain acan-
thomicrorhabds. The affinities of these genera
have been diversely interpreted, resulting in allo-
cation of the family either in the tetractinomorph
Hadromerida, following Bergquist (1978), or in
the ceractinomorph Poecilosclerida following
Topsent (1922),

However, it is unlikely that acan-
thomicrorhabds, which are а distinctive feature of
the family as presently recognised, are homolo-
gous in all of these genera. The gents
Latrunculia, which contains discorhabds, has re-
ceived several species which are certainly atypi-
cal, Two species from the Red Sea, Latrunculia
corticata Carter, 1879 (with its claimed junior
synonym L. magnifica Keller, 1889), and L
purpurea Carter, 1881, were isolated in a new
genus Negambata by de Laubenfels (1936), a
distinction which had been suggested long ago by
Ridley and Dendy (1887) and Thiele (1900), De
Laubenfels differentiated these specimens from
Latrunculia on the morphology of the
microscleres which are spinulate sanidaster-like
acanthomicrorhabds, rather than discate as in
Latrunculia. The position of Axos spinipoculum
(Carter, 1879), transferred to Latrunculia by
Hooper (1986), 1s doubtful. Burton (1934) pro-
posed the new genus Diacarnus to receive Axos
spinipoculum Carter in recognition of the charac-

478

teristic acanthomicrorhabds which consisted of a
straight shaft bearing four whorls of spines,

We have found several sponges in the West
Central Pacific and in the Red Sea which are
characterised by the possession of spinulate
rhabds, hereafter termed spinorhabds. These
microscleres are thin rods with spines or bumps
scattered irregularly or uniformly in 4 whorls
along the shaft. The microscleres of these new
sponges are clearly related to those of Axos
spinipoculum Carter, Latrunculia (=Negombata)
corticala Carter, and L. (=Negombata) magnifica
Keller. These species are re-evaluated in the light
of this new material.

METHODS

Sponges were collected using SCUBA by the
authors and the Coral Reef Research Foundation
(СЕКЕ), Micronesia, from Chuuk (Truk) Siate,
Micronesia, the Australian Great Barrier Recf,
northern and southem Papua New Guinea, New
Caledonia, Fiji, the Philippines, the Red Sea, and
the Gulf of Aden (Fig. 4), On collection, samples
were preserved in 70% ethanol or in 10% forma-
lin and prepared for histological examination as
described in Kelly-Borges et al. (1994). The dried
holotype of Latrunculia (=Negombata) corticata
(Carter) was reconstituted by prolonged soaking
of a fragment in dilute detergent. The colour
notation for living and preserved specimens fol-
lows the Rheinhold Colour Atlas (Kornerup &
Wanscher, 1961). Spicule dimensions are given
as mean length (range of length measurements)
times width (range of width measurements) fol-
lowed by the number of spicule measurements
taken. Primary type materia] has been deposited
in the Queensland Museum, Brisbane, Australia,
paratypes and fragment of holotypes, schizotypes
of some authors, have been deposited in the Nat-
ural History Museum, London, and in the
Muséum National d'Histoire Naturelle (Paris).
Registration numbers are cited in the text.

Abbreviations used in the text: QM, Queens-
land Museum, Brisbane; BMNH, Natural History
Museum, London; MNHN, Muséum National
d'Histoire. Naturelle, Paris; HBOM, Harbor
Branch Oceanographic Museum, Fort Pierce,
Florida; SIO, Scripps Institute of Oceanography,
San Diego; NSRC (UPNG), Natural Science
Resource Centre, University of Papua New
Guinea, Port Moresby; CRRF, Coral Reef Re-
search Foundation, Federated States of Microne-
sia; OCDN, Specimen sample numbers for United
States National Cancer Institute shallow-water

MEMOIRS OF THE QUEENSLAND MUSEUM

collection program contracted to the CRRF. A
complete collection of all ОСОМ specimens is
located at the Smithsonian Institution (United
States National Museum): Q66C, Specimen sam-
ple numbers for United States National Cancer
Institute shallow-water collection m pre-
viously contracted to the Australian Institute of
Marine Science, Townsville. This latter collec-
tion 1 now located at the Queensland Museum.

SYSTEMATICS
Family LATRUNCULIIDAE Topsent

Diacarnus Burton, 1934

Diacarnus Burton, 1934: 549

TYPE SPECIES
Axos spinipoculum Carter, 1879: 286

DIAGNOSIS (émended)

Massive spherical or barrel-shaped, lohate-dig-
itate, repent-branching, tubular or thickly en-
crusting Latrunculiidae. Surface with single or
multiple conules or blunt broad tubercules or
mounds, microscopically smooth, slippery and
rubbery. Ostia radiate in stellate formation in
shallow rounded depressions, oscules apical with
low fleshy raised collars. frequently differentially
coloured cream, always highly contracted and
invisible in preserved specimens. Texture ex-
tremely tough but elastic. Colour usually pale
purple-pink mottled with deep reddish brown and
yellow brown, interior cream. Megascleres sub-
tylostrongyles, with a faint subterminal swelling
бро end, distal end oxeote or more typi-
cally strongylote. Microscleres, two size catego-
ries of spinulate acanthomicrorhabds
(spinorhabds), smallest category always present
and disposed on and below scalloped ectosomc-
choanosome boundary, second larger category,
disposed within the ectosome and choanosome
when present. Skeletal arrangement
plumoreticulate. Primary fibres. conspicuous, ap-
pearing as vertebrate ligaments in soft flesh,
Megascleres are packed and arranged parallel
within axis of fibre, spongin barely visible, fibre
is often hollow. Multiple fine dendritic spicule
tracts emanate from tip of primary fibre as it
passes into ectosome in an umbelliform arrange-
ment, fibres radiating towards surface terminat-
ing in ectosomal megasclere brushes. Primary
fibres, connected by occasional short secondary
fibres, or anastomosing with adjacent fibres,

REVISION OF DIACARNUS AND NEGOMBATA

Ectosome., dense, rubbery, distinct from the un-
derlying choanosome, with layers of collencytes
and collagen fibrils parallel to surface. Choano-
some, dense, with a few large canals surrounded
by a dense layer of contractile cells and collagen
fibrils. Fibres are surrounded by a sheet of elon-

cells and bundles of collagen fibrils. Sponges
incubate huge, bright orange-yellow
parenchymella larvae.

Diacarnus spinipoculum (Carter) comb. nov.
(Figs 1, 4А, 5А, бА, 7A, ЗА; Table 1)

Axos spinipoculum Carter, 1879: 286
Diacarnus spinipoculum; Burton ,1934: 549
Latrunculia spinipoculum; Hooper, 1986- 181

MATERIAL EXAMINED

HoLoTvPE: BMNH 1846.10.14.174 from Port Jack-
son, Sydney, growing on "hard objects" (Carter 1879),
depth unknown, presented by J. B. Jukes, Esq (Fig. 1).
ADDITIONAL MATERIAL: BMNH 1994.5.22.3, west
side of East Fayu Island, Chuuk State, 8°32.61'N,
151°20,01°E, on ledge on outer wall, 24.4 m, collected
by P. L. Colin, CRRF, 14 January 1994; BMNH
1994,5.22.4: East Fayu Island, Chuuk State,
8°32.61'N, 151720.01' E, oceanside, on vertical wall,
10m, collected by P. L. Colin, СЕКЕ, 14 January 1994;
BMNH 1994.8.20.1, (fragment of OCDN 2559-Q):
Blast Pass, 27m, Fiji Islands, collected by C. Arneson,
CRRF, 20 Augusl 1994; Negombo tenuistellata Dendy
- BMNH 1907.2.1.28.

DESCRIPTION
Large spherical or oval barrel-shaped, (Fig.
5A), upto 50cm high, 40cm diameter, with a deep
apical central atrium, 4-5cm diameter. Several
sponges can be joined basally. Oscules, scattered
on interior walls of atrium. Surface, heavily iu-
berculate to mammilate (conulose in the pre-
served holotype due to shrinkage), rubbery and
microscopically smooth to touch, compressible,
elastic in life, barely compressible in alcohol.
Colour in life, mahogany brown (SET), mottled
with pale pinkish red (8C7), rim of apical atrium
and atrium walls cream-white. Cream in ethanol,
Skeleton. Large primary fibres, 500-900 di-
ameter, radiate towards sponge surface, con-
nected by rare short secondary fibres, 250-300um
diameter, or anastomosing with adjacent primary
fibres in deeper choanosome (Fig. 6A). Primary
fibre tip divides into an umbel of numerous sien-
der dendritic spicule tracts, 38-70um diameter,
where the primary fibre passes through ecto-
some-choanosome boundary. Dendritic spicule
tracts occasionally branch below surface and di-
verge and ramify within ectosome, where they

479

form slightly fanned brushes after narrowing to
5-lOum or only 1-4 spicule widths. Megascleres
forma palisade of brushes between dendritic tract
brushes. Megascleres, scattered. singly or in
groups of 1-3 spicules, in a vaguely radiate to
confused arrangement interstitially between pri-
mary fibres. Larger thicker microscleres are scat-
tered predominantly around edges of exhalant
canals in choanosome, around ectosomal lacu-
nae, and just below lower boundary of ectosome
where smaller thinner spinorhabds are also found.
Ectosome, 1200- 150040 deep, extremely dense
with parallel collagen fibrils, easily distinguished
from underlying choanosome which is dense and
soft,

Spicules. Megascleres. Subtylostrongyles with
slight subterminal swelling: 266(232-305) x 4(2-
Тут (see Table 1).

Microscleres: Spinorhabds [: extremely fine
with small sharp spines scattered along shaft;
46(37-55) x <lum, n=10 (Fig. 7A). Spinorhabds
Т: with four groups of spines, terminal spines
frequently strongly recurved, central spines irreg-
ular in shape and length; 57(53-65) x 2(1-5)um,
n=20 (Table 1, Fig. 8A),

SUBSTRATE, DEPTH RANGE, ECOLOGY
Solitary, on oceanside reef slopes, from 10 to
27m depth.

GEOGRAPHIC DISTRIBUTION
Port Jackson, Sydney; East Fayu Island, Chuuk
State, Micronesia; Fiji Islands (Fig. 4A).

REMARKS

The holotype was described by Carter (1879),
and more recently by Hooper (1986), as an "clon-
gate, subcylindrical, solid, cup-shaped with an
apical depression" (Fig. 1). Examination of new
material from Micronesia, and Fiji, reveals that
the species has a large, deep, central atrium in life,
with oscules opening into the atrium. The holo-
type would have almost certainly been spherical
or barrel-shaped in life, with a restricted base of
attachment as in the fresh material, but the spec-
imen is now compressed laterally within the con-
fines of the container, The surface of the
preserved sponge is tuberculate but in life the
tubercules are more distinctly mammilate,

Carter (1879) emphasised the characteristic
"chondroid" nature of the thick ectosome and the
extremely thick, tough fibres set in a relatively
soft, compressible choanosome (Fig. 6A), This
later feature was not emphasised in Hooper's
description, based as it was on an inadequate

480 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Diacarnus spinipoculum (Carter), preserved holotype BMNH 1846.10.14.174, Port Jackson, Sydney.
Scale: 5cm.

REVISION OF DIACARNUS AND NEGOMBATA

TABLE 1. Spicule dimensions of Diacarnus spinipoculum (Carter)
and Diacarnus bellae sp. nov., given as mean length(range). mean

thickness (range), all in pm.

| Megáscleres |Spimorhadel |Spinorhabds Ш
2771232-305) 530-55) TN
255(240-269) |42(38-48) |55(53-60

3.5-5 <1 1-2

BMNH 1994.5.22.4 266(250-278) |43(37-46) a |

amie арр ришти
{Сапег) Н! otype
[BMNH 1846, 14.174

|BMNH 1994,5.22.3

|
ces rit bellae <P. nov. 269(250-280) 42(36-45) 46(43-50)
Holot
BMNY 19945 5.7 e»

| BMNH 1994.5.22.9

| BMNH 1994.5.22.11

older histological section of the holotype, and so
the most diagnostic field characteristic of the
sponge had not been reported in recent literature,

The unusual and characteristic skeletal arrange-
ment of the umbelliform primary fibres within the
ectosome, observable only with detailed serial
histological sectioning, was also not possible to
detect in the slide of the holotype, nor were the
two size categories of microscleres. The ectoso-
mal megasclere brushes are paratangential in ori-
entation in the holotype slide (Hooper, 1986) but
are vertical, as in the living sponge, where the
ectosome has not been squashed due to confine-
ment in its container.

Two specimens from East Fayu Island have
been assigned to Diacarnus spinipoculum (Car-
ter) even though ihe spicule dimensions are
slightly different from the holotype; there is con-
siderable variation between all specimens exam-
ined and spicule dimensions cannot be
considered alone in differentiation of species in
Diacarnus. Even though the spicule dimensions
of these specimens are slightly more similarto the
later described D. bellae, spicule morphology is
very different and identical to that of D,
spinipoculum.

In a revision of Axos Gray from north-western
Australia, Hooper (1986) transferred Aros
spinipoculum to Latrunciia, considering the
general skeletal features of A. spinipoculum to be
similar to those of Latrunculia s.s. The type spe-
cies of Latrunculia, L. cratera Bocage 1869, has
several characteristic features described clearly
by Bocage (1869), and later recognised by Ridley
and Dendy (1887) in the descriptions of
Latrunculia apicalis and L. brevis (Ridley &
Dendy, 1886), Diagnostic features include the

280(250-300) |45(40-50) 45(40-48
9850900) ез» |р
270(250-290) |absent 44(40-48

papo jee ш |

481

possession of dicastra, which form a
single ectosomal layer of erect spic-
ules, their bases implanted in the der-
mal membrane (Fig. 9D). The
dicastra or discorhabds of
Latrunculia s,s, have two central
whorls of serrated discs that are usu-
ally of unequal diameter and which
are invaiiably located closer to either
end of the spicule. The ectosomal ar-
rangement of the dicaslers is well
illustrated by Bocage (1869; pl. 11,
fig. 2b) and the variety of dicasters in
Latrunculia obvious in the
illustrations of Ridley and Dendy
(1887) and Bergquist (1968: 18, fig.
. The lere skeleton is typi-
cally a whispy loose irregular reticu-
lation of indistinct tracts, megascleres
are styles, and microscleres are discorhabds. The
surface of Larrunculia has characteristic mammi-
form ostial sieve-plates. The transfer of Axor
spinipoculum to Latrunculia by Hooper (1986) is
therefore invalid, as it differs considerably in the
form and disposition of the microscleres, the form
and disposition of the fibre skeleton, mesohyl
matrix construction and in surface morphology,
Hooper (1986) also suggested Axos
spinipoculum and Negombo renuistellara Dendy
1905 from Ceylon are synonymous but conceded
that the microscleres of the former are more reg-
ular than those of Negombo. Examination of N.
tenuistellata (BMNH 1907.2.1.28) and histolog-
ical sections of this sponge, reveal that it 1s very
different in morphology and spiculation from A
spinipoculum: N, tenuistellata is а series of com-
pressible thin-walled coalescent tubes, the skele-
ton of which consists of large gently curved styles
in confused longitudinal tracts, reminiscent of the
halichondroid organisation, with minute micro-
spined sanidaster-like microscleres scattered in-
terstitially and in a dermal membrane (Fig. 8H).
The microscleres of N. tenuistellata are acanthose
microrhabds with irregular whorls of spines
along the shaft, and are much smaller than the
microscleres of the Latrunculiidae, Axos
spinipoculum and N. tenuistellata are not synon-
ymous and neither have microscleres typical of
Latrunculia s.s. Negombo was incorrectly placed
in the Spirastrellidae by Dendy (1905), there is по
cvidence of spiralling along the shaft of the spic-
ule, De Laubenfels (1936; 132) used the name
Negombo for an alleged desmoxyid sponge. This
genus name has not been used since, 10 our
knowledge.

482 MEMOIRS OF THE QUEENSLAND MUSEUM

40.4: 6. 5-8.
m dr oU cca. Cota cate

ais Pen MAIS) OE haigh, PIX. es

FIG. 2. Negombata corticata (Carter), dry holotype BMNH 1840.5.6.56-58, three dried fragments of the same
specimen, Red Sea. Scale: 5cm.

REVISION OF DIACARNUS AND NEGOMBATA

Diacarnus bellae sp
(Figs 4B, SB, 5C, 6B, 7B. ЯВ; Table 1)

MATERIAL EXAMINED

Ноготуре: QMG305007: Anaw Wall, oceanside of
reef west of Anaw Channel, northwest side of Chuuk
Atoll, Micronesia, 734.24" N; 151'40.19"E, 24m, col-
lected by P. L. Colin, CRRF, 7 August 1995; FRAG-
MENT OF HOLOTYPE: BMNH 1994.5.22 7.
ADDITIONAL MATERIAL: BMNH 1994,5.22.1] (frag-
ment of OCDN 120-O): west side of Nama Island, 30
nautical miles east of Chuuk Atoll, Micronesia,
6'59.70"N, 152"34.40' E, 10m, collected by P. L. Colin,
CRRF, 3 June 1992; BMNH 1994.5.22.9 (fragment of
OCDN 321-7); oceanside reef, south of northeast Pass,
Chuuk Atoll, Micronesia, 7729,55'N, 151'59.1 E,
I2m, collected by P. L. Colin, CRRF, 8 July 1992;
BMNH 1994.5.22.19 (fragment of SIO-POH 93-005),
Pohnpei, Micronesia, collected by Mary Kay Harper,
510; BMNH 1994.6.14.1 (fragment of OCDN 2503-
G): South Pass Pinnacle, west of South Pass, Chuuk
Atoll, Micronesia, 30 m, 713.49" N; 151"46.25'E, col-
lected by P. L. Colin and Р. Schupp, CRRF, 14 June
1994.

DESCRIPTION

Sub-spherical, solitary sponges (Fig. 5B), 6-
Bem diameter, frequently coalescent with adja-
cent sponges forming a spreading mal upto 60cm
wide (Fig. 5C). Surface with low blunt conules
1-5mm high, separated by 5-7mm, sponge
smooth, rubbery, ostia in differentially
pigmented stellate depressions, Oscules, apical,
single or aligned in rows, with raised cream-col-
oured rims. Texture, compressible but firm, elas-
tic. Colour in life, dull rose (906) mottled with
oxblood red (9E6), large irregular areas and ostial
depressions are oak brown (5D6), Colour of os-
cule rims cream (4A2), in ethanol, uniformly dull
yellowish brown (5B4).

Skeleton: Arrangement plumoreticulate, with
very thick rigid multispicular fibres, 500-
1225um diameter. Secondary fibres, 100-2001 т
diameter, sparse, connect primary fibres to form
an extremely clongate mesh. Megascleres, ar-
ranged parallel to axis of fibre, spongin barely
visible, largest fibres are hollow. Megascleres,
scattered interstitially in choanosome. Curved
branched tracts, 10-30um diameter, emanate
from tips of primary fibres, diverge and ramify
dendritically through ectosome towards surface,
forming fanned brushes (Fig. 6B). Megascleres
form à sparse erect to fanned palisade in outer
ectosome, juxtaposed upon primary fibre
brushes, Thicker category of spinorhabds, found
occasionally within megasclere surface brushes,
lining ectosomal lacunae in greater density, and

483

are common within choanosome, particularly lin-
ing canals. Thinner microscleres, found only
below lower ectosomal boundary. Ectosome,
1225 to 17004m deep. Choanocyte chambers,
approximately 25|um diameter.

Spicules, Megascleres. (Table 1): Sub-
tylostrongyles: 273(240-300) х (3-5)jum, п=50.

Microscleres. (Table 1): Spinorhabd 1: ex-
tremely fine with bumps or very short „pinos
scattered irregularly along shaft; 45(36-50) x
«Ium, n=50 (Fig. 7B); Spinorhabd II: straight,
with distinct short spines along shaft, in four
whorls, or irregularly distributed spines (Fig. 8B).
Degree of spination varies between specimens:
45(40-60) x 1-3um, n=100.

GEOGRAPHIC DISTRIBUTION
Chuuk Atoll, Nama Island, and Pohnpei Island,
Micronesia (Fig. 4B).

SUBSTRATE, DEPTH RANGE, ECOLOGY

Common on oceansides of atoll and island
fringing reefs down to 30m, growing openly
under full illumination.

ETYMOLOGY
Far Lori J. Bell, Cora] Reef Research Founda-
tion, Chuuk, Micronesia.

REMARKS

Diacarnus bellae and D. spinipoculum are very
similar histologically, although the megascleres
and microscleres are consistently slightly smaller
and thinner in D. bellae. The two species can be
easily differentiated hy their gross morphology,
however. Diacamus spinipoculum is a very large
barrel-shaped sponge with a wide, deep, central
atrium lined with exhalant apertures. Diacarnus
bellae is never tubular and occurs consistently in
the field as a small solitary, semi-spherical
sponge, or as several which coalesce to form a
loose cluster, There is no suggestion of an apical
atrium in D. bellae, the oscules are small and
arranged serially or in clusters around the apex of
each sponge.

Diacarnus erythraeanus sp. nov.
(Figs 4C, 5D, 5E, 6C, 7C; Table 2)

MATERIAL EXAMINED

HoLorYPE: OMG305008: near norhem entrance of
Obhor Creck, 30m, Djeddah, Red Sea, 21'50'N,
3910'E, collected by J. Vacelet, 23 February 1983;
FRAGMENTOFHOLOTY PE: BMNH 1995.6.26.1; MNHN
DJV51,

=
5
m
с)
=)
>
a
z
<
m
u
z
а
m
2
©
ш
m
=
ш
©
N
=
©
=
a
z

REVISION OF DIACARNUS AND NEGOMBATA

ADDITIONAL MATERIAL: MNHN DJV52: fore reef
zone, free living on the sand and embodying а piece of
rope, 30m, King Saud Palace, Djeddah, 21'70'N,
38'80'E, collected by J. Vacelet, 8 March 1983;
BMNH 1994.5.22.17: Ras Muhammad, 2-10т, South
Sinai, Red Sea, dry specimen collected by Loya, 1977
{Loya 11); MNHN DJV53: Râs Muhammad, South
Sinai, Red Sea, collected by Loya, 1978 (Loya 254);
MNHN DJV54: Harvey Reef, 10m, near Port Sudan,
collected by C. Wilkinson, August 1978; BMNH
1978.12.14.2: Harvey Reef, 10m, near Port Sudan, C.
Wilkinson, August 1978.

DESCRIPTION

Repent or erect branches, 20-3(imm diameter
(Fig. 5D), variable length and branching pattern,
often with slightly enlarged ends up to 50mm
diameter, often anastomosing to forma sprawling
mass up to 1m diameter, frequently free living on
sand (Fig. SE) or attached to debris. Texture,
tough, just compressible, elastic. Surface, irregu-
lar with broad blunt conules, 2 mm high, on ends
of branches. Ostia, 40-60um diameter, localized
in groups of about 10 in smal! surface depres-
sions. Oscules, 2-3mm diameter, found on upper
surface of branches or at tips of enlarged branch
ends when in upright position, opening at end of
raised cream-coloured fleshy collars. Colour,
rose pink (9106), mottled with oak brown (5D6)
on surface, darker brown in approximately hex-
agonal concave depressions surrounded by cream
ridges, more uniformly brown near end of
branches. Interior in life and whole sponge in
ethanol, cream (442),

SKELETON

Primary fibres, 500-800pm diameter, up to
1800um in centre of branch, radiate towards
sponge surface, connected by sparse short sec-
ondary fibres, 270-420um diameter.
Megascleres, scattered interstitially in choano-
some, Curved tracts, 30-50um diameter, less than
20um diameter when crossing ectosome-choano-
some boundary, emanate from tips of primary

fibres, diverge and ramify dendnitically through
ectosome, occasionally dividing just below sur-
face, giving rise to brushes (Fig. 6C).
Megascleres, form an erect to fanned palisade
juxtaposed upon ectosomal brushes.
Spinorhabds, rarely present, distributed just
below the ectosome in upper choanosome, larger
category absent. Ectosome, 600-900um йеср,
distinct, rubbery. Choanoc yte chambers, approx-
imately 251m diameter.

Spicules. Megascleres (Table 2), Sub-
tylostrongyles, strarght, with a very faint swelling
of proximal end, distal end rounded: 240(190-
269) х (2.5-4.8)рт, п=30.

Micrascleres (Table 2). Spinorhabds I: straight,
with occasional swellings or short spines scal-
tered along shaft and near apices, occasional to
rare; 25(24-30) x 0.8-1.3ит, n= 25 (Fig. 7C).

GEOGRAPHIC DISTRIBUTION
South Sinai, Eilat, Harvey Reef off Port Sudan,
Red Sea (Fig, 4C)

SUBSTRATE, DEPTH RANGE, ECOLOGY

Common on coral substrate or rocks, large
specimens, free living or allached to small corals
fragments in sand, Depth range, 10-30m, al-
though reported to he very common at Eilat in
shallow water of 2-10m (Loya, pers. comm,
1978), Parenchymella larvae, 2mm diameter,
within choanosome of MNHN DJV52.

ETYMOLOGY
The species name indicates that the sponge is
found only in the Red Sea.

REMARKS

This sponge is easily recognised in the field by
its external morphology of repent anastomosing
branches, mottled surface colouration, conulose
surface and smooth rubbery texture, and tough
internal fibres. The small thin spinorhabds are

FIG. 3. Comparison of the developmental and mature stages of spinorhabd microscleres of Sigmoscep{rella,
Diacarnus and Negombata. A-D, Sigmosceptrella fibrosa, Dendy (BMNH 1925.11.1.717), Maria Island,
Tasmania: A, B, rhabd in carliest stage with strongly recurved spines similar to those of Negombata magnifica
(K-L). C, the maturing spinorhabd shows signs of the fused recurved spines on the shaft, D, mature spinorhabd
with a distinctly sigmoid rhabd on the right. Scale; A: 9.lum; B: бшп; C: 10.4um; D: 18.2um. E-G,
Sigmosceptrella quadrilobata Dendy (E-100, X-371), Tuléar, Madagascar. E, specimen E-100, sigmoid rahbd,
Е, specimen X-371, mature spinorhabd. С, specimen X-371, mature spinorhabd with post-fusion sigmoid rhahd.
Scale: E: 9.1um; F: 16.5pm; G: 18.2um. H-J, Diacarnus ardoukobae sp, nov., holotype ОМС305010, Н,

spinulate rhabd, 1, maturing spinorhabd. J, mature spinorhabd. Scale: Н: 9.1ит; 1: 18.84m; J; 14.
egombata magnifica (Keller), MNHN DJY 56. К, rhabd with irregular sharply re-curved spines.

spinorhabd. Scale: К: 5.2]m; L: Sum.

. KL,
, їпайге

486

TABLE 2. Spicule dimensions of Diocarnes
erythraeanus sp. nov., given as mean length (range),
Width, all in рт.

! [Megascleres [Spinorhads1 — |

249(210-269) | .

Daag) [absent |

243(21 1-269)
BMNH197812142 |99014 absent

rare and are easily overlooked in spicule prepara-
tions and histological sections.

Diacarnus levii sp. nov.
(Figs 4A, SF, 6D, 70; Table 3)

MATERIAL EXAMINED

HOLOTYPE: QMG305009; (ORSTOM R1524), Sr.
478, lagoon, Belep, New Caledonia. 4-25m,
19734,03' S. 163'42,03'E, 8 March 1990; FRAGMENT
or HoLorvPE: BMNH 1994,5.22.16; MNHN
DCL3659.

ADDITIONAL MATERIAL: MNHN DCL3660- (OR-
STOM R767), St. 206, Banc de la Torche, New Cale-
donia, 35m, 22'56.60'S, 167°40.00'E; MNHN
DCL23661; (ORSTOM R879). St, 184, 18-25 m, lagoon
ilot Ua, New Caledoma, 2243.00' S, 16649. 10" E. 28
June 1979; MNHN DCL 3662: (ORSTOM R994), St,
225, 42m, Canala Pass, New Caledonia, 21°18.80'S,
165'57.25' E; MNHN DCL3663: (ORSTOM R1247),
St, 270, 8m, Kouare lagoon, New Caledonia,
22'46.50'S, 166'47 90E, 3 March 1980; MNHN
DCL3664: (ORSTOM R1291), St. 303, 6m, forc-recf
zone, Cook Reef, New Caledonia, 19'45.60'S,
161°41.40°E, 23 June 1981; MNHN DCL3665: (OR-
STOM R1528), SL 448, 6-15m, fore-reef zone, Abore
reef, New Caledonia, 22'20.00'S, 166713,15'E;
BMNH 1994.5,22,15 (fragment of Q66C-0248)- west
side of seaward vertical wall of reef, I&m, Big Broad-
hurst Reef, Great Barrier Reef, Australia, 1856,67 S,
147°43.86°E: BMNH 1994,5,22.8 (fragment of Q66C-
0890): Black Reef, Whitsunday Islands, Great Barrier
Reef, 10m, collected by М. Kelly-Borges, 20 October
1987.

DESCRIPTION

Thick erect digitations or lobes, 30-45mm di-
ameter, anastomosing in large masses (Fig. 5F).
Texture, tough, just compressible, elastic. Sur-
face, with low rounded conules, approximately 2
mm high, 2-5mm apart, Ostia, 5Oum diameter,

MEMOIRS OF THE QUEENSLAND MUSEUM

found in small, darker surface depressions, Os-
cules, on apices of digitations, surrounded by a
white margin, 10mm diameter in preserved spec-
imens, Surface with rounded depressions con-
taining ostia in stellate arrangement. Colour in
life oak brown (5D6), darker in ostial depres-
sions, mottled with cream (4A2) in surface
patches and around oscule margin; cream inte-
rior, uniformly white in ethanol.

Skeleton. Plumoreticulate arrangement with
very thick multispicular fibres, 700-1 [251m di-
ameter, joined by small short secondary fibres,
250-375um diameter, at right angles to primary
fibres (Fig. 6D). Small dendritic tracts, 30-50ura
diameter, emanate from tip of fibre, dividing and
radiating through ectosome toward surface, end-
ing in narrow brushes, surface raised in large
bluntconules. Megascleres form an erect palisade
at surface, Interstitial megascleres, abundant.
Spinorhabds, rare, found only in superficial lay-
ers of choanosome when present, Ectosome,
thick, 500-1 100m, composed of parallel colla-
gen fibrils in wavy bundles. Choanocyte cham-
bers, 20 to 25um diameter.

Spicules. Megascleres. (Table 3): Sub-
tylostrongyles, with slight swelling at proximal
end: 258(2 10-300) x 2-5шп, n-30.

Microscieres. (Table 3): Spinorhabd I: straight,
with swellings or small, irregular spines usually
more developed near apices, always rare, may be
absent in some specimens: 58(53-60) x 0.5-
2.4um, п=10 (Fig, 7D).

SUBSTRATE, DEPTH RANGE, ECOLOGY

Found on fringing coral reef slopes down to
35m. Large embryos or parenchymella larvae, up
(о 1.5mm diameter, white or yellow in the
choanosore, are observed in all specimens.

GEOGRAPHIC DISTRIBUTION
New Caledonia; northeastern Great Barrier
Reef, Australia (Fig. 4A)

ETYMOLOGY
For Professor Claude Lévi,

REMARKS

Diacarnus levii appears to be fairly common in
New Caledonia, but has only been collected from
two locations on the Great Barrier Reef. In both
locations the sponge was rare. Specimens from
the Whitsunday Islands and Big Broadhurst Reef,
Australia, have slightly larger megascleres and
the spinorhabds are slightly thicker with better
developed spines, however, they are similar to the

REVISION OF D/ACARNUS AND NEGOMBATA

TABLE 3. Spicule dimensions of Diacarnus levii sp.
nov, given as mean length (range), width (range), all
in jim.

| Sample __ |Megascieres |Spinorhadsl |

|

| Diacarnus levii

| Holo 241190-280) 59(55- 62. 5)
OMG305009 (2.4- "es |
= DCL3663 248(245-315)

| BMNH 1994 5.22.15 азаи 288) ses m
ror VES m
[emmes |

New Caledonian specimens in all other aspects.
Diacarnus levii is differentiated from the Red Sea
species, D. erythraeanus, by gross morphology;
D. levii is more massive than D. erythraeanus
which is ramose and branching. The major spic-
ule difference between D. levii and D.
erythraeanus ts that the spinorhabds in the former
are almost twice as long as those of D.
erythraeanus.

Diacarnus ardoukobae sp, nov.
(Figs ЗН, 31, 3J, 4С, 5G, 6E, 7E, 8C)

MATERIAL EXAMINED

HOLOTYPE: QMG305010, Musha island, Djibouti,
Gulf of Aden, 11*42.50'N, 43708.40' E , fore reef zone,
several specimens observed, some free living on sand,
24m, collected by J. Vacelet, 5 January 1985 (M 8);
FRAGMENT OF HOLOTYPE: BMNH 1994,5.22.14,
MNHN DIV55.

DESCRIPTION

Curved erect or repent branches (Fig. 5G), sep-
arate or forming a sprawling mass up to approxi-
mately 50cm long, uniform 1.5-2.0cm diameter.
Ends of branches markedly conulose, branch sur-
faces irregularly bumpy with occasional conules,
texture compressible, rubbery, Oscules distrib-
uted along branches rather than at tips of branch.
Colour in life pale pink motiled with dark pink in
surface depressions, uniformly cream in ethanol
(4A2),

Skeleton. Primary fibres, 300-1000um diame-
ter, radiate towards sponge surface, connected by
abundant secondary fibres, 60-300рт diameter.
Six to ten short compact fibres, 30-70jim diame-
ter, radiate from tip of primary fibre, occasionally
branching to form irregular sparse surface
brushes (Fig. 6E). Megascleres form an irregular
and often paratangential palisade between fibres.

487

Megascleres, abundant in choanosome. Ecto-
some, generally 300tim deep, ranging from 120
to 540m. Microscleres, two categories, smaller
dispersed under ectosome-choanosome bound-
ary, larger microscleres form a dense layer in
middle of ectosome (Fig. 6E), scattered through-
out choanosome particularly around canals,
never found on the outer layer of ectosome,

Spicules. Megascleres: Phry kqstiowgy les:
255(220-288) x 2.4-6um, n=30

Microscleres. Spinorhabd I: “spines are very
irregular in shape and disposition, terminal spines
occasionally bifurcate, central shaft bumpy, two
major whorls of spines are evident, and closer to
ends of spicule rather than regularly spaced along
axis as in second category: 29(24-34) x 2.441m
(Fig, 3H, 7E); Spinorhabd TI: conical spines are
typically disposed in irregular whorls along shaft,
inner two being usually a greater distance apart
than each are to terminal spine cap (Fig. 31, 3J,
8C). Terminal spine cap and outer whorl fre-
quently merge, these spicules are often shorter
than those with equally spaced whorls; 54(48-
67m, n=10, total maximum width 19-31p1m,
shaft diameter 2-4.5p1m.

GEOGRAPHIC DISTRIBUTION
Known only from Djibouti, Gulf of Aden (Fig.
4C).

SUBSTRATE, DEPTH RANGE, ECOLOGY

Specimens free-living on sand at 24m deep.
Reproductive mode, possibly fragmentation and
release of parenchymella larvae às in other spe-
cies of Diacarnus.

ETYMOLOGY
For the "Ardoukoba" expedition during which
specimens were collected,

REMARKS

External morphology is similar to D.
erythraeanus, which is also ramose and branch-
ing. This latter species forms mats of anastomos-
ing branches which typically have enlarged ends
upon which the oscules are located, Specimens of
D. ardoukebae are separate, elongate branches,
uniform in diameter, with flush oscules along
each branch. Histologically, D. ardoukobae is
distinct from D. erythraeanus, as it has a compar-
atively thin ectosome and less robust fibres which
form a rather tighter meshed skeleton than in D,
eryrhraéeanus; The fibres which diverge from the
apex of the primary fibres in D. ardoukobae are
comparatively thick and short and much reduced

488

А
Namal.

Papua New Guinea Q
"Ts uL
=2

о Micronesia
к: East Ғауџ І.

-®© Chuuk (Truk) Atoll
пара

Australia
, Caledonia

. б)
Port Jackson
р.

v)

Ы -
Albatross Passage руаш,
се, So New Ireland
Watom 155: |
Tr
x JH Duke of York Is.
Bay

-Kimbe
тү Se OD e,
Planet e New Britain

Bismarck Sea

|
|
{е New Guinea
l
|

Fig. 4. Geographic distribution of Diacarnus spp. and Negombata spp. A.
Indo-West Pacific and Australia: & Diacarnus spinipoculum (Carter);
Q Diacarnus bellae sp. nov.; V Diacarnus levii sp. nov. № ;Diacarnus
megaspinorhabdosa sp. nov. B. Papua New Guinea: x Diacarnus
bismarckensis sp. nov.; + Diacarnus tubifera sp. nov.; & Diacarnus
megaspinorhabdosa sp. nov. C. Red Sea and Gulf of Aden: Фф Diacarnus
erythraeanus sp. nov; Ф Diacarnus ardoukobae sp. nov; € Negombata

magnifica (Keller).

in number compared to those of D. erythraeanus.
The major difference on which these two species
are separated, however, is the presence of a sec-
ond category of spinorhabd - a larger, more robust
and heavily spined microsclere, which is dis-
posed in а mid-ectosomal layer.

Diacarnus bismarckensis sp. nov.
(Figs 4B, 5H, 6F, 7F, 8D)

MATERIAL EXAMINED

HoLoTvrE: QMG305011: (fragment of OCDN 705-
Y) Planet Rock, 15m, south of Madang, northern
Papua New Guinea, 5° 15.475'S, 145" 49,12" E, col-

MEMOIRS OF THE QUEENSLAND MUSEUM

lected by P. L. Colin, CRRF, 6
November 1992; FRAGMENT OF
НоготүРЕ: BMNH 1994.5.22.13.
ADDITIONAL MATERIAL: BMNH
1994.5.22.12 (fragment of OCDN
754-C): Banban recf (uncharted),
20m, Kimbe Bay, West New Brit-
ain, Papua New Guinea, 4° 55.53”
S, 150° 55.51°E, collected by P. L,
Colin, CRRF, 14 November 1993;
BMNH 1993,11.5.1: oceanside of
Makada Reef, 15m, Duke of York
Islands, East New Britain, Papua
New Guinea, 4° 6.68'S, 152°
23,87'E, collected by P. L. Colin,
CRRF, 5 November 1993; BMNH
j993.10.29.1: overhang on outer
reef near Albatross Passage, 6-
12m. north-western New Ireland,
Papua New Guinea, 2" 45.23'S,
150* 43.24 E, collected by P. L.
Colin, CRRF, 29 October 1993;
BMNH 1993.11.1.1: south coast
wall near east end of Djaul Island,
western New Ireland, Papua New
Guinea, 2° 58.69'S, 150° 59.33 E,
collected by P. L. Colin, CRRF, 1
November 1993,

DESCRIPTION

Ramose, each branch 1-2cm
diameter, approximately 30cm
length, branches anastomosing,
forming large clumps hanging
off overhangs or growing erect
from coral surface (Fig. 5H).
Texture, slightly compressible,
difficult to tear, rubbery. Sur-
face, conulose but microscopi-
cally smooth, Colour in life
cream flecked with garnet-
brown (9D7) and copper red
(7C7), tips and undersides pale
cream. Ostia, set in garnet
brown stellate surface depressions, set in a paler
copper red surrounding.

Skeleton, Primary fibres, 300-600um diameter,
run parallel with branch, connected by perpendic-
ular short secondary fibres, 120-300um diameter
(Fig. 6F). Dendritic fibres, 30-50um diameter,
radiate from tip of primary fibre, occasionally
branching, forming surface brushes. Interstitial
megascleres, abundant in choanosome. Ecto-
some, 300-450um deep on sides of branches.
Larger category of spinorhabds distributed in an
ordered layer just below surface of sponge, abun-
dant in choanosome and along ectosome-choano-
some boundary in places (Fig. 6F),

REVISION OF DIACARNUS AND NEGOMBATA

Spicules. Megascleres. Subtylostrongyles:
274(250-320) x 2.5-8um, n=30.

Microscleres. Spinorhabd I: spines are rela-
tively regular in shape, disposed in four equidis-
tant, frequently incomplete whorls: 39(33-48) x
2.4um (Fig. 7F). Spinorhabd П; sharp conical
spines of equal length, disposed in regular whorls
along shaft, each equidistant from the other:
58(46-67)um, п=10, total maximum width 14-
26um, shaft diameter 5-10um (Fig. 8D).

GEOGRAPHIC DISTRIBUTION

Bismarck Sea (Fig. 4B), very common on bar-
rier patch reefs of western New Ireland, New
Britain, and Madang, north coast of mainland
Papua New Guinea, particularly common around
Djaul Island off the northwest tip of New Ireland
(L. J. Bell, pers. comm, 1994).

SUBSTRATE, DEPTH RANGE, ECOLOGY

Found predominantly on ledges, overhangs, on
cave walls within 10 to 30m. Reproduction by
production of large yellow-pigmented larvae.

ETYMOLOGY
For the Bismarck Sea,

REMARKS

General morphology, growth form, and col-
ouration of this northern Papua New Guinean
species 15 remarkably similar to that of Diacarnus
ardoukobae from the Gulf of Aden, Both are
ramose, have a relatively thin ectosame, and pos-
sess a second larger category of spinorhabd dis-
posed in a central to upper ectosomal layer and
throughout the cheanosome. The major differ-
ence between these two species is the spicule size
and morphology of the largest spinorhabds.
Megascleres and microscleres in D. bismarcken-
sis are longer than in 2. ardoukobae, and the
latter are more abundant in D. bisinarckensis. In
D. bismarckensis, microsclere spines are very
regular, of equal Jength, and disposed equidis-
tantly along the shaft of the spicule. In D.
ardoukabae. spines are irregular, with the central
whorl of spines being longer, and disposed closer
to the ends of the spinorhabd, These differences
are also apparent in the smaller spinorhabds of D.
ardoukobae,

489

Diacarnus tubifera sp. nov.
(Figs 4B, 51, 5], 6G, 7G, 8E)

MATERIAL EXAMINED

HOLOTYPE: QMG305012: oceanside reef near Pizion
Island, Chuuk Atoll, Micronesia, 45m, 7^ 1 1.10" N,
151° 50.20'E, collected by P. L. Colin, СЕКЕ, 7 Janu-
ary 1994; FRAGMENT OF HoLoTYPE: BMNH
1994,5.22,2

ADDITIONAL MATERIAL: BMNH 1994,5,22,1; west
side of East Fayu Island, 30.5m, 60nautical miles
north-northwest of Chuuk Atoll, Micronesia, 8°
32.61°N, 151° 20.01'E, collected by Р. L. Colin,
СЕКЕ, 14 January 1994; BMNH 1994.5 22.10 (frag-
ment of (CDN 716-M): west side of barrier reel sur-
rounding Bagabag Island, 28m, 30nautical miles
offshore on northem Papua New Guinea coastline, 4°
47.15' S, 146° 10.96 E, collected by P. L. Colin, CRRF,
9 November 1993; 066C-6138; attached to vertical
wall, 40m, "The Grotto", southeast Watom Island, off
western coast of New Britain, Papua New Guinea, 4°
6.2'8, 152" 5.7°E, collected by M. Kelly-Borges, 16
October 1991; BMNH 1993.11.1.2: south coast wall
near east end of Djaul Island, western New Ireland,
Papua New Guinea, 2" 58.69°S, 150° 59.33'E, col-
lected by Р, L, Colin, CRRF, 1 November 1993;
BMNH 1993.10.22,]: oceanside of Eastern Fields
Atoll, 9m, Coral Sea, 90 miles southwest of Port
Moresby, southern Papua New Guinea, 10° 00.805,
145° 40.24'Е, collected by P. L. Colin, СЕКЕ, 22.
October 1993; BMNH 1995.7.4.5: Tubbataha Reefs,
south of Cagayan Islands, Sulu Sea, Philippines,
8°40°N, 120*E, disturbed rubble and sand, 10m, col-
lected by L. Sharron, 21 April 1995; BMNH
1995.7.4.3: Tubbataha Reefs, south of Cagayan Is-
lands, Sulu Sea, Philippines, 8°40"N, 120°E, on verti-
cal coral wall, 37m, collected by L. Sharron, 22 April
1995; BMNH 1995.7.4.2: Puerto Princessa, Palawan,
Philippines, 9?50"N, 118°30'Е, 12-18m, collected by
P.L. Colin, 15 April 1995.

DESCRIPTION
Single tubular to vase-shaped, very heavy,
thick-walled (Fig. 5I), average height SOcm, 15-
30cm wide, most frequently twice as tall as wide,
with large lobate processes on the outer surface,
surface tuberculate. Atrium, apical, typically up
to I5cm wade in vase-shaped sponges (Fig. 5J),
typically about 8cm diameter in tubular sponges,
Smaller specimens extremely conulose, often 10-
14cm diameter, 9-10cm high. with a Sem wide
atrium, Texture, very firm, rubbery, mesohyl ma-
trix very dense, fleshy. Colour in life carrot red
(6B7) mottled with bright reddish brown (9D8),
very patchy, interior of tube bright white. beige
in ethanol.
Skeleton. Primary fibres, 450-600,m diameter,
sparse, run longitudinally, anastomosing along
axis of tubular body towards sponge apex. Large

UN

em

z
2
2
z
2
2
-
о
m
Е
б
0)
=
т
Д
2

REVISION OF DIACARNUS AND NEGOMBATA

branches, 150-300jtm diameter, emerge some
distance below ectosome-choanosomal boundary
from upper 5-10mm of primary fibre. Fibre ex-
tensions radiate towards sponge surface, branch-
ing many times before entering ectosome
(900-2400um deep; average depth 1500pm, with
lacunae 30-60um wide), forming an immense
number of very fine, long, dendritic fibres which
meander through choanosome and ectosome ul-
timately forming small surface brushes in outer
ectosome (Fig. 6G). Immense umbells raise sur-
face into rounded lobes. Abundant megascleres
form a palisade superimposed over the surface
brushes. Primary fibres anastomose, also con-
nected by short secondary fibres 120-300yum di-
ameter. Interstitial megascleres, abundant,
arranged in loose broad bundles between primary
fibres. Large spinorhabds, scattered in outer ecto-
some, slightly more abundant in outer third of
ectosome, absent from outer 50um, common
throughout choanosome. Megascleres and
microscleres abundance variable between speci-
tinens. Smaller category of spinorhabds dispersed
under ectosome-choanosome boundary.

Spicules. Megascleres. Subtylostrongyles:
304(250-345) x 2.5-Sum, n=30.

Microscleres. Spinorhabd 1: extremely fine
with fine spines dispersed along shaft: 38(38-41)
x 2.5um, n=10 (Fig. 7G); Spinorhabd TI: distinct
whorls of spines: 60(55-67) x maximum width
17-19, shaft width 3.5-Sum, n=30 (Fig. 8E).

GEOGRAPHIC DISTRIBUTION

Chuuk Atoll and East Fayu Island, Micronesia;
Bismarck Sea, widely and sparsely distributed on
barrier patch reefs of western New ireland and
New Britain, and Madang on northern coast of
mainland Papua New Guinea (L. J. Bell, pers.
comm.); Eastern Fields Atoll, southern Papua
New Guinea (Fig. 4B),

SUBSTRATE, DEPTH RANGE, ECOLOGY

Solitary and uncommon in any one locality,
they are found on vertical walls and steep-sloped
oceanic fringing reefs to depths of 45m, Mode of

491

jon not observed, sponge surface fre-
quently completely infested with harnacles,

ETYMOLOGY
Reflects the consistently tubular morphology.

REMARKS

Diacarnus tubifera is conspicuous and easily
recognised in the field as large long solitary or
double tubes, with an extremely heavy, rubbery
texture. Smaller specimens are also tube-shaped
and extremely conulose, also with a deep central
atrium, Histologically, the sponge is well differ-
entiated from all species described thus far. The
sponge has a very thick ectosome which is rami-
fied with abundant dendritic fibres which ema-
nate well within the choanosome, arising to form
a lobed surface, The megascleres and large cate-
gory of spinorhabds are larger than in previously
described species, and the large spinorhabds are
scattered throughout the ectosome rather than
being restricted to a distinct mid-ectosomal band
such as in D. ardoukobae and D. bismarckensis.

Diacarnus megaspinorhabdosa sp, nov.
(Figs 4B, 5K, 6H, 61, 7H, 8F)

MATERIAL EXAMINED

HOLOTYPE: QMG305013: fringing reef 150m south of
Motupore Island, 10m, Bootless Bay, Papua New
Guinea, 9" 31.6'S, 147° 16.6 E, collected by M, Kelly-
Borges, 6 December 1985; FRAGMENTS OF HOLOTYPE:
BMNH 1994.5 ,22.6, NSRC(UPNG) 90,

ADDITIONAL MATERIAL: BMNH 1994.5.22.5: (frag-
ment of OCDN 673-P): lagoon side of Tab (Pig) Island,
10m, on Madang barrier reef, Papua New Guinea, 5°
10.21'8, 145° 50,37" E, collected by P, L- Colin, CRRF,
3 November 1992; HBOM 002:00925: fringing recf
south of Motupore Island, 10m, Bootless Bay, Papua
New Guinea, 9" 31.6'S, 147° 16.6°E, collected by M.
Kelly-Borges, 24 January 1994; BMNH 1994.7,25,1
(fragment of OCDN 2693-Му: attached to rabble. 7m.
Gamao Point, Batangas, Philippines, 13° 38,39" N, 120°
56.86'B, collected by C. Arneson, CRRF, 25 July
1994; BMNH 1995,5.7,4.1; Zambales, West Luzon,
Philippines, 15*50" N, 120°Е, silty reef, 18m, collected
by P.L. Colin, 2 May 1995; BMNH 1995.7.4.4:
Zambales, West Luzon, Philippines, 15*50'N, 120^ E,

FIG. 5. Species of Diacarnus in situ. Sec descriptions is text for size in life; A. Diacarnus spinipoculum (Carter),
Fiji, BMNH 1994,8,20,1. B-C. Diacarnus bellas sp. nov., Chuuk Atoll, Micronesia: B. solitary form, BMNH
1994.5.22.9. C. coelescent form, holotype QMG305007. D-E. Diacarans erythraenus sp. nov., Djeddah, Red
Sea, holotype QMG305008. F. Diacarnus levii sp. nov., Belep, New Caledonia, holotype QMG305009, G.
Diacarnus ardoukobae sp, nov., Djibouti, holotype QMG305010, Н. Diacarnus bismarckensis sp nov., Djaul
Island, Papua New Guinea, BMNH 1993.11,1.1. 1-J. Diacarnaur tubifera sp. nov., Ојаш Island, Papua New
Guinea: L tubular form, BMNH 1993,1 |. 1,2; J. vase-form, К. Diacamas megaspinorhabdora sp. nov., Balangas,

Philippines, BMNH 1994,7.25.1.

492 MEMOIRS OF THE QUEENSLAND MUSEUM

REVISION OF DIACARNUS AND NEGOMBATA

43m, collected by P,L. Colin, 3 May 1995; BMNH
1995,9.1.1, BMNH 1995.9.1.2; south side of Puerto
Princessa Bay, Puerto Princessa, Pallangbato, Philip-
pines, 9°41.62'N, 118°4.51'E. Collected by P.L. Colin
CRRF, 15 April 1995,

DESCRIPTION

Thickly encrusting, lobate, some specimens
with restricted base of attachment, 6-15cm diam-
eter, lobes up to 4cm high (Fig, 5K). Lobe apices
covered in tough sharp conules approximately 2
mm high, 2-5mrn apart, Oscules, 1-3mm diame-
ter, with smooth raised cream collars, scattered
apically on lobes or irregularly over surface. Tex-
ture, barely compressible, tough, harsh to touch
but microscopically velvety between conules.
Colour in life oxblood red (9E8), mustard (SC8)
internally, uniformly cream (4A2) in ethanol,
BMNH 1994,5.22.5 from Madang was pale
brown (4B4) in life due to shading effects, Very
large bright yellow embryos are abundant in
BMNH 1994.5.22.6 (Fig. 6I).

Skeleton. Primary fibres, sparse, tough, up to
900um diameter, raise surface into well-sepa-
rated tough, sharply pointed conules (Fig. 6H).
Primary fibres joined by short, very thick, sec-
ondary fibres, up to 500jtm thick. Sparse spicule
tracts, 60-100nm wide, diverge from primary
fibre well within choanosome, branching, occa-
sionally forming well separated tracts that radiate
within ectosome. Tracts end in dense brushes at
surface where they interdigitate with a dense
loose palisade of megascleres. Interstitial
megascleres, numerous in choanosome, dis-
persed without order between fibres. Larger fi-
bres are surrounded by a sheath of bundles of
collagen fibrils, 60-80um thick, which also con-
tains loose megascleres arranged longitudinally
along fibre. Large spinorhabds, abundant in ecto-
some and disposed in an undulating band in upper
portion of ectosome surrounding lacunae, also
abundant above lower ectosomal boundary and
abundant in choanosome, sometimes patchily,
sometimes concentrated around canals (see Fig.
6T). Small spinorhabds, below ectosomal-
choanosomal boundary.

493

Spicules. | Megascleres. Subtylostrongyles:
318(298-346) x 7-12um, approximately 201m
shorter (average) in BMNH 1994,5.22.5 from
Madang.

Microscleres, Spinorhabds I: slender rods with
regular whorls of small spines; 48(43-55) x 3-
5pm, shaft lum wide (Fig. 7H); Spinorhabds Il:
spines blunt, central shaft very thick, occasion-
ally malformed with central shaft bare or spines
re-curved along shaft; 84(74-96)um, n=30, max-
imum width 36(34-48), shaft width: 15(10-
19)um (Fig. ВЕ).

SUBSTRATE, DEPTH RANGE, ECOLOGY

Found on silty, fringing coral reefs down to
10m. Choanosome contains large conspicuous
embryos of parenchymella type, up to 1.5mm
diameter. Easily observed in specimens in the
field, they are bright orange-vellow, contrasting
with cream mesohyal matrix, they are abundant,
present at any time of collection, Larvae, loosely
embedded within the central part of body (Fig.
61), many contain thin megascleres dispersed
without order in central region. Microscleres, ah-
sent from embryos. Round or ovoid inclusions,
50-80um diameter, with a fibrillar content, abun-
dant in the choanosome, possibly spermatocysts,

GEOGRAPHIC DISTRIBUTION

Northem (Madang), southern (Motupore Is-
land) Papua New Guinea; Batangas, Philippines
(Fig. 4B)

ETYMOLOGY
Reflects large size and abundance of largest
category of spinorhabds.

REMARKS

The major character separaüng Diacarnus
megaspinorhabdosa from other species of
Diacarnus is the possession of extremely large
spinorhabds, and the largest average megasclere
dimensions. These microscleres form dense ag-
gregations in the choanosome and are disposed in
an upper and lower layer within the ectosome.

FIG. 6. A-L, Skeletal arrangements of Diacarnus and Negombata spp. A, Diacarnus spinipoculam (Carter),
holotype BMNH 1846.10.14.174, 30 x. B, Diacarnus bellae sp. nov., BMNH 1994,5,22.11, 30 x, C, Diacarnus
eryrhraeanus sp. пду., MNHN DJV52, 30 x. D, Diacarnus levii sp, nov., holotype QMG305009, 30 x. E,
Diacamus ardoukobae sp, nov., holotype QMG305010, 30 х. Е, Diacarnus bismarckensis sp. nov,, BMNH
1994.5.22.12, 30 x. G, Diacarnus tubifera sp, nov., BMNH 1994,5.22,10, 30 x, H-1, Dicanius
megaspinorhabdosa sp.nov., [o QMG305013. Н, skeletal arrangement, 30 x. L larva, 120 x. J, Negombata

corticata (Carter), holotype BMN

1840.5,6.56-58, 30 x. K-L, Negombata magnifica (Keller): holotype

fragment BMNH 1908.9.24.118, 30 x. L, MNHN DIVS6, 120 x.

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 7. A-H, Scanning electron micrographs of spinorhabd I microscleres of Diacarnus spp. A, Diacarnus
spinipoculum (Carter), holotype BMNH 1846.10.14.174. Scale: 21.2um. B, Diacarnus bellae sp. nov., BMNH
1994.5.22.11. Scale: 18.5pm. C, Diacarnus erythraeanus sp. nov., MNHN DJV52, Red Sea. Scale: C1: 12.6um;
C2, C3: 9.1ym. D, Diacarnus levii sp. nov., D1 = holotype QMG305009, D2 = BMNH 1994.5.22.8. Scale:
21.1um. E, Diacarnus ardoukobae sp. nov., holotype QMG305010. Scale: 9.3um. Е, Diacarnus bismarckensis
sp. nov., holotype QMG305011. Scale: 13pm. G, Diacarnus tubifera sp. nov., BMNH 1994.5.22.10. Scale:
15.1um. Н, Dicarnus megaspinorhabdosa sp. nov., holotype QMG305013. Scale: H1: 16.2um; H2: 28рт.

REVISION OF DIACARNUS AND NEGOMBATA

Negombata de Laubenfels
Negombara de Laubenfels, 1936: 159

TYPE SPECIES

Latrunculia corticata Carter, 1879: 298 (by
subsequent designation, de Laubenfels, 1936:
159)

DIAGNOSIS

Erect, massive, lobate to elongate, finger-form-
ing, branching. Compressible, with an extremely
smooth surface through which large pores are
clearly visible. Megasclere skeleton consists of a
uniform, elongate, to square-meshed reticulation
of clear spongin cored fairly irregularly with
thick, slightly curved oxea with strongylote ends,
and a clear hollow axial canal. Ectosome, with
thin fusiform wavy oxeas in tracts that fan within
a thick collagenous ectosome, ultimately forming
surface brushes. Microscleres, irregular
spinorhabds in two sizes, largest very irregular,

ick, without a distinct shaft, smaller, with a
distinct straight or curved shaft with spines of
uneven length arranged roughly in two central
whorls and two terminal bunches, terminal spines
commonly double. Immature microscleres,
straight rods in which one or more spines from
apices are strongly recurved, disposed predomi-
nantly on very outside of ectosome, but can also
be very dense within choanosome.

Negombata corticata (Carter)
(Figs 2, 6J)

Latranculia corticata Carter, 1879: 298; Keller, 1889:
40!

MATERIAL EXAMINED

HorprvPE: — Latruünculia соггісага BMNH
1840,5,6.56-58; "from the Red Sea", three dried frag-
ments from the same specimen (Fig. 2).

DESCRIPTION

Carter (1879) described the sponge as being

"erect, solid, tobate. apparently subsessile”.
Type specimen consists of short, ridged, broad
fans of different lengths with a restricted base of
attachment (Fig. 2). Colour, yellowish white in
dry sponge. Surface texture of dry holotype, fi-
brous, roughened due to shrinkage, Carter (1879)
described the fresh sponge as "chondroid" and
“smooth as varnish to the unassisted eye" with
pores 20pm diameter and 741m apart.

Skeleton, Reconstitution of the dry holotype
allowed examination of the skeletal organization

495

of this sponge. The skeleton consists of a central
axis of rectangular meshes formed by spongin
fibres, 300-600um diameter, not clearly differen-
tiated into primary and secondaries (Fig. 6J).
Large oxea are embedded within the fibres, and
also cecur interstitially. Megascleres are not ar-
ranged uniformly within fibres, but rather, are
scattered singly or in groups, and oblique or oc-
casionally perpendicular to fibre axis, An ectoso-
mal skeleton of wavy oxea is present, but this has
collapsed in the holotype, tendering the arrange-
ment difficult to determine. Microscleres,
densely packed on sponge surface and also in
choanosome, almost obscuring choanosomal fi-
bres,

Spicules. Megascleres. I. Oxeas of main fibro-
reticulation: thick, straight or slightly curved with
rounded strongylote ends, centrally thickened
with a distinct hollow axial canal: 346(317-384)
x 10-1 2um, n=30; П. Ectosomal oxea: thin, fusi-
form irregularly curved, wavy, disposed in sur-
face plumose brushes: 422 (394-451) х 2.4-6,1m,
n=30.

Microscleres, 1; Irregular, shaft rarely distinct,
overall shape of microsclere straight, curved
slightly or spiralled, spines of uneven length ar-
ranged roughly in two central whorls and two
terminal bunches, terminal spines commonly
double: 26(19-31)um, n230; maximum width 14-
19m, shaft width 3-6um, n=10; П: Extremely
irregular spinorhabds, almost oval when heavily
centrally thickened, shaft indistinct and spins tn-
evenly distributed along sides of the spicule:
35(26-43)um, n230, maximum width 17-24um,
shaft width 7-1011m, nz M0.

SUBSTRATE, DEPTH RANGE, ECOLOGY
There js no information given in Carter (1879)
on the ecology nr habitat of this species.

GEOGRAPHIC DISTRIBUTION

Carter (1879) noted that the sponge was sup-
plied hy а dealer who stated that the sponge came
from the Red Sca.

REMARKS

Examination of a spicule preparation of the
holotype of Latrunculia corticata Carter (BMNH
1340.5.6.56-5%) indicates that this sponge differs
considerably from species recognised within
Diacarnus s.s., Sigmosceptrella $.8., oar
Latrunculia s.s. The megascleres of Negombata
are two forms of wavy oxea, instead of sub-
tylostrongyles as in Diacarnus and styles in
Latrunculia. Examination of the reconstituted

MEMOIRS OF THE QUEENSLAND MUSEUM

P 5 HM

— а; = \ /
all nA. B

REVISION OF DIACARNUS AND NEGOMBATA

skeleton of the lype specimen reveals
megascleres embedded in spongin forming a rec-
tangular mesh from which short ectosomal tracts
of thinner, wavy oxeas arise to form plumose
brushes within the ectosome. In Diacarmus the
fibres are clearly differentiated into very large
primary fibres, with smaller secondary fibres giv-
ing rise to dendritic tertiary fibres in a plumose
umbelliform arrangement, Although the
microscleres of Negombata are also spinulate,
they differ from those of other latrunculiid gen-
era. The largest category are so irregular they
look like spined Jumps of silica, the smaller cate-
gory are also irregular with dense, frequently
double terminal spines that are arranged only
vaguely in whorls. The overall spicule is fre-
quently curved or spiralled, with a distinct shaft
only rarely visible, The smaller category of
spinorhabds derive from straigh! rods in which
one or more spines from the apices are strongly
recurved. Carter’s illustrations of regular
microscleres are thus misleading.

Latrunculia purpurea Carter, from Bass Strait,
southern Australia, was also regarded by de
Laubenfels (1936) to be a species of Negombata.
However, Latrunculia purpurea appears to be a
valid species of Latrunculia. Carter (1881) de-
scribed a "flat, compressed, circular, thin, cake-
like or fungiform" sponge, brown-purple in
colour, with a "ragged and proliferous upper sur-
face". There is no mention, however, of à
"chondroid dermal region" and "fibroreticulate
internal structure” described for Negombata
corticata, rather, the intemal structure is "сот-
pact and densely spiculous." The microscleres of
Latrunculia purpurea are typical of Latrunculia
as the two inner spined discs are disposed towards
one end of the spicule, and the microscleres are
disposed "to the circumference on the upperside
and darker portions, arranged perpendicularly in
juxtaposition with the spinous disk of one end
outwards (Carter, 1881)." The megascleres, how-
ever, are described and illustrated as being oxcote
as in Negombata certicata and unlike the poly-
tylote or uniform styles of Latrunculia. However,

497

without examination of the type material, it is
impossible to speculate further on the affiliations
of this specimen,

De Laubenfels (1936) also considered the pos-
sibility that Negombata was synonymous with
Negombo Dendy. It is clear from examination of
the type species Negombo tenuistellata Dendy
(BMNH 1907.2.1.28) that this is not the case; the
megascleres of Negombo tenuistellata are gently
curved large styles arranged in vague longitudi-
nal to reticulated tracts within thin hollow-walled
tubes, ànd the microscleres are different, as al-
ready shown (see remarks for Diacarnus
spinipoculum).

Negombata magnifica (Keller)
(Figs 3K, 3L, 4C, 6K, 6L, 8G)

Latrunculia magnifice Keller, 1889: 402

MATERIAL EXAMINED

FRAGMENT OF HOLOTYPE: Latrunculia magnifica
BMNH 1908.9.24.118 (fragment from the Berlin Mu-
seum) from Suükin, Sudan, Red Sea,

ADDITIONAL MATERIAL: MNHN DJV56: Musha Js-
land, Djibouti, 24m, 11°42.50°N, 43'08.45' E. col-
lected by J. Vacelet, 23 Jantary 1985 (M 3), (fragment
deposited as BMNH 1994.5 22.18)

DESCRIPTION

Keller (1839) described the sponge as having
many long cylindrical digits arising from a short
spreading stalk. A specimen 30cm high, was de-
scribed as having 25 branches of equal length
with a uniform diameter of 6-10mm. Although
the digits do not commonly divide to form
branches, occasional short stumps or incipient
branches are seen. The texture was described as
compressible bul firm and elastic. The surface of
the sponge was completely smooth, slightly
shiney, with a translucent surface, and punctured
with regularly spaced pores 100-150um diame-
ter. Oscules were uncommon, tear-shaped and
raised slightly on collars. The colour in life was
described as deep red-orange which remained in
the preserved specimen for a considerahle time

FIG, 8, Scanning electron micrographs of spinorhabd I microscleres of Diacarnus spp, Negoribata magnifica
(Keller), and acanthose microrhabds of Negombo tenuistellata Dendy: A, Diacarnus spinipoculum (Carter),

holoty

BMNH 1846.10,14.174. Scale; А1: 24.2um; A2: 28.2um. B, Diacarnus bellae s

, nov., BMNH

1994.5,22.11, Scale: 18.20 m. C, Diacarnus ardoukobae sp. nov., holotype QMO305010. Scale: C1: 21.3; C2.
16,6um; C3: 14. 640m, D, Diacarnas bismarckensis sp. nov., holotype QMG305011, Scale: 213m. E,
Diacarnus tubifera sp. nov., BMNH 1994.5.22,10. Scale: El: 18.2рт; E2: 24.2um. F, Diucarnws
megaspinorhabdosa sp. nov. holotype QMG305013. Scale: F1: 341m; F2: 30. Зит; F3: 34pm. G, Negornbare
magnifica (Keller) MNHN DJV 56, Scale; GI: 9.]im; G2: 12,5um. Н, Negombo lenuistellata Dendy 1905,
holotype BMNH 1907.2.1.28, acanthose microrhalxl microseleres. Scale: 5.20 m,

498

before eventually fading. The ectosome was more
intensely coloured than the interior of the sponge.
Embryos of the parenchymella type were present
and large (825-1125jum diameter),

Skeleton. Details of gross morphology, histol-
ору, arrangement of the choanosomal skeleton,
and spiculation were accurately described and
illustrated by Keller (1889), However, for consis-
tency these are re-interpreted here with details of
the ectosomal skeleton not evident 1n the holo-
type Negombata corticata. The skeleton consists
of a central axis of thick strongylote oxeas bound
into an irregular round-meshed reticulation with
abundant stratified spongin. Interstitial
megascleres are common. Primary fibres cannot
be clearly distinguished from secondary fibres
which average 125um diameter (range: 60-
300um). Keller (1889) notes and illustrates the
presence of a "parasitic" algae, thought to be a
species of Calithamnlon, living within the spon-
gin fibres. This was not observed in any of our
specimens. A distinct ectosomal skeleton is pres-
ent and consists of regularly and closely spaced
compact spicule tracts 30-70рт wide which di-
vide and diverge towards the ectosome where
they form fanned brushes (Fig. 6K). Megascleres
of the ectosomal skeleton are wavy fusiform
oxeas rather than the shorter thicker strongylote
oxea embedded in the spongin fibre. The tips of
these spicules protrude through the dermal mem-
brane in patches. Microscleres are common
within the choanosome and present as a distinct
layer in the upper third of the ectosome. The
ectosome is distinct, collagenous, approximately
180m deep (range 240-430; т), and perforated
by vertical poral canals 150-200j.m wide which
lead tô subdermal lacunae.

Spicules. Megascleres. Y: Oxeas of main fibro-
reticulation, thick with hollow central axis,
straight or slightly curved, with shghtly restricted
strongylote ends: 346 (298-385) x 10(4.8-12)ит,
n=30; II; Ectosomal oxea, fusiform irregularly
curved, wavy, disposed in surface plumose
brushes: 435 (403-490) x 1-3um, n=30,

Micrascleres, I, Slender, straight, curved
slightly or spiralled, spines of even length and
arranged in two discrete central whorls with ter-
minal bunches, terminal spines commonly dou-
ble, occasionally recurved, shaft clearly
distinguished between spines: 23(19-26)]m,
n-30; maximum width 10-12um, shaft width
3pm, n=10 (Fig, ЗК, 3L); П: Spicule very stout
with short scattered irregular spines, spines ab-
sent in some parts of the microsclere: 33(24-

MEMOIRS OF THE QUEENSLAND MUSEUM

41)um, n=30; maximum width 22-261um, shaft
width 7-201 m, n=10 (Fig. 8G).

SUBSTRATE, DEPTH RANGE, ECOLOGY

The sponge was abundant on coral reefs and
vertical coral walls, and is one of the most prom-
inent sponges in the Gulf of Eilat between depths
of 3-30m (Kashman et al.. 1980). Very large
embryos were abundant in à sponge collected in
July of 1985 (Fig. 6L). Itis not known whether
these ure pigmented bright yellow-orange as in
Diacarnus.

GEOGRAPHIC DISTRIBUTION
Suákin, Sudan, Red Sea; Djibouti, Gulf of
Aden (Fig. 4C)

REMARKS

Negombata magnifica is distinct from N.
corticata in gross morphology, N. magnifica
forming long, cylindrical, whip-like branches,
the later being a clump of broad ridged fans.
Histologically, the species are rather similar, but
with discemable dimensional and morphological
differences between the megascleres and
microscleres. The later are extremely dense in the
choanosome of №, corticata. The spongin skele-
ton of N, corticata is more robust.

DISCUSSION

Diacarnus and Negombata are distinct within
the Latrunculiidae as they possess a fibro-reticu-
late skeleton with an unusual umbelliform
"ectosomal" skeleton developed to a greater ог
lesser degree, and a highly collagenous mesohyl
matrix structure with tough fibres which superti-
cially resemble vertcbrate ligaments in soft tis-
suc. Diacaraus is remarkably homogenous with
the skeletal arrangement, huge fibres and spicule
dimensions of all species being similar. We have
emphasised the gross morphology of the sponge,
coupled with the presence or absence of a second
larger category of microsclere, and the morphol-
ogy of these spicules, in recognizing new species.
The disposition of these microscleres throughout
the sponge, and spicule and fibre dimensions are
less reliable characters for species differentiation.
Diacamus spinipoculum and D. tubifera are both
vasiform but the former is barrel-shaped rather
than tubular. These two species are further sepa-
raled on the presence of large spinorhabds in D,
tubifera, in addition to very abundant dendritic
tertiary spicule tracts which arise from well
within the choanosome of this species. Diacarnus

REVISION OF DJACARNUS AND NEGOMBATA

bellae from Micronesia and D. levii from New
Caledonia are very similar but are separated on
gross morphology and spicule dimensions. The
three ramose species D. erythraeanus, D.
ardoukobae, and D. bismarckensis, are separated
geographically, and the two Red Sea species are
further separated by the presence of large
spinorhabds in D. ardoukobue. Diacarnus
megaspinorhabdosa has the largest megascleres
Of all species thus far described, and highly diag-
nostic spinorhabds,

Although there are many similarities between
Negombata and Diacarnus, there are several
major differences which separate them, including
the presence of diactinal megascleres and a regu-
lar üght-meshed "axial" fibrous skeleton in which
ibe fibres are only lightly cored by megascleres.
The skeletal arrangement of Negombata superfi-
cially resembles that of several myxillid, raspailid
and microcionid poecilosclerid genera which em-
phasise an axial fibroreticulation and cctosomal
brush-forming skeleton. In Diacarnus, the fibres
are packed regularly with spicules with no visible
spongin surrounding them. Negombata corticata
and N. magnifica are differentiated primarily on
gross morphology, spicule differences, and pos-
sibly the development of the ectosomal skeleton.

Sponges previously identified as Latrunculia
magnifica (Keller) from the Red Sea uniquely
contain compounds known as latrunculins (Nee-
man etal, 1975; Kashman etal., 1980). With the
transfer of this species to Negombata here, the
latrunculins become à potentially diagnostic
chemical marker for sponges of the genus Nego-
mbate, Several latrunculin - containing voucher
specimens from the Philippines and Indonesia
supplicd hy M-K. Harper (Scripps Institute of
Oceanography} and the Coral Reef Reseach
Foundation, were examined as this report went to
press, While these sponges are clearly related
morphologically to the two known species of
Negombata, there are considerable differences,
requiring the examination of further material be-
fore a complete identification can be made. Spe-
cies identified as Latrunculia brevis and L
conulosa from Australia (in Buller & Capon,
1991; 1993), Sigmosceptrella laevis (Alberici et
al., 1982), and Diacarnus bellae sp. nov. (F. J.
Schmitz, D. J, Faulkner, pers. comm.), contain
norseésterterpene peroxides, Confirmation of pre-
vious identifications for the specimens whose
chemistry is known and has been published, is
presently being carricd out (Kelly-Borges and
Мает, in prep.).

400

The possession of large tracts of diverging
megascleres, megasclere brushes at the surface,
spined acanthomicrorhabds, and large bright yel-
low larvae, in Diacarnus and Negombata, is rem-
iniscent of Sigmosceptrella Dendy. Examination
of the type species of Sigmosceptrella, S.
quadrilobata (BMNH 25.11.1.1641), and
Spirastrella (=Sigmosceptrella) fibrosa Dendy
(1897), which Dendy (1922) regarded as the
тое’ type species of the genus Sigmosceptrella,
reveals several characteristics which distinguish
Sigmosceptrella from Diacarnus and Nego-
mbata.

The largest microscleres of Sigmosceptrella are
very similar to those of Diacarnus and Nego-
mbata; but the spines of the two inner whorls of
the rhabd axis are longer than those of the termi-
nal whorls, which are more or less in the same
plane as the axis of the spicule, and they are closer
to the terminal whorls than they are to each other,
imparting a dumbell appearance to the spicule
(Fig. 3D, 3F. G). The spinorhabds of Diacarnus
and Negombata are more regular with spines of
equal length in whorls that are separated equidis-
tantly, in most species, along the axis of the
spicule (Fig. 31, J, L). In all specimens of
Sigmoscepirella examined, these microscleres
are packed in confusion in a dermal crust at the
surface (Fig, 9A,B) of the sponge, rather than
scattered in a band within a broad collagenous
detachable ecrosome, as 1n Diacarnus and Nego-
mbata.

The developmental stages of the spinorhabds of
these three genera also differ. The rhabds of
Sigmosceprrella are distinctly sigmoid in shape
(Fig. 3D, E, G). These were illustrated by Dendy
(1905: Plate 18 Figure 4c: Dendy 1921: 122) who
regarded these as the major diagnostic character
for Sigmosceprrella. These are not to be mistaken
for the "true" sigmas which can also be found in
the BMNH histological slide of the holotype.
There were also small anchorate isochelae and
tylostyles in the section and thus the lot is obvi-
ously foreign. The earliest protorhabd forms of
Sigmosceprrella and Negombata are rather more
similar to cach other than they are to the pro-
torhahds of Diacarmus, which are simply spinul-
ate as in the mature spicule (Fig. ЗН). The earliest
spines of the rhabds of Negombara and
Sigmosceptrella are strongly recurved spines
with kinked tips which eventually become the
spines along the shaft. However, where the shaft
of Sigmosceptrella becomes sigmoid (Fig. 3B, E)
(Dendy, 1921), those of Negambata remain
straight (Fig. 3K).

500

FIG. 9. A-D, Skeletal arrangements of Sigmoasceprrella and Latrunculia, A,
Sigmosceptrella quadrilobata Dendy, holotype BMNH 25.11.1.1641, In-
dian Ocean, 30x. B, Sigmosceptrella sp., Q66C-2779, Sisters Reef, Murray
Reefs, 3m, Wambro Sound near Rockingham, Perth, Western Australia,
32° 21.38 115° 41,3'E, 30 x, C-D, Latrunculia sp., Q66C-2463, Horse-
shoe Reef 3km WNW of Margaret Brock Lighthouse, 18m, Cape Jaffa,
Kingston, South Australia, 36°56.8"S, 139*35.0'E, C, the skeleton is a Podospongia, are in complete con-

whispy reticulation of loose tracts of styles, with a layer of erect discale
microrhabds at surface of sponge, 30 x, D, single discorhabd with terminal g
ectodermal membrane, above a layer of

spines embedded beneath the
tangential styles, 2000 x.

The megascleres of Sigmosceptrella are ar-
ranged in plumose tracts which arise from the
base of the sponge, branching continuously to
form very fine fibres which terminate in brushes
at the surface (Fig. 9A,B), unlike the
plumoreticulate arrangement of huge compact
spongin-bound fibres with apical umbells, as in
Diacarnus. The fibres of Diacarnus are clearly
visible to the unaided eye, and are separable from
the mesohy! matrix, resembling ligaments, There
is also no continuous dermal palisade of
megascleres at the surface of Sigmosceptrella.

The family Latrunculiidae has an unstable his-
tory and has been switched between the orders
Hadromerida (¢.g., Topsent, 1922: Dendy, 1922;
Bergquist, 1978), and Poecilosclerida (e.g. Lévi,
1973; Van Soest, 1984, see Hooper, 1986). As it
is currently perceived, the family is heteroge-
neous, sharing features that belong to both orders,

MEMOIRS OF THE QUEENSLAND MUSEUM

Diacarmus, Negombata and
Sigmosceptrella incubate huge
parenchymella larva, and embryos
Sum of diameter have been
found in a Mediterranean species
of Latrunculia (Vacelet, 1969),
while all known hadromerid
sponges are oviparous. However,
— in contrast to the obvious affinity
of these genera with poecilosclerid
sponges Which incubate larvae,
Diacarnus and Negombata, in par-
ticular, are similar to many
hadromerid genera such as Tethya,
Spirastrella and Polymastia,
which emphasise heavily collage-
nous tissue, particularly in the cor-
lex.

Although there is no clear reso-
lution of the affinities of these gen-
era with other demosponges,
several patterns have emerged
from this study of morphological
characters. It is quite clear that
Diacarnus, Negombata and
Sigmosceptrella are more closely
related to each other than they are
to Latrunculia, type species of the
family Latrunculiidae. The skele-
tons of these three genera, includ-
ing another latrunculiid

trast to that of Larrunculia, all gen-

era emphasising a complex
plumose fibro-reticulation, while
the skeleton of Latrunculia s.s. is а
wispy reticulation of very loose
tracts of styles with a tangential surface layer of
megascleres (Fig. 9C). Although the morphology
of the acanthomicrorhabd microscleres in
Sigmosceptrella, Diacarnus, and Negombata ap-
proach those of Latrunculia in general form, the
microscieres of the later are discate rather than
spinulate. The microscleres of Latrunculia s.s.
are arranged in a peculiar erect palisade of
microscleres with their terminal disc embedded
beneath the ectosomal membrane, the remainder
of the microsclere free of tissue outside the
sponge (Fig. 9C, D). In Sigmoscepirella and
Negombata, microscleres are densely packed in
the outer cortex, while in Diacarnus these are less
dense and contained in a band mid-cortical band.

Morphological data suggest that Latrunculia
s.s., and possibly Barbozia with its palmate
isochelae, may be more closely related to

REVISION OF DIACARNUS AND NEGOMBATA

iophonid poecilosclerids such as Zyzzya, as they
share several features such as a tangential surface
layer of тераѕсјегеѕ, erect surface spicules (al-
(hough in Zyzzya these spicules are tangential

lotes or strongyles, and acanthoxeas, respec-
tively), deep brown or green colouration, and in
the possession of fistules, It seems likely from
morphological data, also, that Diacarnus, Nego-
mbata, Sigmoscepirella, and Podaspongia,
should not be included in the family
Latrunculiidae. Although it is clear that these
genera are similar in some respects lo some
hadromerids (collagenous tissuc, radial symme-
try in Podospongia, ctc.), the vast majority of
characters are shared with poecilosclerids (plu-
mose-reticulate fibre skeleton, chemistry, repro-
duction), suggesting that this group also
constitutes a poecilosclerid family, but separate
from the Latrunculiidae, rather than a hadromerid
family. However, as there is no clear resolution
of this hypothesis we retain them as incertae sedis
within the Family Latrunculiidae until further
data is available. Secondary metabolite datà and
DNA sequence data acquisition and analysis are
currently in progress towards this end {Kelly-
Borges, unpublished data).

Diacarnus 1s predominantly a shallow-water
genus with a broad Jatitudinal distribution within
tropical West Central Pacific marine environ-
ments (Fig. 4). Diacamus spinipoculum is very
widely and sparsely distributed in temperate
south-eastem Australia and tropical Micronesia
and Fiji, The longitudinal distribution is however,
remarkably disjunct, as two species of Diacarnus
also occur in the Red Sea (D. erythraeanus and
D. ardoukobae) with the two known species of
Negombata, Except in the Red Sea and Gulf of
Aden, Diacarnus has not been recorded further
west than the Philippines despite extensive recent
sampling within the broad Indo-Pacific region.

This disjunct Red Sea-Indo-Pacific distribution
has been indicated for several Indo-Pacific spe-
cies of sponge, including well known species
such as Psammaplysilla purpurea, Hyrtios
erecta, Dysidea herbacea, — Phyllospongia
papyracea, and Theonella swinhoei. Sixty per-
cent of the Red Sea records in Lévi (1958) were
of species known previously only in the Indo-Pa-
cific. A number of Red Sea coral reef fishes show
a similar level of differentiation from related spe-
cies found in the Indo- West Pacific (Allen, 1979;
Steene, 1977), Histological examination of the
obviously very closely related sister-species
groups which contain D, erythraeanas in the Red
Sea, and D. leyii in New Caledonia, and D.

50!

drdoukobae in the Red Sea and D, bismarckensis
in the Indo-Pacific, reveal small but consistent
differences through geographic separation, 11 is
likely that subtle differences will also be found
for the species indicated in earlier literature to
have a Red Sea / Indo-Pacific distribution, espe-
cially those genera which have few reliable char-
acters lo used іп species separation such as
Psammaplysilla, Dysidea, and Hyrtios. It is be-
coming increasihgly obvious that species pre-
viously thought to be “cosmopolitan” actually
consist of cryptic sibling species (Bergquist &
Kelly-Borges 1991; Hooper et, al., 1992; Kelly-
Borges & Bergquist, 1994; Solé-Cava et al.,
1991},

KEY TO SPECIES OF DIACARNUS

la. Large tubular or spherical sponge wilh deep
wide apical atrium . 5... earns
ib, Massive subspherical or digitate sponges, soli-
tary orcedlescemt .....„........- К
lc, Trregular sprawling branches
2a. ‘Tube or vase-shaped sponge, up to 80cm long
and 30cm wide, 8-15cm apical atrium, surface
with labate projections, extremely abundant den-
Gritic ectosomal fibres and large irregular
spinorhabds, found in Micronesia, common in
the Bismarck Sea from Madang to Kavieng,
New Ireland, and rarely in southern Papua New
Guinea. ,.... . Dicamus tubifera sp. mòv.
2b. Barrcl-shaped sponge, up to 50cm high and
40cm diameter, deep, wide, apical átrium, slen-
der megascleres and only very thin microscleres
of the smallest category that can be easily over-
looked, found in Micronesia, Fiji, and in south
Australia. . . Diacarnus spinipoculum sp, nov.
3a. Spherical sponge, 6-8cm diameter, frequently
joined to adjacent sponges to form a small
clump, oscules white-rimmed and arranged api-
cally in а cluster or serially, slender megascleres
and thin microscleres that can be easily over-
looked, only recorded from Micronesia so far
Li А Ht Diacarnus bellae sp, nov,
3b. Thick erect digits or lobes, 3-4.5cm in diameter,
anastomosing to form a large sprawling mass, 0s-
cules on ends of digits, slender megascleres and
only very thin mieroscleres of the smallest cate-
gory that can be easily overlooked, found in
New Caledonia and rarely on the central Great
BarrierReef.,.... Diacarnus levii sp, nov.
3c. Thickly encrusting to lobate, huge spinorhabds
present. Wide, sparse distribution - found in
southern Papua New Guinea, Philippines and
Madang coast of Papua New Guinea... .. -
‚+. Diacarnus megaspinorhabdosa sp. nov.
4a. Large regular spinorhabds present, found on
Madang coast, and west coast of New Ireland,

502
Pnpva New Guinea |... eese
V da^ add a Diacarnus ‘bismarckensis sp ov,
db, Found in Red Seå . .............. 5
5a. Large spinorhabds present... .... i.

‚д ied б Ріасатиѕ ardoukobae sp, nov,
Sb, Large spinorhabds absent. ..........
Lee 02 Diacarnus eryrhraeanus sp, nov.

ACKNOWLEDGEMENTS

We thank personnel of CRRF, Micronesia,
AIMS, Townsville, and ORSTOM, New Caledo-
nia, for collection of samples, and the personnel
of Motupore Island Research D ent, Uni-
versity of Papua New Guinea, for field and labo-
ratory support whilst in Papua New Guinea, We
thank Clive Wilkinson for specimens from the
Red Sea, and Mary Kay Harper, SIO, La Jolla, for
providing a specimen from Pohnpei Island, Mi-
сгопезіа. The specimens from Djibouti were col-
lected during the expedition "Ardoukoba" and
{юзе from Djeddah were collected during a trip

ized by the Faculty of Marine Sciences in
Djeddah. We thank Klaus Kelly-Borges (prepa-
ration of material for histological examination,
and translation of Keller (1889) from German to
English), Clare Valentine (location of speci-
mens}, Peter York and Phil Crabb (micro and
macro photography), all of the NHM, Londen.
We thank CRRF, Micronesia, for underwater
photography of Micronesian, Papua New Оџіп-
ean, Fiji and Philippine specimens of Diacarnus,
and Chantal Bézac and Isabelle Mahieu, for their
help with SEM microphotography. Thanks to
Professor Ralph-Heiko Mattem for discussions
on the chemistry of these sponges. This is Harbor
Branch Oceanographic Institution Contribution
No. 1090.

LITERATURE CITED

ALLEN, G. R. 1979. Butterfly and Angel fishes of the
оз Volume 2. (Wiley and Sans, New York),

ALBERICCI, M.. BRAEKMAN, J, C., DALOZE, D.,
& TURSCH, B. 1982, Marine Invertebrates.
ХІМ, The chemistry of three norseste
peroxides from the sponge Sigmioscepiretia levis,
Tetrahedron 38; 1881-1890.

BERGQUIST, P. R. 1968. The Manne Fauna of New
Zealand: Porifera, Demospengiac, Part |
(Tetractinomorpha and Lithistida) New Zealand
Department Scientific and Industria]. Research
Bulletin] 88(New Zealand Oceanographic Enstitu-
tion Memoirs 37) : |05pp.

1978. Sponges, London: Hutchinson: 268pp

BERGQUIST, P.R. & KELL'Y-BORGES, M. m An
evalualion of the genus Teriye (Porifera:

MEMOIRS OF THE QUEENSLAND MUSEUM

Demospongiae: Hadromerida) with descriptions

of new species from the Southwest Pacific, The

Beagle, Records of the Northern Territory Mu-
seum of Arts and Science, 199] 8:37-72.

BOCAGE, J. V. B. DU. 1869. Eponges siliceuses nou-
velles de Portugal et de l'ile Saint-Iago (archipel
de Cap vert). Jornal de Scienzas Nationale
Academie Lisbonne 2: 159-162.

BURTON, M. 1934. Sponges. Scientific Report on the
Great Barrier Reef Expedition 4(14):513-621.

BUTLER, M. 5. & К. J, САРОМ, 199]. Norterpene
dienes from an Australian marine sponge
Latrunculia brevis. Australian Journal of Chemis-
try 44:77-85,

1993. Trunculin F and Contrunculin A and B; novel
oxygenated norterpenes from a southern Austra-
lian marine sponge, Latrunculia conulosa, Aus-
tralian Journal of Chemistry 46: 1363-1374.

CARTER, H, J. 1879, On Holostrella, a fossil sponge
of the Carboniferous era, and on Hemiasterella, à
new genus of recent sponges. Annals and Maga-
zine of Natural History (5)3:141-150.

1881. Supplementary report on specimens dredged
up from the Gulf of Manaar, together with others
from the sea in the vicinily of Ihe Basse Rocks
and from Bass's Straits respectively, presented to
the Liverpool Museum by Capt, Н, Cawne War-
ren. Annals and Magazine of Natural History
(5)7:361-385.

DENDY, А, 1897, Catalogue of the non-calcareous
sponges collected by J. Bracebridge Wilson in the
neighbourhood of Port Phillip Heads. Part 3, Pro-
ceedings of the Royal Society of Victoria 2: 230-
259.

1905, Report on the sponges collected by Professor
Herdman, at Ceylon, in 1902. Report to the Gov-
ernment of Ceylon on the Pearl Oyster Fisheries
ofthe Gulf of Manaar. 3(Supplement 18):57-246,

1921. The tetraxonid sponge-spicule: а study in
evolution. Acta Zoologica 2: 95-152,

1922. Report on the Sigmatotetraxonida collected
by the H.M.S."Sealark" in the Indian Ocean: In
Report of the Percy Sladen Trust Expedition to
the Indian Ocean in 1905, Volume 7, Transac-
tions of the Linnean Society of London, Zoology
18:1-164,

HOOPER, J. М. А. 1986, Revision of the marine sponge
genus Axis Gray (Demospongiae: Axinellida)
from the north-west Australia. The Beagle, Occa-
sonal papers of the Northern Territory Museum
of Arts and Science, 1986 3: 167-189,

HOOPER, J. М. A., САРОМ, R. J., KEENAN, C. P.,
PARRY, D, L. & SMIT, N. 1992. Chemotaxon-
omy of Marine Sponges: Families Microcionidac,
Raspailiidae and Axinellidae, and their relation-

ships with other familes in the orders
Poecilosclerida and Axmellida (Porifera:
Беширов) Invertebrate "Taxonomy 6:261-

KASHMAN, Y , GROWEISS, A, AND SHMUELI.
U, 1980, Latrunculin; a new 2-thiazolidinone

REVISION OF DIACARNUS AND NEGOMBATA

macrolide from the marine sponge Latrunculia
magnifica. Tetrahedron Letters 21: 3629-3632.

KELLER, C. 1889, Die Spongienfauna des Roten
Meeres I, Zeitschrift für wissenschafliche
Zoologie 48: 311-405.

KELLY-BORGES, M., ROBINSON, E. V.,
GUNASEKERA, S., GUNASEKERA, M.,
GULA VITA, N. & POMPONI, S. A. 1994. Spe-
cies differentiation in the marine sponge genus
Discodermia (Demospongiae, Lithistida): The
utility of secondary metabolites as species-spe-
cific markers. Biochemical Systematics and Ecol-
ogy 22(4):353-365,

KELLY-BORGES, M. & BERGQUIST, P. R. 1994, A
redescription of Aaptos aaptos (Schmidt), with
additions to the genus Aapros in northern New
we Journal of Zoology, London 234: 301-
323.

KORNERUP, A. & WANSCHER, J. H. 1961.
Rheinhold Color Atias. (Reinhold Publishing
Corporation: New York). 224pp.

LAUBENFELS, M. W. DE 1936. A discussion of the
sponge fauna of the Dry Tortugas in particular and
the West Indies in general, with material for the
revision of the families and orders of the Porifera,
Publications of the Carnegie Institution, Papers

. from Tortugas Laboratory. 30 (467):1-225.

LEVI, C. 1958. Spongiaires de Mer Rouge recueillis par
la "Calypso" (1951-1952). Résultats scientifiques
des Campagnes de la "Calypso" 3: 3-46

1973. Systématique de la classe des Demospongia-
ria (Demosponges). In "Traité de Zoologie. IIT.
Spongiaires (P. -P. Grassé, ed)", Masson et Cie
:571-631.

NEEMAN, I., FISCHELSON, L. & KASHMAN, Y.
1975. Isolation of a new toxin from sponge
Latrunculia magnifica in the Gulf of Aquaba.
Marine Biology (Berlin) 30: 293-296,

503

RIDLEY, S. O. & DENDY, A. 1886, Preliminary report
on the Monaxonida collected by H.M.S.
"Challenger. Annals and Magazine of Natural
History 5(18): 325-351, 470-93,

1887. Report on the Monaxonida collected by
H.M.S. 'Challenger' during the years 1873-1876.
Vol 20: Pp.1-275, In: Report on the Scientific
Results of the Voyage of the H.M.S, ‘Challenger’
during the years 1873-1876. (Her Majesty’s Sta-
tionary Office: London, Edinburgh, Dublin).

SOEST, К. W. M., VAN 1984. Marine sponges from
Curagao and other Caribbean Localities, Part III.
Poecilosclerida. Studies on the Fauna of Curagao
and other Caribbean Islands 199:1-167.

SOLE-CAVA, A.M., KLAUTAU, M., BOURY-ES-
NAULT, N., BOROJEVIC, R. & THORPE, J.P.
1991, Genetic evidence for cryptic speciation in
allopatric populations of two cosmopolitan spe-
cies of the calcareous sponge genus Clathrina.
Marine Biology (Berlin) 111: 381-386,

STEENE, R. C. 1977. Butterfly and Angel Fishes of the
World. Vol. 1. (John Wiley and Sons: New York).
144pp.

THIELE, J. 1900. Kieselschwámme von Ternate, I.
Sonderabdruck aus den Abhandlungen der

Senckenbergischen naturforschenden Gesell-
Schaft 25(1): 19-80.

TOPSENT, E. 1922. Les mégascléres polytylotes des
Monaxonides et la parenté des Latrunculiines.
Bulletin de I’ Institut Océanographique de Mo-
naco 415: 1-8.

VACELET, J. 1969. Eponges de la Roche du Large et
de l'étage bathyal de Méditerranée (récoltes de la
Soucoupe plongeante Cousteau et dragages).
Mémoires du Muséum National d'Histoire
Naturelle, A, Zoologie 59; 145-219.

OBSERVATIONS OF THE AMETHYST PYTHON
(MORELIAAMETHYSTINA) FEEDING ON RAINBOW
BEE-EATERS (MEROPS ORNATUS). Memoirs of the
Queensland Museum 38(2), 504. 1995:- Observations on
feeding by arboreal snakes are poorly documented. Members

af Morelia are primarily oocturmal, rock inhabiting and/or

athoreal snakes that commonly bask during daylight hours,
Amethyst pythons (Morelia amerhystina) eat a wide variety
of Vertebrates (Cogger, 1992), including birds and mammals
(Wilson & Knowles, 1988),

Amethyst pythons were observed esting rainbow boe-
cars (Merops ornas, length 230-230mm (Lowe, 1989)
measured as the distance from the lip uf the hill to the np of
the tail, rounded to the nearest Smm, and where a bill or tail
is unusually Jong, as with the rainbow bee-eater, its form is
incloded in the length) on Milman Island (11°10.2'S,
143°00,8°E: Great Barrier Reef Marine Park Code 11-007), a
vegetated sand cay Jocated approximately ]]2km soutlneast
from Thursday Island, Torres Strait and 45km northeast From
Drford Ness, Queensland, Australia.

АЙ feedmg observations took place between 0600h and
0900h on 19 March and 21 March, 1994, Up to four pythons
had been observed simultaneously stretched out on the defo-
hated branches of Premna serratifolia, first on 10 March. and
then 18 - 23 March, The snakes appeared camouflaged, their
bodies curving so they blended with branches of the shrub,

The first observation was of an adult rainbow bec-ester
[extended central tail feathers) within the coils of a python al
D645h an 19 March. The python moved approximately 2 m
down the branch before swallowing the bird. This sequence
took approximately 25 minutes. Then the snake moved back
to the top of the branch. Another rainbow bee-eater landed on
the branch above the snake, but flew off as the python moved
towards iL The snake retreated down the branch (approxi-
mately Zm) and curled wp in a fork in the shrub. The followin
day (20 March) the snake did nat appear to move. On 2
March, at 08 15h the same python caught à juvenile rainbow
bee-ealer (central tail feathers not extended). The snake swal-
lowed this bird in 5.5 minuces, ‘The snake was the same one
observed on 19 March, because the bulge made from eating.
the first bird was still visible, and the snake was on the same
branch. After swallowing the juvenile bird, the snake slithered
back up the branch and waited for 40 minutes, apparently
trying to catch another bird. Even though other rainbow
bec-eaters were flying around its head, the python did nol
catch any. Rainbow bee-eaters catch their prey (flying insects)
by "hawking", rumes to perch and batter their pey before
ingesting it (Mac Donald, 1973), This method would bring the
birds into contact with hunting snakes. White-breasied
woodswallows (Artamus leucorhynchus, length 170
180mm), yellow-bellied sumbirds (Nectorinia jugylaris,
length 110-1 15mm), and a spangled drongo (Cirurus
braces: length 280-320mm) (Lowe, 1989) all landed
nearby. These may also be preyed upon. The white-breasted
Weodswallows appeared to swoop avnear the snake ih alarm

Although (hesnake was nol caught and messural, the sre
of the python (1.25-1.50m in total length} observed feeding
on rhe rainbow Dec-caters appeared to be in ihe smaller range
for records on Milman Island. Six females measured had a
total length ranging from 124.5 - 261.0cm and three unsexed
pythons measured had a total length minging from 6040 =
220 km.

Rainbow bee-caters are a migratory species in the western
South Pacific Ocean area. They move from their southern
breeding areas in Australia tà over-winter in Torres Strait and
Papua Now Guinea (Blakers е! al., 1984). Although the dis-
Iribulion of rainbow hee-earers is dependent on the abundance

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1. Summary of Amethyst Python (Morelia
amethystina} measurements collected from snakes caught on
Milman Island, northern Great Barrier Reef, Queensland,
Australia, Sex was determined by examination of spur size
and attempted eversion af hemipeni. ? = Sex was not deter-
mined. Abbreviations: HL-Head length, SVL-Snout length,
VTL=Vent to tail length, TL=Total length. All measure-
ments are in centimetres.

[Dae — [uL |зи |vr. |r. (бех |
[sri | | | вот |
| [uso [mo [imo |r |
[toFeb 1992 | — [23.0 {380 |2610 [Female
(22 ғеһ 1992 | — |1500 faio |I8LO |Female|
|j9Febl992 | Jigso [aso [mmo |v |
(т.Маг 1994 |s5 |1843 |355 |2200 [Female]

35.5
(17101995 | — |1490 |260

of insects (Blakers et al,, 1984), i! is nol known how long the
rainbow bee-eaters stay.at Milman Island. The island supports
a seasonal nesting colony of Torres Imperial Pigeons (Ducula
bicolor, length 380-440mm) (King, 1990), as well as numer

ous pairs of yellow-bellied sunbirds. All of the birds men-
tioned are small enough to be possible food soprces for the
snakes, although consumption of these other species has nol
been observed. When rainbow bee-eaters are not present on
Milman Island, the pythons must prey on other species, Be-
couse, there are no mammals-on Milman Island, considered a
common prey for amethyst pythons (Cogger, 1992), the py-
thans are probably feeding «m other birds, inclading ground
nesting species, found on the island.

ир ГУЗ
К
т
т
8

Acknowledgements

The U.S, Army Corps of Engineers provided a grant fo Dr.
André M, Landry №. of Texas A&M University - Galveston
which assisted K.A,L, with research on the island

Literature Cited

Blakers, M., Davies, S. J. J. Р. & Reilly, P. N, 1984. "The Апаз
of Australian Birds", (Melbourne Uni versiry Press. Vic-
form).

Cogger, Н, О. 1992. "Reptiles and Amphibians of Australia!
(Reed International Books: Chatswood, NSW).

King. B. 1990, Distribution and status ef the Torresian Imper-
ial Pigeon, Ducula bicolor, in North-eastern Queens-
land: Cooktown to Cape York. Emu 90:248-253.

Lowe, К. W, 1989, "The Australian Bird Bander's Manual
(CPP Communications Ltt Fyshwick, ACT).

MacDonald, 1. D, 1973. " Birds of Australia’, (Kyodo Priming,
Co. LIE Tokyo].

Wilson, S. К. & Knowles, D.G. 1988. ' Australia's Reptiles
(Collms Publisher Sydney, NEW h

A А, laap, Gerensland Deparnpsenr af Enviranmenr am
Hermie. PO Rox 5390, Tewnyville, Queensland 4810, Ans-
iral: Present Арета lnstimte: of Marine Life Sriencer,
Tear A&M (бшмеу кїї, 471) Ave Un Bide 205, Galet,
TX 77551 USA; J, D, Miller and D, Pollart, Queensland
Depurineur of nvirownenr aut Heiter, PO Box 539],
Vowessville, Queensland SEJA, Аш инна, f August #995

OBSERVATIONS OF HUMPBACK WHALES (MEGAPTERA NOVAEANGLIAE) ON A

CRUISE TO NEW CALEDONIA AND THE CHESTERFIELD REEFS
PETER C. GILL, ELIZABETH J. EYRE, CLAIRE GARRIGUE & WILLIAM H. DAWBIN

Gill, P.C., Eyre, E.J., Garrigue, C. & Dawbin, W.H. 1995 12 01; Observations of humpback
whales (Megaptera novaeangliae) on а cruise to New Caledonia and the Chesterfield Reefs,
Memoirs of the Queensland Museum 38(2):505-51 ]. Brisbane. ISSN 0079-8835,

During a yacht-based survey in the austral winter of 1992, low numbers of humpback whales
were observed around the main island of New Caledonia, and humpback song was recorded
there. The song, the first recorded in New Caledonian waters, showed similarities to song
recorded off eastern Australia, No humpback whales were detected at Chesterfield Reefs in
the eastern Coral Sea, despite their possible status as a breeding area. Only a small portion
of the Chesterfield Plateau was surveyed, Reports of sightings there in recent years indicate
al Jeast occasional visitation by humpback whales.

Peter C. Gill, Oceanic Research Foundation, c/- 148 Copeland Rd Eas, Beecroft NSW 2119
Australia; Elizabeth J. Eyre, Marine Mammal Department, Taronga Zoo, PO Box 20,
Mosman, NSW 2028 Australia; Claire Garrigue, ORSTOM. BP A5, Noumea, New Caledo-
nia; William Н. Dawbin, Australian Museum, 6-3 College St. Sydney, NSW 2000 Australia;

15 June 1995,

Group V humpback whales feed in Antarctic
Area V (1307 E-170" W) during the austral sum-
mer and segregate into tropical breeding areas in
the southwest Pacific during winter (Dawbin,
1966). Catch data of 19th century American pe-
lagic whalers indicated wintering, and presum-
ably breeding, concentrations around Tonga, Fiji,
and the Chesterfield Reefs in the Coral Sea
(Townsend, 1935), French whalers killed hump-
backs in the Loyalty Islands of New Caledonia,
as well as at the Chesterfield Reefs (du Pasquier,
1982). Many American whalers also visited parts
of New Caledonian waters during the 19th century
{Langdon, 1984). Humpbacks were marked by
Dawbin using "Discovery" tags around Vanuatu
and the Loyalty Islands during the late 1950s, but
nomarks were recovered (Dawbin, 1964), Recent
incidental sightings data, unreported at the oulset
of this study, indicate that humpback whales,
including newborn calves, are now frequently
seen in New Caledonian waters during winter and
spring (Garrigue and Gill, 1994), There has been
speculation, based on Townsend's (1935) charts,
as {о whether Chesterfield Reefs are still a hump-
back breeding destination (Dawbin and Falla,
1949: Paterson, 1991).

This paper describes a yacht-based cruise dur-
ing the austral winter of 1992, initially to investi-
gate whether humpback whales migrate to New
Caledonia and the Chesterfield Reefs. While in
New Caledonian waters the study focused on
humpback song, never recorded in the region, for
companson with other areas.

METHODS

The 15-metre cutter "Iniquity" was equipped
with GPS navigation, depth sounder (max. depth
100m), radar, weatherfax, wind instruments and
seawater thermometer. With a minimum of 2
observers. continuous visual monitoring was caf-
tied out (except when interrupted by sailing du-
ties) from deck 2-3m above sea level (distance to
horizon 5.6-6.7km) while the vessel was making
way. In sheltered lagoon waters where whales
were more likely to be encountered, when the
vessel's motion allowed, an observer was posi-
tioned up the mast 12m a.s.1, (distance to horizon
13.5km).

"Iniquity" departed Coffs Harbour, New South
Wales on 7 July 1992, and sailed northeast to
Noumea, Nearly 4 weeks were spent in New
Caledonian waters from 14 July, and the passage
westward across the Coral Sea to Queensland
took place during 11-31 August (Fig. 1). This
timing allowed the vessel to be in these areas
around the estimated peak of the east Australian
humpback breeding season (Simmons and
Marsh, 1986; Paterson, 1991).

Humpback song may be detected over tens of
kilometres (Cato, 1991), a much greater range
than visual observation permits. It is commonly
heard in breeding areas and on coastal migration
routes, even in darkness. On a previous yacht-
based study in Australian waters, humpbacks
were detected 3 times more often by song than by
sight (Dawbin and Gill, 1991). For these reasons

506

MEMOIRS OF THE QUEENSLAND MUSEUM

Marian Rf

2055
Kenn Ri, PLATEAU

* Saunas КЇ

AUSTRALIA

[
А

|

}

agai RI

CHESTERFIELD ,

South a RÍ

Great Nonh Lagoon
“a
F,
4,
tr )
Quvea % 1
v Lifou Ял,
RES. ©

NEW З ® ` ‘eu, Mare

е А. У
m n

Soulhesst Laguun

LEGEND
—»— Track of vessel
ww Reef

P О Listening station
(no whales heard)

' Pá A — Humphacks seen
* Middleton KI Ф 7 Humpbacks heard
305 Срб» Harbour
DL aaa 3 200
Nautical miles

/ Lord Howe Is. (1 ——À— —— 100

/

150F

Kilametres

160E 170E

FIG. 1. Map showing the route taken by the vessel, locations of humpback whale sightings, and listening stations.

acoustic monitoring was regarded as the primary
detection method for this study.

Hydrophone soundings were taken for about 10
min every 3-4 hours while at sea (approximately
every 30km) when conditions allowed, in order
to ensure that the same animals were not detected
at more than one listening station. Soundings
were taken not only near coastlines, islands and
reefs, but also distant from land, in order to ascer-
tain possible migration routes (Clapham &
Mattila, 1990). Monitoring was also conducted
when whales were sighted. A minimum of 20 min
was recorded whenever possible, to ensure cov-
ering at least 1 full song cycle. Acoustic equip-
ment consisted of an Edmund Scientific 41759
hydrophone, connected via a 40dB preamplifier
with high pass filter to a Sony WMD6-C cassette
recorder. System response was 30 Hz-6 kHz.

Song from New Caledonia was compared with
song from Eden (southbound migration, 1991)
and Coffs Harbour (northbound migration,
1992), New South Wales (Table 2). Song analysis
was both aural and spectrographic. Cato (1991)
has noted that humpback whale sounds are well
suited to human aural perception. Songs are com-

posed of units, phrases and themes (Cato, 1991).
For this comparison, units were denoted by sub-
jective descriptive terms (e.g., "yap", "moan") in
the order in which they occur (Table 4). Phrases
were used as the measure of aural comparison.
Sonograms of selected segments of song were
made with a Kay Elemetrics DSP Sono-Graph
model 5500-1 (Fig. 2). Analysis filter bandwidth
was 15 Hz.

RESULTS

DETECTION OF WHALES
No humpback whales were detected at Ouvea
and Lifou in the Loyalty Islands, although only

TABLE |. Humpback whales sighted, New Caledonia

[Date [Position [Locality [Number]

| 19 Jut 1992 1607 Southeast Lagoon E
зы - OR T NEN EN
2 Aug 1992 | 2755, Г; Southeast Lagoon "mH

HUMPBACK WHALE OBSERVATIONS 507

NEW
CALEDONIA
EDEN
COFFS
HARBOUR ie- —
N
N
Pc
S
ELI ee Oe a eee аа
Tat
NEW 1
SA PENA — n M pIANBHU
voe a M a vw li РА
— p 7 17
hers 2 дии tf —
rm: e t |
Н 2 m. i
: x E44 rz
EDEN cite a cau 4. 3o NA. |o
wm MS TW ER TT |
уш Sef ОШ —®=, T
2 x Е
БА] =
COFFS = x — i
HARBOUR Zo apre
E “i т
— -=$
a Nu i

FIG. 2. Sonograms showing comparison of segments of song phrases from New Caledonia and two locations on
the east coast of Australia. Samples shown are not consecutive.

TABLE 2. Locations of humpback song heard, number
of whales singing, and duration of art

m eepe IS
mes базе | [шш
06 Jui 92. юта [cats tow | Coffs Harbour санын + fom |

а. |А Сейли:
оше:
Eie Tee as E La oon (Cap
19?44'8 Gres Northe
| 16 Aug 92 | 153°55'E [рин нойет || fiomin |
Great North

саз

lie Аш 92 © La
292 1S E den
"s Great North
19°30°S
ma 3. AN

small areas around these islands were examined
due to strong winds. Five humpbacks in 3 pods
were sighted in the east and southeast of the main
island of New Caledonia during 463km travelled
in lagoon waters (Fig. 1; Table 1). These sightings
have been included in the incidental sightings
study of Garrigue and Gill (1994). Wind strengths
exceeded 30 kmh- for more than half of the
period spent in New Caledonian waters, making
both visual and acoustic detection of whales dif-
ficult at these times.

Song was detected and recorded on four occa-
sions in New Caledonian waters, off both south-
east and northwest extremities of the main island
(Table 2). Song was heard during only one of
three sightings due to wind-induced water noise
during the other two. Humpbacks were detected
in the Great North Lagoon of New Caledonia by
sound alone; visual monitoring in good ta excel-
lent conditions detected no whales.

No humpback whales were detecied during à
22-hr visit to South Bellona Reef (21'53'S,
159728'E) at the southern end of Chesterfield
Plateau in calm clear conditions on 20-21 August.
The vessel was anchored during this period, and
visual and acoustic observations were made at

2-hously intervals. No humpback whales were
detected in the Chesterfield Reefs lagoon
(19°55°S, 158°25°E) during 22-26 August, de-
spite reasonable to excellent conditions, Though
the vessel traversed the lagoon twice, it was an-
chored most of the time. The entire lagoon was
visible, and was scanned several times per day
from the vessel's mast, while at anchor. А hydro-
phone was continuously deployed, and was mon-

MEMOIRS OF THE QUEENSLAND MUSEUM

огей for 10 min every 2 hour except during
periods of sleep.

No humpback whales were detected in open
ocean waters, or during visits to Middleton and
Marion Reefs in the Coral Sea (Fig. 1). After
entering the Great Barrier Reef at 20 15'S on 31
August, humpback song was frequently heard
between 20'04'S, 149'05'E and 22'54'S,
151*05'E, where monitoring was discontinued on
20 September. At times up to 4 whales could be
heard singing simultaneously, and several hump-
backs were seen, including 4 cow-calf pairs. À
notable exception was within the Whitsunday
Islands, wherc heavy vessel traffic was evident,
and no whales were seen or heard.

SoNG ANALYSIS

Songs from the three areas compared here
(New Caledonia Southeast Lagoon, Eden, Coffs
Harbour) each have three themes, and show a
high degree of shared phrases, as determined by
aural comparison (Table 3). Coffs Harbour
shared 89% of phrases with both Eden and New
Caledonia, while Eden and New Caledonia
shared 78% of phrases. A transcript of the aural
comparison is shown in Table 4,

Spectrographic analysis of selected phrases
(B2, C3, C4) shows similarities in structure and
pallern of humpback sound types from the three
areas (Fig. 2).

DISCUSSION

New Caledonia. Useful information was ob-
tained from acoustic monitoring during this
study. Song was heard on four occasions, and
humpback whales sighted on three. Acoustic
monitoring extended the known distribution of
humpbacks into the Great North Lagoon, a large

TABLE 3. Shared and unique song phrases.

Harbour Caledonia

|
|
|
Jae] о [o fas рое хх |а

>

|
| с
|

4A

HUMPBACK WHALE OBSERVATIONS

509

TABLE 4. Aural comparison of humpback song, using subjective descriptive terms. Symbols: (R) repeated; (P)
pause; Tupward; (2) unit repeated. Unit in bracket repeated,

Bloop-low moan-gloop(2)-low moan(2)-cries-cows

ayne cries-shorter cnes into chirps

gloop- woo cry VP) wo cry-woo cry (P)-lowercries into whistles] - (R)

тпоап-моо cry

) woo cry (P) woo cry-lower cries into whistles] - (R)

whisltle chirps

Tmoan( 2)-long low moan - T moan-long low moan
very short moan-undulating medium groan] - (R)

very short moan-ratchet moans] - (R)

| Song characteristics:

| dominent (most frequent) theme 15 B
* phrase B2 repeated most often
» medium-length son;

plop low moan-gloop (2) - low сшы

ayne cnes-shorter cries into chirps] (

gloop (2) - eries-cows] - (Е)

loop-woo cry (Р) woo cry (P) yog сту lower cries into whistles] - (R)

moan-woo cry p woo cry

moan-long moan lon

- | moan-woo cry-lower cries into whistles] - (R)

moan - raspy whistle ch

T moan (2)- long low moan - T moan-long low moan
very short moan-undulating medium moan] - (R)

(52у short тоал-гаїсһеї moans] + (К)
upward whistle-ya

| Song characteristics
| * short song; dominant theme probably A
* themes quite short

* B2 has extra cry since Eden

loop-low moan-gloop (2)-low moan-gloop (2)-cries-cows] - (К)

ayne cries-shorter cries into chirps] -

T moan-woo cry- lower cries into whistles] - (R)

S Puer) moan rasp

whistle chirps

moan-long low moan- T moan-long low moan

very short
very short moan-ratchet moans] - R
pward whistle - yaps

Song characteristics
| © short song; dominant theme probably A

moan-undulating medium moan] - (R)

* some units no longer repeated e.g. moan of Cl, long moan of B2

| * Bl of pre

and rarely-visited area where no sightings had
been reported. It also permitted the first song
comparison that we know of between songs from
two Group V breeding arcas.

Although song was recorded after the vessel
entered north Queensland waters, it is not used
here for comparison with New Caledonian song.
The songs from Eden and Coffs Harbour were
most similar in pattern to New Caledonian song,
while the north Queensland song had evolved
substantially since July 1992. Change of song
with time may be complex even along a single
migration path (Cato, 1991), let alone between
breeding areas as well as seasons. Here we at-
tempt only to reporí similarities between songs
from New Caledonia and east Australia, and these

similarities are most apparent in the examples we
have used.

Humpback song is considered to be an indicator
of stock identity (Payne & Guinee, 1983); songs
tend to show decreased similarity with increased
geographical separation within ocean basins, arid
little or no similarity between ocean basins (Winn
et al. 1981; Helweg et al., 1990; Dawbin & Eyre,
1991). While some differences were found be-
tween the songs compared here (such as the num-
ber of repetitions, or the omissions of units or
phrases), such variation can occur between indi-
viduals in the same area, or even between differ-
ent song renditions by an individual (Dawbin &
Eyre, 1991). But the song from the three arcas
was basically similar. The Coffs Harbour song

510

shared as many phrases with New Caledonia as
with Eden (Table 3), while the closest resem-
blances in spectrographic structure appear to be
between New Caledonia and Eden (Fig. 2).

Given the relatively close proximity of New
Caledonia and east Australia, and that Group V
breeding grounds extend at least to Fiji (Dawbin,
1966) and probably Tonga, the similarities in
song reported here are not surprising. An antici-
pated migratory connection between New Cale-
donia and other Group V breeding grounds was
established in 1993 by a photo-identification
match with Hervey Bay, Queensland (Garrigue
& Gill, 1994). The similarities in song reported
here add further evidence for migratory exchange
between humpback whale sub-populations of
New Caledonia and east Australia.

Chesterfield Reefs. There аге numerous reefs on
the Chesterfield Plateau, Townsend's (1935)
chart shows a wide spread of points representing
humpbacks killed by American whalers between
July and October (approximately 45% of them in
August), centred around Chesterfield Reefs, to-
ward the northern end of the Plateau. French
pelagic whalers operated there at least in 1862
and 1864 (du Pasquier, 1982). The Chesterfield
Reefs lagoon ts the most sheltered body of water
on the Plateau, enclosing roughly 85 sq km. It
meets the criteria cited by Whitehead & Moore
(1982) as humpback breeding habitat, offering
shelter, having suitable depths (average 50m) and
warm water (23°С in August 1992). It would have
provided whaling vessels with safe anchorage
and easy access to whales in or near thé lagoon.
Numerous whale bones, some of them with marks
from chopping implements still visible, have
been located by divers in the lagoon (B. Crouch,
pers. comm.).

Only two reliable reports have been obtained of
humpbacks at Chesterfield Reefs during this cen-
tury, In June-July 1972, 5-10 humpbacks were
seen, and song heard by divers, inside the Ches-
terfield Reefs lagoon (R. & V. Taylor, pers.
comm.), More recently, a humpback mother and
calf were observed in the same lagoon over sev-
eral weeks during September-October 1990 (B.
Crouch, pers. comm.) This observer has not seen
humpbacks during other frequent visits to Ches-
terheld Reefs in recent winters. Several French
Government (ORSTOM) scientific expeditions
during the past decade, including bathymetric
surveys of the Plateau itself, have reported no
sightings in winter and spring months (B. Richer
de Forges, pers, comm.).

MEMOIRS OF THE QUEENSLAND MUSEUM

Due to bad weather the vessel did not traverse
the main body of Chesterfield Plateau during this
cruise, visiting only South Bellona and Chester-
field Reefs, for 22 hours and 5 days, respectively.
During these periods no whales were observed or
heard in good to ideal monitoring conditions.
‘These observations are inconclusive, however, as
whales may have been present elsewhere on the
Plateau, or at Chesterfield Reefs earlier or later
than the yacht's visit. The determination of the
current status of Chesterfield Plateau's reefs as a
hum k breeding area therefore awaits à more
comprehensive study.

ACKNOWLEDGEMENTS

Field assistance from Tim Edkins, Diana Da-
vidson, Peter Drew, Chook Wall-Smith, and Deb-
bre Milham made the study possible. Ken Schultz
and Dr Doug Cato assisted with sonograms. Dr
Peter Corkeron offered useful comments on the
text. Ms B. Jantulik prepared Figure 1. The study
was partly funded by a grant from the М.У. Scott
Estate, and was conducted under Whale Protec-
tion Act Permit No, P1992/027, issued by the
Australian National Parks and Wildlife Service,
Canberra,

LITERATURE CITED

CATO, D.H. 1991. Songs of humpback whales; the
Australian perspective. Memoirs of the Queens-
land Museum 30(2): 277-290.

CLAPHAM, P.). & MATTILA, О.К. 1990. Humpback
whale songs as indicators of migration routes,
Marine Mammal Science 6(2): 155-160.

DAWBIN, W.H. 1964. Movements of humpback
whales marked in the south west Pacific Ocean
x to 1962, Norsk Hvalfangst-Tidende, 3: 68-

1966. The seasonal migratory cycle of humphack
whales. 145-170. In K.S. Norris (ed,), ‘Whales,
Dolphins and Porpoises', (University of Califor-
mia Press: Berkeley).

DAWBIN, W.H. & FALLA, R.A. 1949, A contribution
to the study of the humpback whale based on
observations at New Zealand shore stations. Pro-
cedin E of the 7th Pacific Science Congress. 4:
373-382.

DAWBIN, W.H. & EYRE, EJ. 199]. Humpback whale
songs along the coast of Westem Australia and
some comparison with east coast songs. Memoirs
of the Queensland Museum 30(2): 249-254.

DAWBIN, W.H. & GILL, P.C. 1991. Humpback whale
survey along the west coast of Australia: à com-
parison of visual and acoustic observations, Mem-
oirs of the Queensland Musuem 302): 255-257,

HUMPBACK WHALE OBSERVATIONS

DU PASQUIER, T. 1982. 'Les baleinieres Francais au
XIXieme siecle, 1840-1868'. (Terre et Mer 4 Sei-
gneurs, Grenoble).

GARRIGUE, C. & GILL, P.C. (1994). Observations of
humpback whales (Megapterea novaeangliae) in
New Caledonian waters during 1991-1993. Bio-
logical Conservation 70:211-218.

HELWEG, D.A., HERMAN, L.M., YAMAMOTO, S.
& FORESTELL, P.H. 1990. Comparison of songs
of humpback whales (Megaptera novaeangliae)
recorded in Japan, Hawaii and Mexico during the
winter of 1989. Scientific Reports of Cetacean
Research 1: 1-20.

LANGDON, R. 1984. “Where the whalers went’. An
index to the Pacific ports and islands visited by
American whalers (and some other ships) in the
19th century. (Pacific Manuscripts Bureau, Re-
search School of Pacific Studies, Australian Na-
tional University: Canberra),

PATERSON, R.A. 1991. The migration of humpback
whales Megaptera novaeangliae in east Austra-
lian waters. Memoirs of Queensland Museum
30(2): 333- 341,

511

PAYNE, RS. & GUINEE, L.N. 1983. Humpback whale
(Megaptera novaeangliae) songs as an indicator
of "stocks". 333-358. In R. Payne (ed.), ‘Commu-
nication and Behaviour of Whales’. (AAAS Se-
lected Symposia Series, Westview Press:
Boulder).

SIMMONS, M.L. & MARSH, H. 1986. Sightings of
humpback whales in Great Barrier Reefwaters.
Scientific Reports of the Whales Research Insti-
tute, Tokyo 37: 31-46.

TOWNSEND, C.H. 1935. The distribution of certain
whales as shown by log book records of American
whaleships. Zoologica 19: 1 -50.

WHITEHEAD, H. & MOORE, M.J. 1982. Distribution
and movements of West Indian humpback whales
in winter. Canadian Journal of Zoology 60: 2203-
2211.

WINN, H.E., THOMPSON, T.J., CUMMINGS, W.C.,
HAIN,J., HUDNALL, J., HAYS, H., & STEI-
NER, W.W. 1981. Song of the humpback whale -
population comparisons. Behavioural Ecology
and Sociobiology 8: 41-6.

512

PREDATION ON THE CANE TOAD (BUFO
MARINUS) BY THE BLACK KITE (MILVUS
MIGRANS). Memoirs of the Queensland Museum 38(2):512.
1995:- Since the introduction of the Cane Toad into Australia
in 1935 there have been several reports of native fauna that
have been adversely affected by preying upon them (Cov-
acevich & Archer, 1975; Covacevich & Couper 1992) how-
ever several native animals [Water Rats (Hydromys
chrysogaster), Crows (Corvus spp.), Koels (Eudynamys
scolopacea) and Keelback Snakes (Tropidonophis mairii)]
are known to prey successfully on them with no obvious toxic
effect (Covacevich & Archer, 1975; Covacevich & Ingram,
1990). Herein we report predation upon Cane Toads by nest-
ing Black Kites in July 1995.

During surveys of the Ross River Dam area AJ observed
several Black Kite nests with toad remains on the ground
below them. Detailed observations were made on a single
Black Kite nest in a paperbark tree (Melaleuca nervosa)
beside a farm dam approximately 24km South West of
Townsville, north-east Queensland (Map No. 8259 Grid Ref.
715 480). While there were no young in the nest, six Black
Kites were perched around the nest and in adjacent trees.

Over twenty toad remains were observed lying on the
ground below the nest. Furthermore, there were toad remains
hanging from branches beside the nest suggesting that the
Black Kites had fed portions of Cane Toad to the young in the
nest. Close inspection of the toads revealed that they had had
portions of their internal organs removed. The toads appeared
to have been opened from the ventral surface presumably
allowing the Kites to avoid the toxic glands on the dorsal
surface, This observation corroborates with Lavery (1969)
who reported toad remains in gut-analyses of Black Kites in
Townsville (along with Hylid and Lymnodynastine frogs).

One of us (AJ) has observed Black Kites hovering around
the edge of dams presumably looking for prey and as Cane
Toads are known to hide in hoof prints near the water during

MEMOIRS OF THE QUEENSLAND MUSEUM

the day (JMH, pers obs.) we suspect that they were actively
searching for them.

Our observations suggest that Black Kites have adapted
their behaviour to exploit this intoduced amphibian despite its
toxicity. Firstly it has learned to find and capture Cane Toads
in their diurnal refuges. Secondly it has learned to eat the
internal organs of the Cane Toad avoiding the toxins concen-
trated on the dorsal surface. Black Kites are an abundant
predator in the Townsville region that could influence Cane
Toad populations. We hypothesize that this predator, along
with other native animals which are adapting their behaviour
to eat Cane Toads, will reduce toad populations and hence the
impact of this introduced amphibian on the Australian envi-
ronment.

Literature Cited

Covacevich, J. & M. Archer. 1975 The distribution of the
Cane Toad Bufo marinus, in Australia and its effects on
indigenous vertebrates. Memoirs of the Queensland
Museum 17(2):305-310.

Covacevich, J. & С. J. Ingram. 1990 Tropidonophus mairi Vs
Bufo marinus. Memoirs of the Queensalnd Museum
29(2):396

Covacevich, J. & P. J. Couper. 1992 The Carpet Python
Morelia spilota (Lacepede), another unsuccessful pred-
ator of the Cane Toad, Bufo marinus (Linnaeus), in
Australia.

Lavery, H. J. 1969 Collisions between aircraft and birds at
Townsville, Queensland. Queensland Journal of Agri-
cultural and Animal Sciences 26:447-455.

David Mitchell, Arthur Jones & Jean-Marc Hero, Department
of Zoology, James Cook University, Townsville Queensland
4811, Australia; 19 October 1995.

A NEW LOCALITY FOR THE HASTINGS RIVER MOUSE, PSEUDOMYS ORALIS, IN

SOUTHEAST QUEENSLAND
IAN C. GYNTHER AND PETER S, O'REILLY

Gynther, LC, & O'Reilly, P.S. 1995 12 01: A new locality for the Hastings River Mouse,
Pseudomys oralis, in southeast Queensland. Memoirs of the Queensland Museum 38(2):5] 3-
518, ISSN 0079-8835.

A Hastings River Mouse, Pseudmnys oralis, was captured near O'Reilly's Rainforest
Guesthouse on the western edge of Lamington National Park in December 1994. This is only
the second locality at which the species has been trapped in Queensland in 25 ycars. The
capture site, in the McPherson Range, is almost 70km east of the P. øralis population recently
discovered in Gambubal State Forest. The steep topography and comparatively dry vegeta-
tion at this new location differ considerably from those reported for previous Hastings River
Mouse capture sites. These findings suggest that a wider variety of potential localities and
environments should be targeted during future surveys of this species. [ ] Pseudomys oralis,
new locality.

Ian Gynther, NatureSearch, Queensland Department of Environment and Heritage, PO Box
155, Brisbane, Albert Street, Queensland 4002, Australia; Peter $. O'Reilly, O'Reilly's
Rainforest Guesthouse, Lamington National Park Road, via Canungra, Queensland 4275,

Australia; 15 September 1995,

The Hastings River Mouse, Pseudomys oralis
(Rodentia: Muridae), is patchily distributed in
mideastern Australia at altitudes between 400m
and 1250m (Read, 1993a,b, pers. comm .;
Tweedie & York, 1993), Until this study, thirty
capture localities were known for the species -
five in southeast Queensland and the remainder
in northeast New South Walcs (Hastings River
Mouse Recovery Team, 1993; Fox et al., 1994).
The first records of the Hastings River Mouse in
Queensland were from the perind 1969-1970,
when animals were trapped at four separate local-
ities southeast of Warwick (Kirkpatrick & Mar-
tin, 1971). Despite additional surveys (Read.
1988), the species was not seen again in this State
for over 20 years. In 1993, it was ‘rediscovered’
near the original capture locations, but at higher
altitude, in Gambubal State Forest (Poole, 1994).

Fossil evidence indicates that the present rarity
of P. огай is only the result of a relatively recent
reduction in the species” distribution and abun-
dance, Skeletal remains of the Hastings River
Mouse were common from Holocene cave depos-
its associated with owl roosts along the Great
Dividing Range in New South Wales and Victo-
ria (Wakefield, 1972; Hall, 1974; Kirkpatrick,
1983), Furthermore, the co-occurrence of P. or-
alis and Rattus rattus remains at certain localities
(Hall, 1974; Lee, 1995) suggests that even at the
time of European settlement the distribution of
the species was much grcater than at present.

In light of this rapid contraction in range of the
Hastings River Mouse and the paucity of infor-

mation on the species’ distribution and ecology
(Read, 1993b), conservation efforts have focused
on locating remaining populations and identify-
ing environmental features that may be used as
indicators to predict additional localittes for this
rare rodent (King, 1984; King & Mackowski,
1986; Read, 1988, 1993a,b; Hastings River
Mouse Recovery Team, 1993; Tweedie & York,
1993). In addition, dietary studies have recently
been undertaken to assist in determining more
precise habitat requirements of the Hastings
River Mouse (Fox et al., 1994).

The present paper, documenting a newly dis-
covered locality for P. oralis in Queensland,
broadens existing knowledge of both the species’
geographical distribution and the range of habi-
tats in which it occurs, Importantly, this paper
suggests that in Queensland, at least, the current
‘search jmage’ used by field workers to select
potentially suitable Hastings River Mouse habi-
lats may be loo narrow,

METHODS

Dunng à general mammal survey conducted in
the vicinity of O' Reilly's Rainforest Guesthouse
between December 1 1 and 16, 1994, a precipitous
ridge known as Castle Crag near the western edge
of Lamington National Park was chosen as one of
many locations to sample for small mammals.
Trapping cffort at this particular site was low,
with 24 size A Elliott traps, baited with salami
and a mixture of peanut butter and rolled oats,

514

Beechmont

ALBERT

.
Hillview

LAMINGTON
NATIONAL PARK]

Tyalgum »

FIG. 1. Location of Castle Crag.

being set along the ridge crest over the two nights
of December 15 and 16.

SITE DESCRIPTION

The trapping site (28°14 13"S, 153'07'06"E),
located u km WSW of O'Reilly's Rainforest
Guesthouse, lies at an altitude of 790m on the
west-aligned Castle Crag ridge (Fig. 1). To the
north, Morans Creek is some 250m below and to
the south, the Left Branch of the Albert River 1s
400m lower in elevation. A wall of exposed basalt
lies along the ridge crest (Fig. 2) and small rock
outcrops and cliff faces are features of the south-
ern slope

Mean annual rainfall at O’Reilly’s is 1643mm,
however, the trapping period coincided with the
end of a severe drought, With precipitation total-
ling only 1013mm, 1994 was the fourth driest
year since 1917 when records at the Guesthouse
began.

The vegetation at the trapping site consists of
an open dry sclerophyll forest, with a canopy
height of 8-10m, growing on a shallow, brown

"NERANG

MEMOIRS OF THE QUEENSLAND MUSEUM

s Neranwood

"Nurminbah
Valley

Murwillumbah
Ф

10 15 кт

kraznozem (Fig. 3). The forest is dominated by
Eucalyptus banksii and Allocasuarina torulosa,
with an occasional Lophostemon confertus and a
sparse shrub layer of Xanthorrhoea glauca. The
steepness of the terrain (variable, but averaging
c. 70°) and several fires within the last 14 years
ensure a minimum of accumulated deadfall. The
ground cover is primarily dense Danthonia
longifolia growing to a maximum height of about
40cm, although close to the ridge crest the cover
provided by this grass is more sparse. Other spe-
cies in the immediate vicinity of the trapping site
were Dendrobium kingianum, forming a large
clump approximately 25cm high, Daucus
glochidiatus, Plectranthus graveolens,
Wahlenbergia graniticola, Eupatorium ad-
enophorum, E. riparium, Brachyscome as-
cendens, Helichrysum bracteatum, Rhodanthe
anthemoides, Podolepis neglecta, Lepidosperma
laterale, Dianella caerulea var. assera, Bulbine
vagans, Cymbopogon refractus and Doodia as-
pera. Lomandra longifolia was poòrly repre-
sented in the understorey near the ridge top, but
grew in scattered clumps some 30m downhill of
the trap line. The growth of mosses and small
ferns lower on the slope indicated that soil mois-

HASTINGS RIVER MOUSE

HG. 2. Р. oralis capture site (arrowed) near basalt wall
along the Castle Crag ridge crest.

ture is greater there than near the ridge crest. The
vegetation of the much warmer northern side of
the ridge was relatively sparse and included Eu-
calyptus tereticornis, E. melliodora and occa-
sional Acacia melanoxylon.

Approximately 450m to the east, the ridge line
broadens and the dry sclerophyll forest grades
into very tall (35m) wet sclerophyll forest with a
dense fern understorey. This gives way to
notophyll уте forest after a further 130m. The
nearest permanent water to the trapping site is the
section of Morans Creek abave Morans Falls,
approximately 900m to the northeast, A small
seepage area 800m to the northeast would only
provide moisture after periods of rain,

RESULTS AND DISCUSSION

An adult Hastings River Mouse (Fig. 4) was
captured on Castle Crag on the night of December
15, 1994. The successful trap was positioned on
a small grassy ledge at the top of the southerly
aspect, close to the exposed basalt wall (Fig. 2).
No other mammals. were caught in the trap line,

515

although a total of five Major Skinks, Egernia
frerei, was also captured over the two nights. The
P, oralis individual, a lactating female, was re-
moved from the site and weighed, measured and
photographed, before being released shortly af-
terwards at the point of capture. Its head and body
(to vent) length was 135mm, tail to vent length
141mm, hind foot length 30mm and weight 85g.
The identification was subsequently confirmed
from the photographs by S. Van Dyck of the
Queensland Museum.

Castle Crag is only the second locality at which
P. oralis has been trapped in Queensland since
the initial records of Kirkpatrick & Martin (1971)
near Warwick in 1969-70. Lying approximately
68km cast of the site where a population was
recently discovered in Gambubal State Forest
(Poole, 1994), 3t is well away from the general
vicinity of previous captures, Indeed, the present
locality on the northern fall of the McPherson
Range (Fig. 5) is unique because all other speci-
mens of the Hastings River Mouse in eastem
Australia have been caught in close proximity to
the Great Divide (Read, 1993a; Fox et al., 1994).
Earlier, ‘outlying’ records of P. oralis do exist -
identifiable bones were collected in 1976 from
‘relatively fresh’ owl pellets near Mapleton in the
Blackall Range of southeast Queensland (Kirk-
patrick, 1983; Read, 1988) and also in 1986 from
a Grass Owl, Tyto capensis, pellet near Wardell
in coastal New South Wales (G. Holmes, pers.
comm.; Lee, 1995) - but the species has not vet
been recorded alive at these localities,

The present record is of particular interest be-
cause of the nature of the topography and vege-
tation on Castle Crag. The altitude and rainfall at
the site lie within the ranges recorded for previous
Hastings River Mouse capture localities (King,
1984; King & Mackowski, 1986) and the general
habitat is an open dry sclerophyll forest with a
grassy understorey, as 15 typical for the species
(e.g, King & Mackowski, 1986; Read, 1993b;
Tweedie & York, 1993; Fox et al., 1994), An
obvious difference, though, ts that the forest can-
ору is much lower (8-10m) than the 20-40m
upperstorey heights reported for other P. oralis
sites (King, 1984; King & Mackowski, 1986;
Townley, in press), due to the location on a nar-
row, exposed ridge on shallow soils. However,
Read (1993b) concludes that the type or abun-
dance of tree cover are probably not important
factors in determining suitable habitat for the
Hastings River Mouse, and so details of the forest
structure on Castle Crag may not be especially
critical. A more significant difference is that most

516

FIG. 3. Southern slope of Castle Crag fram P. oralis
capture site,

other capture sites possess stands of sedges
(Cyperaceae and Juncaceae, particularly Carex,
Cyperus and Juncus) growing in association with
either permanent water in creeks and gullies or
with bogs, soaks or seepage areas on ridges and
mid-slopes (Read, 1988, 1993a,b; Hastings River
Mouse Recovery Team, 1993; Tweedie & York,
1993: Lee, 1995), Such moist conditions do not
exist on Castle Crag. Permanent water is not
accessible and the lack of terracing on the steep
slope has prevented the formation of boggy areas
which would enable the growth of stands of these
sedges (cf. Read, 1993a,b). Also, there are no
seepage areas nearby and ground moisture is
apparently insufficient to support species of
Carex, Cyperus or Juncus, The only sedge pres-
ent, Lepidosperma laterale, is represented by in-
dividual plants sparsely distributed through the
understorey,

Of the ground cover species recorded at Castle
Crag, only Lomandra longifolia, Lepidosperma
laterale and Doodia aspera have been reported
in the understorey at other captures sites (King,

MEMOIRS OF THE QUEENSLAND MUSEUM

1984; King & Mackowski, 1986; Hastings River
Mouse Recovery Team, 1993; Read, 1993b; Fox
et al., 1994), suggesting a difference in habitat
type at this new locality. The sedge L. laterale has
never been recorded in the diet of P. oralis, but
seed head material from Lomandra longifolia and
pollen from an unidentified species of mat-nish
arc known to be eaten by the Hastings River
Mouse (Fox et aL, 1994; A. Smith & D. Quin,
unpublished data), although Read (1993b) does
not consider L. longifolia to be an indicator of
suitable P. oralis habitat where it occurs on hill
sides, On separate occasions, S. Townley (pers.
comm.) has observed radio-tracked animals in
Billilimbra State. Forest in New South Wales
eating Doodia aspera and an undetermined spe-
cies of Plectranthus, Leaf and seed head material
lentatively identified as originating from a
Plectranthus sp. was also found during a dietary
analysis of the Hastings River Mouse in the same
State Forest (A. Smith & D. Quin, unpublished
data). Plectranthus graveolens is present at Cas-
tle Crag and may, together with Doodia, repre-
sent a food resource for the Hastings River Mouse
there, Clearly, an analysis of faecal pellets from
animals at the Lamington National Park site
would provide valuable information about what
plant species and items are eaten and would assist
in refining current knowledge of habitat features
that are critical for P. oralis at this and other
locations.

The Hastings River Mouse capture site in
Gambubal State Forest shares some similarities
with the present one in that it is also distant from
surface water and is adjacent to a rocky escarp-
ment (Hastings River Mouse Recovery Team,
1993; Poole, 1994). S. Townley (pers. comm.)
bas found that individual animals in Gambubal
State Forest use cracks and crevices in rocks of

FIG. 4, Adult female P. oralis captured al Castle Crag.

HASTINGS RIVER MOUSE

SOUTH WALES

~

i Armidale &

A

FIG. 5. The location of Castle Crag (asterisk) in relation
to previous P. oralis capture sites in eastern Australia
(dots). The area shaded grey is land below 500m
elevation. Four sites in New South Wales discovered
since 1993 (Fox etal., 1994) аге not shown. (Adapted
from Read, 19933).

the cliff face for shelter. The same may be true on
Castle Crag, where potentially suitable nesting
locations in areas of exposed rock were numer-
ous. There are, however, fundamental differences
between these two Queensland sites. Substan-
tially more habitat area is available for the species
at Gambubal State Forest, most occurring on flat
or gently sloping terrain, and conditions there are
more lush, with the grassy, tall open forest abut-
ting notophyll vine forest across an abrupt eco-
tone. The open forest of this ecotonal area has
recently been found to support significant quan-
tities of the sedge Carex breviculmis (Y. Ross,
pers. comm.). By contrast, as previously stated,
no Carex spp. occur at the capture site in
Lamington National Park and the nearest
notophyll vine forest is almost 600m away.
Given that the female Hastings River Mouse
reported here was in a reproductive state and,

517

therefore, unlikely to be a transient individual аї
the site, this record indicates that Castle Crag
supports a viable, even if isolated, population of
P. oralis. Furthermore, it extends the diversity of
habitat types the species is known to utilise and
suggests a broader range of localities and envi-

| ronments should be targeted during future sur-

veys for the Hastings River Mouse, particularly
in Queensland. Additional work to determine the
population size and distribution of P. oralis in the
Lamington National Park area is currently being
undertaken to clarify the status of the species in
this State.

ACKNOWLEDGEMENTS

The O'Reilly's Rainforest Guesthouse initiated
and facilitated the trapping program, which was
undertaken by IG as part of the NatureSearch
project. We are grateful to Steve Van Dyck,

; Queensland Museum, for the loan of additional
| traps and for much helpful discussion. We would

also like to thank Greg Gordon and Noleen Kunst,
Queensland Department of Environment and
Heritage, and Sally Townley, Southern Cross

_ University, for their comments on an earlier draft

of this paper. Staff of the Queensland Herbarium
provided identifications of the herbaceous plant
specimens and David Read kindly supplied the
original artwork to enable us to produce Fig. 5.

LITERATURE CITED

FOX, B. J., READ, D. G., JEFFERYS, E. & LUO, J.
1994. Diet of the Hastings River Mouse
(Pseudomys oralis). Wildlife Research 21: 491-
505.

HALL, L. S. 1974. A recent bone deposit at Marble
Arch, N.S.W. Pp. 35-46. In ‘Proceedings of the
Tenth Biennial Conference’, (Australian Speleo-
logical Federation; Sydney).

HASTINGS RIVER MOUSE RECOVERY TEAM
1993. Interim Hastings River Mouse habitat iden-
tification guide. NSW National Parks and Wildlife
Service.

KING, С.С. 1984, Habitat utilized by Pseudomys oralis
Thomas (Rodeniia: Muridae). Australian Mam-
malogy 7: 139-147.

KING, б. C. & MACKOWSKI, C. М. 1986. Two new
localities for Pseudomys oralis Thomas
(Rodentia: Muridae) in New South Wales. Aus-
tralian Mammalogy 9: 63-65.

KIRKPATRICK, T. H. 1983. Hastings River Mouse
Pseudomys oralis. In Strahan, R. (ed.), "The Com-
plete Book of Australian Mammals". (Angus and
Robertson: Sydney).

518

KIRKPATRICK, T. H. & MARTIN, J. H. D. 1971.
Uncommon native fauna. Queensland Agricul-
tural Journal 97: 114-115.

LEE, A. К. 1995. ‘The Action Plan for Australian
Rodents'. (Australian Nature Conservation
Agency: Canberra).

POOLE, M. A. 1994. The Hastings River Mouse,
Pseudomys oralis, from Gambubal State Forest,
southeast Queensland. Memoirs of the Queens-
land Museum 37(1): 280.

READ, D. G. 1988. Surveys for the rare Hastings River
Mouse in south-east Queensland. A report to the
Queensland National Parks and Wildlife Service.

1993a. Body size in Hastings River Mouse
Pseudomys oralis (Rodentia: Muridae) from new
and old locations. Australian Zoologist 29(1-2):
117-123.

MEMOIRS OF THE QUEENSLAND MUSEUM

1993b. Prescriptions for the identification of habi-
tats of the Hastings River Mouse Pseudomys
oralis (Rodentia: Muridae). Forestry Commis-
sion of New South Wales. Technical Report No.
58.

TOWNLEY, S. In press. Home range, movement and
microhabitat distribution of Hastings River
Mouse Pseudomys oralis (Rodentia: Muridae).
Wildlife Research.

TWEEDIE, T. D. & YORK, A. 1993. Survey guidelines
for the Hastings River Mouse (Pseudomys oralis).
Forestry Commission of New South Wales. Tech-
nical Paper No. 62.

WAKEFIELD, N. A. 1972. Palaeoecology of fossil
mamrmal assemblages from some Australian
caves. Proceedings of the Royal Society of Victo-
ria 85: 1-26.

THE BUTTERFLY TYPES OF W, H. MISKIN IN THE QUEENSLAND MUSEUM
(LEPIDOPTERA)

DAVID L. HANCOCK

Hancock, D.L. 1995 12 01: The butterfly types of W.H. Miskin in the Queensland Museum
(Lepidoptera), Memoirs of the Queensland Museum 38(2); 519-528, Brisbane, ISSN 0079-
8835,

Notes are provided on the type specimens of the 65 species of Australian butterflies described
between 1874 and 1891 by №. Н. Miskin. [ ] туре specimens, Lepidoptera.

David L, Hancock, Department of Primary Industries, Meiers Rd, Indooroopilly, Queensland

4068, Australia; 15 September 1995.

William Henry Miskin published 19 papers on
Australian butterflies between 1871 and 1891.
Descriptions of 65 new butterfly taxa were pub-
lished in 11 papers (Miskin 1874c, 1876, 1884b,
1889a, 1889c, 1889d, 1889e, 1890a, 1890b,
1890c, 1891), whilst a further 8 papers (Miskin,
1871, 1874a, 1874b, 1883, 1884a, 1888, 1889b,
1890d) appeared in which no new taxa were
described.

Miskin did not place type or determination
labels on his specimens and in most cases the
number of specimens used for each description is
unknown. Were it not for his meticulous hand-
written notebooks, now preserved in the Queens-
land Museum, in many cases it would be
impossible to determine which specimens ate in
fact types. Fortunately, Miskin applied 4 species
number to each specimen in his collection and
this corresponds with those listed foreach species
in his notebooks. In many cases locality labels are
also absent bui for those specimens from his own
collection the locality appears to be Brisbane. In
the list below, numbers given in brackets after the
species’ citations are those allocated to the spe-
cies in Miskin's notebooks. Species without a
number were not present in his collection.

In July 1910, G, A. Waterhouse (Australian
Museum) and C. J. Wild (Queensland Museum)
added type labels to many of Miskin's syntypes,
but in 2 cases they also added labels to specimens
that are not types. A copy of G. A. Waterhouse's
notés on the collection made during his 1910
examination is kept at the Australian National
Insect Collection, Canberra. Reference to these
notes has been made where appropriate. The

number of specimens listed for the various spe-

cies in Miskin's Coliection by Waterhouse in
1910 and the number currently located. some-
times differ but in at least some cases
Waterhouse's lists also include specimens added

to Miskin's Collection subsequent to his descrip-
Lions.

Of the 65 nominal species described by Miskin,
holotype, lectotype or syntype material has been
located in the Queensland Museum for 53 spe-
cies. Seven species were described from the
Lucas Collection, 1 from the Australian Museum
Collection. Two of the Lucas species are repre-
sented by tvpes in the South Australian Museum.
Types of 4 species supposedly in the Queensland
Museum have not been located. Lectotypes have
been designated directly by Sands (1986) and by
De Baar & Hancock (1993) and "by inference of
Holotype", under the rules of the International
Commission on Zoological Nomenclature, by
Eliot & Kawazoe (1983), Sands (1980) and Wa-
terhouse (1933, 1937).

Thirty-five taxa remain valid at the species or
subspecies level, the remainder are synonyms.
Determination labels and Type numbers
{QMT 12342 to QMT12440) have been added to
each specimen by the present author.

TYPE SPECIMENS

Family PAPILIONIDAE

Papilio egipius Miskin, 1876:451 (190)

Described fram 3 males and 1 female collected by М
Kuntsler [in late 1875 or early 1876] at Rockingham
Вау (= Cardwell), Qld. Only the 2 males from Miskin's
Collection have been located; the pair originally depas-
ited in the Queensland Museum (Miskin 1876) was not
located by Waterhouse in 1910 and still has not been
found.

SYNTYPES: 2 d (QMT12342, QM T12343): Н.К. Car-
dwell/190 3,

CURRENT STATUS: Princeps ambrax egipius (Miskin).

520

Family PIERIDAE
Delias nigidins Miskin, 1884b:93 (214)

Described from an unknown number of females col-
lected by Miskin at Rockingham Bay and Johnstone
River, Qld. Although the original description stated
"d". it was actually a female being described; the
discussion noted that only females were known. The
male was described subsequently (Miskin 1889a:
1515). Two females and a male have been located
ЅҮМТҮРЕЅ: 29: (ОМТ12344, QMT12345). -/5/82
Cardwell/214, One 9 subsequently labelled: Misk,
Type 2; 28/7/10, GAW, CJW (on reverse).
METALLOTYPE (not an original type): 1d. (QM):
Johnstone R/214. Subsequently labelled; Misk. Type
d;28/7/10, GAW, CIW (an reverse)/ 1/3011.
CURRENT STATUS: Delias ennia nigidius Miskin.

Elodina perdita Miskin, 1889e:263 (44)

Described from an unknown number of specimens
collected by Augustus Simson at Port Denison (=
Bowen), Qld. Two of the 3 specimens noted by Water-
house are regarded as types. These were designated
Lectotype and paralectotype by De Baar and Hancock
(1993). The paralectotype female has the hindwing un-
derside yellow. The third specimen, a female with the
hindwing underside white, 1s labelled "Bowen/184". In
Miskin’s notebook, species number 184 is listed as
Elodina egnatia and this specimen cannot be regarded
as part of Miskin's original type series.

LECTOTYPE: d (QMT12346), A.S., В, Bowen/ 44.
Subsequently labelled: Lectotype 4, Elodina perdita
Miskin, desig, M. de Baar & D. L. Hancock 1992 [red
label].

PARALECTOTYPE: ? (QMT12347), A.S., C.
Bowen/44, Subsequently labelled: Paralectotype р
Elodina perdita Miskin, desig. M. de Baar & D. L
Hancock, 1992.

CURRENT STATUS: Elodina perdita Miskin.

Tachyris asteria Miskin, 1889a:1514

Described from a male from Port Douglas, Old in the
Lucas Collection. Miskin's notebook indicates only 1
specimen; it has not been locatext

CURRENT STATUS: Synonym of Appias melania
(Fabricius, 1775).

Terias immaculata Miskin, 1889d:258 (84)

Described from an unknown number of specimens
collected at Rockhampton, Qld. One of the 4 specimens
noted by Waterhouse has been located,

SYNTYPE: 1d (QMT12348), -/1/75 Rockhampton/84.
The following labels were added subsequently: Misk.
Type @ 7; 27/7/10, САМ, CJW (on reverse)/ L/3013/

? Terias immaculata.

MEMOIRS OF THE QUEENSLAND MUSEUM

CURRENT STATUS: Synonym of Eurema herla (W.S.
Macleay, 1826).

Terias lineata Miskin, | 889d:257

Described from 2 specimens from North Queensland
in the Lucas Collection. Not located.

CURRENT STATUS; Synonym of Eurema sana (Butler,
1877).

Terias varius Miskin, 1889d:259 (14)

Described from an unknown number of specimens
collected at Brisbane and Rockhampton, Three of the
5 specimens noted by Waterhouse have been located,
Specimens from Dawson River (Expedition Range,
inland from Rockhampton) appear to have been in
Miskin's Collection since at least 1875 (see discussion
of Ogyris barnardi) and the female is therefore in-
cluded as a syntype
Syntyres: 1d (QMT 12349): С.В. [7 or G.P.], A.
Rockhn./14; 1 d(QMT12350); 14; 1 9 (QMTI2351):
Dawson В./0/14. The unlabelled male is probably
from Brisbane.

CURRENT STATUS: Synonym of Eurema smilax (Don-
ovan, [805).

Family NYMPHALIDAE
Subfamily DANAINAE

Euploea amycus Miskin, 1890a:1044 (187)

Described from an unknown number of both sexes
collected at Cape York, Qld, Two specimens haye been
located,

ЅҮМТҮРЕЅ: 10 (QMT12352); Cape York/187, Subse-
quently labelled: Misk, Type 5 ;28/7/10, GAW, CJW
(on reverse)/ L/3003/ Е. amycus $ ; This specimen was
taken for Fig. 18 in "The Butterfiies of Australia" ‚һу
Waterhouse & Lyell (on reverse). 12 (QMT12353):
F.M. -/5/75, Cape York/187. Subsequently labelled:
Misk. Type 9; 28/7/10, GAW, CIW (on reverse)
L/3002.

CURRENT STATUS: Synonym of Euploea algea violetta
(Butler, 1876) (Dunn & Dunn 1991).

Euploea boreas Miskin, 1890a:1043 (189)

Described from an unknown number of both sexes
collected by H. Kuntsler at Cardwell, Qld. Three spec-
imens have been located. Waterhouse's 1910 notes
indicate that a further male from Herbert R. best fitted
the description of the type but this localily was not
mentioned by Miskin (18902) and the specimen may
have been added subsequently.

SYNTYPES: 2d (QMT12354, QMT12355 ): Н.К. Car-
dwell/189 d, 1 2 (QMT12356): Н.К. Cardwell/189 ©.

MISKIN BUTTERFLY TYPES

The female subsequently labelled: Misk. Type 9;
28/7/10, GAW. CJW (on reverse).

CURRENT STATUS: Synonym of Euploea alcathoe
eichhorni Staudinger, 1884.

Euploea crithon Miskin, 1890a:1042 (188)

Described from an unknown number of males col-
lected at Cape York, Qld. Two specimens have been
located.

SYNTYPES; | G(QMTI2357): Cape York/188, Subse-
quently labelled: Е. doriecha ? crithon; This specimen
was taken for Fig. 32 in "The Butterflies of Australia".
by Waterhouse & Lyell (on reverse). 18 (QMT12358):
[no original labels]. Subsequently labelled: Misk, Type
д; 28/7/10. САМ, CIW (on reverse)/ 1/3002.
CURRENT STATUS: Form of Euploea sylvester
(Fabricius, 1793).

Euploea dardanus Miskin, 1890a:104 (204)

Described from (probably one) male collected at Cape
York, Qld,

SYNTYPE: 18 (QMT12359); Cape York/204. Subse-
quently labelled: Misk, Type 3: 28/7/10, GAW, CIW
(on reverse)/ L/2998.

CURRENT STATUS; Form of Euploea sylvester
(Fabricius, 1793).

Euploea euclus Miskin, 18903:1045 (203)

Described from an unknown number of both sexes
collected at Cape York, Old. Two specimens have been
located.

SYNTYPES: 1¢(QMT12360): C. York/203;
19(0МТ12361): Cape York/203/6. Both subsc-
quently labelled: Misk. Type d [or ? ]; 28/7/10, САМ,
CJW (on reverse) L/2999/ E. chauafa corinna ab.
euclus & [or 9]; This specimen was used for Fig, 21
[or23]in "The Butterflies of Australia", by Waterhouse
& Lyell (on reverse),

CURRENT STATUS: Form of Euploea core corinna
(W.S. Macleay, 1826).

Euploea hippias Miskin, 1890a:1040 (206)

Described from (probably one) male collected at Cape
York, Qld.

SYNTYPE: id (ОМТІ2262): E.W. Cape York/206.
Subsequently labelled; Misk. Type д; 28/7/10, GAW,
CIW (on reverse)/ L/3000/ E. hirdafa hippias d ; This
specimen was taken for Fig. 17 in "The Butterflies of
Australia", by Waterhouse & Lyell (on reverse),
CURRENT STATUS: Synonym of Euploea usipeles
Hewitson, 1858.

Euploea misenus Miskin, 1890a:1039 (183)

Described from (probably onc) male collected at Cape
Yark, Qld.

SYNTYPE: 1 Ó (QMT12363): Cape York/183. The fol-
lowing labels added subsequently: Misk, Type d;
28/7/10, GAW, CJW (on reverse)/ L/3001/ E. misenus
d; This specimen was taken for Fig, 14 in "The Bul-
terflies of Australia". by Waterhouse & Lyell (on re-
verse).

CURRENT STATUS: Synonym of Euploea alcatitew
monilifera (Moore, 1883).

Subfamily NYMPHALINAE
Atella propinqua Miskin, 1884b:94 (215)

Described from an unknown number of specimens
collected at Rockingham Bay, Qld. Two of the 3 spec-
imens noted by Waterhouse have been located,

SvurvPES: 1¢(QMT12364); Cardwell/ 215;
i 2(QMT12365): -/5/82 Cardwell/ 215/Female. The
female subsequently labelled: Misk, Type; 28/7/10,
GAW, CIW (on reverse),

CURRENT STATUS: Vagrans egista propinqua (Mis-
kin),

Pyrameis lucasii Miskin, 1889a:1515

Described from 1 specimen from Fernshawe, Victoria,
in the Lucas Collection. It has not been located,
CURRENT STATUS: Synonym of Vanessa kershawi ab.
suffusa (ОПГ, 1888).

Subfamily SATYRINAE
Epinephile rawnsleyi Miskin, 1876:454 (149)

Described from 1 specimen collected by H. C.
Rawnsley at Maroochy River, near Brisbane, Qld,
HoLoTYPE: & (QMT12366): Maroochec/HCR. 149.
Subsequently labelled: Misk. Type d ; 28/7/10, GAW,
CIW (on reverse),

ОГРН Status: Tisiphone abeona rawnsleyl (Mis-
in).

Heteronympha digglesi Miskin, 1876:454
(192

=

Described from an unknown number of specimens (all
males) collected at Brisbane, Old. Two males have
been located.

SvNTYPES: 2 d (QMT12367, QMT12368): 192. One
subsequently labelled: Misk. Type с; 28/7/10, САМ,
CIW (on reverse).

CURRENT STATUS: Synonym of Heteronympha
mirifica (Butler, 1866).

Mycalesis modestus Miskin, 1890¢;29 (198)
Described from an unknown number of specimens

collected at Cooktown, Qld. Two of the 3 specimens
noted by Waterhouse have been located.

522

SyNTYPES: | d. (QMTI2369): D.K. Cooktown/ 198:
19 (QMT12370): D.M.B. Cooktown/198

CURRENT STATUS: Synonym of Mycalests perseus
(Fabricius, 1773).

Xenica kershawi Miskin, 1876:452 (200)

Described from an unknown number of both sexes
collected by W, Kershaw in Victoria, Two specimens
have been located.

SYNTYPES: 1G(QMT12371), 19 (QMT12372): W.K.
Vietoria/200/d [or 9j, The male subsequently la-
belled: Misk. Type 3; 28/7/10, GAW, CIW (on re-
yerse).

CURRENT STATUS: Üreixenica kershawi (Miskin).

Family LYCAENIDAE

Arhopala cyronthe Miskin, 1890c:43 (178)

Described from 2 males collected at Cape York and
Bowen.

SYNTYPES: 15 (QMT12373): Cape York/178. Subse-
quently labelled Misk. Type à: 28/7/10, GAW, CJW
(on reverse)/ L/3010. 1 8 (QMT12374): Bowen/ 178,
CURRENT STATUS: Synonym of Arhopala micale
amytis (Hewitson, 1862). The Bowen specimen is
referrable on geographical grounds (0А, micale amphis
Waterhouse, 1942, placed asa synonym of A. m. атуѓіх
by Dunn & Dunn (1991).

Arhopala eupolis Miskin, 1890c:42 (136)

Described from an unknown number of both sexes
collected at Cape York, Cooktown and Cardwell. Six

specimens have been located.
SyNTYPES 14(0МТ12375}: Cape York/136;
Id (QMTi2376): H Cardwell/136;

29 (QMT12377, QMT12378): - /5/82. Cardwell/136;
19(QMTI2379): Јн, Cooktown/136;
12(QMT12380): 136. The male and 1 female from
Cardwell subsequently labelled; Misk, Type & [or 9 J;
28/7/10, GAW, CJW (on reverse).

CURRENT STATUS; Synonym of Arhopala centaurus
(Fabricius, 1775).

Arhopala wildei Miskin, 1891,71]

Described from | male and 1 female collected by C. J,
Wild at Caims, Qld. Miskin (1891) consistently mis-
spelt Wild's name as Wilde.

Syntypes: 14 (QMT12381), 1 9 (QMT12382): [no
original labels]. Subsequently labelled: L/3006/ Misk.
Type d [or 9]; 28/7/10, GAW, CIW (on reverse).
Female also labelled: Arhopala wildei 9 .

CURRENT STATUS: Arhopala wildei Miskin.

MEMOIRS OF THE QUEENSLAND MUSEUM

Chrysophanus aenea Miskin, 1890c:33 (175)

Described from an unknown number of both sexes
collected in Queensland as far north as Bowen. Three
of the 4 specimens noted by Waterhouse have been
Incated; the male from Bowen has not been found.
SyNTYPES: 15 (QMT12383), 12 (QMT12384): 175,
рт labelled: Misk, Type б [ог 9 J; 28/7/10,

CJW (on reverse), 1 $ (QMT12385): Moggill/
no All these specimens are probably from Brisbane.
CURRENT STATUS: Synonym of Paralucia pyrodiscus
(Doubleday, 1847) (Edwards 1991).

Danis coelestis Miskin, 1891:50 (239)

Described from an unknown number of both sexes
collected by С. J, Wild at Caims, Qld. Two males have
been located, Aithough the species was described from
specimens in the Queensland Museum, the specimen
labelled Cairns (from Miskin's own collection) i is un-
doubtedly from the type series since Miskin gave col-
lection number 240 to specimens of Danis serapis
described at the same time. Only those species in his
own collection were numbered.

ЅҮМТҮРЕЅ: — ld (QMT12386): Cairns/239;
1 d (QMT12387): [no locality data]/ Danis coelestis 3
Misk. The latter specimen subsequently labelled: Misk.
Type д; GAW (on reverse).

CURRENT STATUS: Jamides aleuas coelestis (Miskin),

Danis serapis Miskin, 1891:49 (240)

Described from numerous specimens of bath sexes
collected at Cairns and by H. Kuntsler at Cardwell, Qld.
Three of the 4 specimens noted by Waterhouse have
been located.

Syntypes; 18 (QMT12388), 29(QMTI2389,
QMT12390): Н.К. Cardwell/240. One specimen of
each sex subsequently labelled: Misk. Type д [or Ẹ J;
28/7/10, GAW, CIW (on reverse).

CURRENT STATUS: Danis danis serapis Miskin.

Danis syrius Miskin, 1890c:34 (224)

Described from an unknown number of both sexes
(actually all males) collected m North Queensland.
Two males have been located; one was described as a
female.

SyNTYPES: 1d (QMTI2391): C. York/224. Subse-
quently labelled: Misk, Type d ; 28/7/10, GAW, CIW
(on reverse). 1 d (QMT12392): М. Queensd./224. Sub-
sequently labelled: Misk. Type @ = a male; 28/7/10,
GAW, CJW (on reverse),

CURRENT STATUS: Danis danis syrius Miskin,

Deudorix democles Miskin, 1884b:95 (222)

Described from an unknown number of specimens
collected at Basilisk Range (Johnstone R.), Qld. Wa-

MISKIN BUTTERFLY TYPES 52

terhouse (1902) noted there were 2 males; his 1910
notes record the additional data 5/83,

SYNTYPES: 28 (QMT12393, QMT12394); Basilisk
Range/222. One subsequently labelled: Misk. Type 4;
27/7/10, GAW, CIW (on reverse); bath also labelled:
Fig. 276 (upperside) [or 277 (underside)] in "The But-
terflies of Australia" by Waterhouse & Lyell was taken
from this specimen.

Current STATUS: Virachola democles (Miskin).

Deudorix simsoni Miskin, 1874c:165 (102)

Described from an unknown number of both sexes
collected by Augustus Simson at Port Denison (—
Bowen) and hy Miskin at Brisbane, Qld. Five syntypes
have been located, A female in QM labelled
"Dovercourt [Miskin's home in Toowong, Brisbane],
Ap '74/ 102/ Misk. Type 9; 28/7/10, GAW, СІМ (on
reverse)" is not a type; the original description is dated
18 March 1874, before the specimen was collected.
SYNTYPES; ] d (QMT12395), | ? (QMT12396): 1028
for 7], 10 (QMTI2397), 19 (QMT12398): 102. The
first исеп subsequently labelled: Misk. Type 3;
28/7/10, GAW, CJW (on reverse); all these specimens
аге probably from Brisbane. | C(QMT12399): ALS, А.
Bowen 102.

CURRENT STATUS! Rapala vara simsoni (Miskin).
Holochila albosericea Miskin, 1891:65 (159)

Described from an unknown number of both sexes
collected by George Barnard at Expedition Range (=
Dawson R.), inland from Rockhampton, Qld, Two of
the 3 males noted by Waterhouse have been located.
These appear to have been in Miskin’s Collection since
at least 1875 (see discussion of Ogyris barnardi).
ЗҮМТҮРЕЅ: 20 (QMT12400, QMT12401): Dawson
R./159. One subsequently labelled: Misk. Type;
28/1/10, GAW, CJW (on reverse).

CURRENT STATUS: Nesolycaena albosericeéa (Miskin),

Holochila androdus Miskin, 1890c: 41 (164)

Described from an unknown number of both sexes
from Cape York and Cooktown, Qld. Two specimens
have been located,

ЅҮМТҮРЕЅ; 15 (QMT12402): Cape York; Cockerell,
My '73 (on геуегѕе)/164; 1? (QMTI2403): Cook-
town/164, Both subsequently labelled: Misk. Type d
[or 9 ; 28/7/10, GAW, CIW (on reverse).

CURRENT STATUS: Synonym of Candalides helenita
(Semper, 1879).

Hypochrysops apollo Miskin, 1891: 85 (243)

Described from a supposed female (actually a male)
collected by Rowland Turner al Herbert River, Qld, 1
is in poor condition, having been flattened between the
pages of a book.

р]

Ноцотуре: d (QMT12404); К.Т, Herbert К./243,
Subsequently labelled: Misk. Type 2= d; 28M/T0,
GAW, CIW (on reverse L/3004/ Specimen photog.
for CHECKLIST AUST LEP Film 186/11.

CURRENT STATUS: Hypochrysaps apollo Miskin.

Hypochrysops epicurus Miskin, 1876:455
(114)

Described from am unknown number of bath sexes
collected al Brisbane, Old. Two specimens have been
located. These were designated Lectotype and
paralectotype by Sands (1986), with the collection
number recorded incorrectly as 124,

LECTOTYPE: $ (QMT12405): 114, Subsequently In-
belled: Misk. Type d; 27/7/10, GAW, CJW (on re-
verse)! Lectotype Hypochrysops epicurus Miskin,
1876, designated 1986, D. P A. Sands (red label).
PARALECTOTYPE: 12 (QMT12406): [label glued to
insect and number not visible], Subsequently labelled:
Misk. Type 9; 27/7/10, САМ, CJW (on reverse
Paraleclotype Hypochrysops epicurus Miskin, 1876,
designated 1986, D. P. A. Sands (blue label).
CURRENT STATUS: Hypochrysops epicurus Miskin.

Hypochrysops euclides Miskin, 18892; 1517

Described from an unknown number of both sexes
from Gippsland, Victoria in the Lucas Collection. 'The
type locality is evidently erroneous (Sands 1986) since
this taxon occurs only in North Queensland. Syntypes
not located; believed destroyed (Waterhouse 1903).

RENT STATUS: Hypochrysops pythias euclides
Miskin.

Hypochrysops Moralis Miskin, 1884b:94
(218)

Described from (probably one) female collected by W.
Kershaw in Victoria, designated Lectotype by Sands
(1986), Miskin (1889a; 1516) subsequently described
the male from the Lucas Collection.

LECTOTYPE: 9 (QMTI2407): W.K. Victoria/218,
Subsequently labelled: Misk. Type 2; 27/7/10. САМУ,
CJW (on reverse)! Lectotype Hypochrysops hecalius
Miskin, 1884, designated 1986, D. P, A. Sands (red
label).

CURRENTSTATUS: Hypochrysops byzos hecalius Mis-
kiñ.

Hypochrysops olliffi Miskin, 1889a:1518 (237)

Described from a single male collected by A. S. Olliff.
The original description gave the type locality as both
Newcastle, NSW and Fremantic, WA. The Holotype
was stated to be in the Australian Museum (Miskin
18892) but Waterhouse and Lyell (1914) noted that no
specimens from Newcastle were present there and re-
stricted the type locality to SW Western Australia. A
specimen Without locality data (KL 23213) in the Aus-

524

tralian Museum is possibly the holotype (Sands 1986).
The locality Newcastle must be considered an error,
although Miskin (1889a:1520) evidently believed the
species occurred in eastern coastal Australia. It is pos-
sible that the locality Fremantle was intended as a
correction rather than an addition.

In the QM Collection are 2 specimens (14,1), la-
belled "Olliffi, NSW/237", These are specimens of
typical H. ignitus ignitus (Leach) and were evidently
added to Miskin's collection after his description of H.
ОШ, since he noted (18898) that the female was
unknown,

CURRENT STATUS: Hypochrysops ignitus olliffi (Mis-
kin).

Talmenus eubulus Miskin, 1876: 457 (91)

Described from an unknown number of specimens
from Rockhampton, Qld. Three specimens have been
located, The Dawson River males appear to have heen
in Miskin's Collection since at least 1875 (see discus-
sion of Ogyris barnardi) and are regarded as synt ypes.
The Rockhampton female probably came from further
inland, since this taxon is not known from coastal arcas
(Common & Waterhouse 1981); like the Dawson R,
specimens, it may have been collected by George Bar-
nard.

Syntypes: 1 9 (QMT12408): О.Р. [? or О.В.], A.,
Rockhampton/91, Subsequently labelled: Misk, Type
2:28/7/10, GAW, CIW (on reverse), 28 (QMT 12409,
QMT12410); Dawson R./91.

CURRENT STATUS: Jalmenus evagoras eubulus Mis-
kin.

Talmenus itonus Miskin, 1890c: 41 (152)

Described from an unknown number of males col-
lected at Cape York, Qld. Waterhouse's 1910 notes list
2 males; 1 has been located.

SYNTYPE: 1d (QMTI2411); Саре York/152, Subse-
quently labelled: Misk. Type с ; 28/7/10, GAW, CIW
(an reverse].

CURRENT STATUS: Synonym of Jalmenus ejchhorni
Staudinger, 1888.

Lycaena canescens Miskin, 1890c:35 (163)

Described from (probably one) specimen collecied by
(?) Charles French in Tasmania.

SYNTYPE: 15 (QMT12412): C.F. Tasmania/163. Sub-
sequently labelled: Misk, Type d; 28/7/10, GAW,
CJW (on reverse).

CURRENT STATUS: Synonym of Candalides acastus
(Cox, 1373).

Lycaena gracilis Miskin, 1890c:37 (162)

Described from an unknown number of both sexes
collected from Brisbane to Cooktown, Qld. Two spec-

MEMOIRS OF THE QUEENSLAND MUSEUM

imens. (from Rockhampton and Brisbane) of the 7
noted hy Waterhouse have been located,

SvwrYPES 18 (QMTI2413): Rockh"/162;
1 2(QMT12414): Toowong, Mar. "73/162.

CURRENT STATUS: Synonym of Famegana alsulus
(Herrich-Schaeffer, 1869).

Lycaena hobartensis Miskin, 1890с:38 (223)

Described from an unknown number of specimens
collected by Gervase F. Mathew at Hobart, Tasmania,
Waterhouse's 1910 notes list a pair but only the male
has been located.

SywTYPE: 1d. (QMT12415): G,F.M, Hobart/ 223.
Subsequently labelled: Misk. Type 4; 28/7/10, САМ,
CJW {on reverse),

CURRENT STATUS: Neolucia hobartensis (Miskin),

Lycaena mackayensis Miskin, 1890c:35 (230)

Described from (probably one) male collected at
Mackay, Qld. Waterhouse’s 1910 notes also record the
data R. T[urner], 2/89.

SYNTYPE: 18 (QMT12416); Mackay/230. Subse-
quently labelled: Misk. Type 3; 28/7/10, GAW, CIW
(on reverse) L. mackayensis Misk. 230,

CURRENT STATUS: Synonym of Prosotas dubiosa
(Semper, 1879).

Lycaena mathewi Miskin, 1890c:38 (89)

Described from (probably one) specimen collected by
Gervase F. Mathew at Sydney, NSW. Waterhouse’s
1910 notes record the additional data 12/82,
SYNTYPE: 19 (QMT12417); G.F.M. Ѕуйпсу/89, Sub-
sequently labelled: Misk. Type 9;28/7/10, GAW,
CJW (on reverse),

CURRENT STATUS: Neolucia татем (Miskin),

Lycaena nigra Miskin, 1890c:36 (216)

Descnbed from an unknown number of "females"
collected at Cardwell, Qld. Waterhouse's 1910 notes
lista pair but only the male (described as a female) has
been located.

LECTOTYPE: d (QMTI2418): -/5/82 Cardwell/216,
Subsequently labelled: Misk, Type 9 = d; 28/7/10,
GAW, CIW (on reverse). Called Holotype by Eliot and
Kawazoe (1983), thereby designated as Lectotype "by
inference of Holotype" under the rules of the ICZN,
CURRENT STATUS: Megisbe strongyle nigra (Miskin),

Lycaena sulpitius Miskin, 1890c:37 (181)

Described from (probably one) specimen collected at
Rockhampton, Qld.

SYNTYPE: 1d. (QMTI2419): Rockh"/181 [red ink].
Subsequently labelled: Misk. Type d; 28/7/10, GAW,
CIW (on reverse).

CURRENT STATUS: Theclinesthes sulpitiuy (Miskin).

MISKIN BUTTERFLY TYPES

Lycaena tenella Miskin, 1891:63

Described from (probably one) female collected by C
J. Wild at Caims.

LECTOTYPE: 9 (QMT12420): [no locality datal/ Lyc.
fenella n. sp. 9 . Subsequently labelled: Misk. Type 2;
28/7/10, GAW, CJW (on reverse). Called Holotype by
Eliot and Kawazoe (1983), thereby designated as
Lectotype "by inference of Holotype".

CURRENT STATUS: Udara renella (Miskin).

Lycaenesthes tasmanicus Miskin, 1890c:40
195

Described from (probably one) male collected by Syl-
vester Diggles supposedly in Tasmania, The type lo-
cality was doubted by Miskin (1891) who recorded the
species from Cairns. Waterhouse's 1910 notes indicate
amale from "Cape York or Tasmania", The type-local-
ity is evidently erroneous.

SYNTYPE: 1d (QMTI2421): S.D. 22/11/75 Tasrna-
nia/195. Subsequently labelled: Misk. Type d;
28/7/10, GAW, CJW (on reverse).

CURRENT STATUS: Erysichton palmyra tasmanicus
(Miskin).

Lycaenesthes turneri Miskin, 1890c;39 (180)

Described from an unknown number of both sexes
collected at Cape York, Card well and Mackay (1 pair).
Four specimens have been located, at least one col-
lected by Rowland Turner at Mackay.

SyntyPes: 1d (QMTI2422): Cape York/180;
19(QMT12423): -/5/82 Cardwell/180;
1 (QMT12424): Mackay/180; 1 ? (QMT12425): К.Т.
Mackay/180. The Mackay specimens subsequently la-
belled: Misk. Type d [ог 9 ]; 28/7/10, САМУ, CJW (on
reverse),

CURRENT Status: Synonym of Anthene lycaenoides
godeffroyi (Semper, 1879).

Ogyris barnardi Miskin, 1890b:27 (174)

Described from an unknown number of both sexes
collected by George Barnard at Dawson River (= Ex-
pedition Range), Old. Two specimens have been lo-
cated. These appear to have been in Miskin's
Collection since at least 1875, since collection numbers
up to 200 appear for species described by Miskin
(1876) and species numbers 187 (Euploea amycus) and
195 (Lycaena tasmanicus) were collected in May and
November 1375 respectively.

SvNTYPES: 18 (QMT12426), 1 ? (QMT12427): Daw-
son R./174. Both subsequently labelled: Misk. Type с
[ог 9 1; 28/7/10, GAW. CJW (on reverse). Male also
labelled: Specimen photog. for CHECKLIST AUST
LEP Film 186/12; female also labelled; L/3009, The
female has lost its hindwings.

CURRENT STATUS; Ogyris barnardi Miskin.

щл
T2
n

Pseudodipsas brisbanensis Miskin. 1884b:95
(202)

Described from (probably one) female collected at
Brisbane, Qld.

LECTOTYPE: ? (QMT12428): 202. Subsequently la-
belled: Misk. Type 9; 28/7/10, GAW, CJW (on rc-
verse)! T6193, Type [red label] Specimen photog. for
CHECKLIST AUST LEP Film 186/10. Called Holo-
type by Sands (1980), thereby designated Lectotype
"by inference of Holotype”.

CURRENT STATUS: Acrodipsas brisbanensis (Miskin),

Pseudodipsas fumidus Miskin, 1889c:264

Described from an unknown number of both sexes
from Brisbane, Qld, in the Lucas Collection, Not lo-
cated

CURRENTSTATUS: Synonymof Psendodipsas серћепех
Hewitson, 1874.

Pseudodipsas innotatus Miskin, 1874c:165
(93)

Described from an unknown number of both sexes
collected by Miskin at Brisbane, Qld. None of the
original specimens has been located; the 2 males and |
female listed by Waterhouse in his 1910 notes may
have been added subsequently. It is species number 98
in Miskin's notebook,

This species was treated as a synonym of P. ilias
(Felder) by Miskin (1891).

CURRENT STATUS: Philiris innotara (Miskin).

Family HESPERIIDAE
Apaustus minimusMiskin, 1889c: 153 (227)

Described from an unknown number of specimens
collected in Victoria and Western Australia. None
ofthe 4 males listed hy Waterhouse in his 1910 notes
(where he recorded the type as à Victorian male) has
been located, although Waterhouse (1937) called à
specimen from Western Australia (then 1n QM) the
Holotype, thereby designating the Lectotype "by infer-
ence of Holotype". The species is numbered 227 in
Miskin's notebook.

CURRENT STATUS: Synonyms of Taractrocera papy-
ria papyria (Boisduval, 1832) (Victorian specimens)
and Т, papyria agraulia (Hewitson, 1868) (WA speci-
mens, including Lectotype). This synonymy was also
suggested by Miskin (1891: addenda).

Hesperilla atromacula Miskin, 1889c: 148
(234)

Described from an unknown number of males col-
lected in Victoria. Two have been located,

526

Waterhouse's 1910 notes also list a female; this was
presumably added later to Miskin's Collection,
LECTOTYPE: d (QMT12429): Victoria/234. Subse-
quently labelled: Misk. Type d ; 27/7/10, GAW, CIW
(on reverse). Called the Holotype by Waterhouse
(1937), thereby designated as Lectotype "by inference
of Holotype".

PARALECTOTYPE: 1 d (QMT12430): Victona/234.
CURRENTSTATUS: Synonym of Signeta flammeata
(Butler, 1882).

Hesperilla croceus Miskin, 1889c;150 (130)

Described from an unknown number of both sexes
collected from Brisbane to Cooktown. Two specimens
have been located.

LECTOTYPE: d (QMT12431): G.J.M. Cooktown/130,
Subsequently labelled: Misk. Type d; 27/7/10, САМ,
CIW (an reverse)’ Neohesperilla crocea (Miskin).
Called the Holotype by Waterhouse (1937), thereby
designated as Lectotype “by inference of Holotype".

PARALECTOTYPE: 1 P (QMT12432): 130. This speci-
men is probably from Brisbane,

CURRENT STATUS: Neohesperilla crocea (Miskin)
(Lectotype) and synonym of Neohesperilla
xanthomera (Meyrick & er, 1902) (Paralectotype).

Hesperilla fulgidus Miskin, 1889c:151 (137)

Described from (probably one) female (actually а
male) collected at Brisbane, Qld. No located although
Waterhouse (1937) recorded it (then in QM) as the
Holotype male, thereby designating it as Lectotype "by
inference of Holotype". A female in QM labelled
"Mackay/ 137" is mentioned in Miskin (1891) but not
in the original description (Miskin 1889c) and is not a
type. Waterhouse's. 1910 notes list 3 further males
which presumably were also added later.
CURRENTSTATUS: Synonym of Parnara amalia (Sem-
per, 1879).

Hesperilla humilis Miskin, 1889c:150 (150)

Described from an unknown number of both sexes

collected at Brisbane, Qld. None located, although

Waterhouse's 1910 notes list 2 males and 1 female.

Waterhouse (1937) recorded the Holotype male (then

in QM), thereby designating it as Lectotype "hy inler-

are of Holotype". It is number 150 in Miskin's note-
юк.

CURRENT STATUS: Synonym of Toxidia parvula
(Plotz, 1884),

Hesperilla senta Miskin, 1891:85

Described from | female collected by C, J. Wild at
Herberton, Qld.

НОГОТҮРЕ: 9 (QM12433): Herberton, ij, 1891, C. J.
Wild/ Hesperilla senta Miskin. Subsequently labelled:

MEMOIRS OF THE QUEENSLAND MUSEUM

Misk. Type 9; 27/7/10, GAW, CJW (on reverse)/
1/3016,
CURRENT STATUS: Neohesperilla senta (Miskin).

Hesperilla tasmanicus Miskin, 1889¢; 149
(128)

Described from an unknown number of "females"
collected in Tasmania. Waterhouse's 1910 notes list 2
specimens; only a male (described as a female) has
been located,

LECTOTYPE: & (QMT12434): G.B. Tas[mania] [label
torn off]/128. Subsequently labelled: Misk. Type 9;
27/1/10, GAW, CIW (on reverse). Called Holotype by
Waterhouse (1937), thereby designated Lectotype "by
inference of Holotype".

CURRENT STATUS: Pasma tasmanica (Miskin),

Pamphila albifascia Miskin, 1889c:148 (43)

Deseribed from (probably one) male collected at Her-
bert River, Qld. Waterhouse's 1910 notes indicale that
this species was evidently added at à late stage to the
Miskin Collection, being placed early in the notebook
wo fill a vacant number. Miskin (1891) placed this
species as a synonym of Carystus caesina Hewitson.

LECTOTYPE: 2 (QMTI2435): Herbert R./43, Subse-
quently labelled: Misk, Type $; 27/7/10, GAW, CIW
(on reverse). Called Holotype by Waterhouse (1937),
thereby designated as Lectotype "by inference of Ho-
lotype".

CURRENT STATUS: Sabera caesina albifascia (Mis-
kin),

Pamphila autoleon Miskin, 1889c:147 (226)

Described from an unknown number of "males" in the
Miskin Collection and females in the Lucas Collection,
both collected at Cardwell, Qld. Waterhouse's 1910
notes list 2 females from Cardwell, 7/84, in the Miskin
Collection; only one (described as а male) has been
located,

Lectoryre: © (QMT12436): Cardwell/ 226. Subsc-
quently labelled: Misk, Type à = 9; 27/7/10, GAW,
CIW (on reverse). Called Holotype by Waterhouse
(1937), thereby designated as Lectotype "by inference
of Holotype".

CURRENT STATUS; Sahera dobbor autoleon (Miskin),

Pamphila fuliginosa Miskin, 1889c:147 (197)

Described from an unknown number of females col-
jected at Cardwell, Qld. Waterhouse's 1910 notes list
2 females from Cardwell; only 1 has been located. The
male was described subsequently by Miskin (1891: 76)
from Cairns.

LECTOTYPE: 9? (QMT12437): D.M.B. Cardwell/ 197.
Subsequently labelled: Misk. Type 9; 27/7/10, САМ,

MISKIN BUTTERFLY TYPES 32

CJW (on reverse). Called Holotype by Waterhouse
(1937), thereby designaled as Lectotype "by inference
of Holotype".

CURRENT STATUS: Sabera fuliginosa (Miskin).

Tagiades gamelia Miskin, 1889c:146 (127)

Described from an unknown number of both sexes
collected at A Сори York, Qld. Three specimens have
been located

LECTOTYPE: d (QMT 12438): Cape York/127. Subse-
quently labelled: Misk, Type d ; 27/7/10, САМ, CJW
(an reverse)/ L/3015. Called Holotype by Waterhouse
(1937), thereby designated as Lectotype "by inference
of Holotype" .

PARALECTOTYPES: | Ф (QMT12439): Cape York/127,
Subsequently labelled: Misk, Ty уре 9; 27/7/10, САМ,
EN on reverse)! LJ3015. 1 (QMT12440): Саре

or

CURRENT STATUS: Synonym of Tagiades japetuis
janerta Butler, 1870.

Trapezites idothea Miskin, 1889c:152

Described from 1 specimen from Victoria in the Lucas
Collection. The Holotype female is deposited in the
South Australian Museum (Waterhouse 1933, 1937).
CURRENT STATUS: Hesperilla idothea (Miskin),

Trapezites phillyra Miskin, 1889c: 153.

Described from an unknown number of both sexes
from Victoria in the Lucas Collection. Waterhouse
(1937) recorded the Holotype male at the South Aus-
tralian Museum, thereby designating the Lectotype "by
inference of Holotype".

CURRENT STATUS: Synonym of Trapezites phigalia
(Hewitson, 1868).

ACKNOWLEDGEMENTS

I wish to thank С. B. Monteith and С. Thomp-
son (Queensland Museum) for their assistance
with this study and access to specimens, and Е,
D. Edwards (ANIC) for the loan of G. A.
Waterhouse's unpublished notes on the Miskin
Caliection.

REFERENCES

COMMON, І, F, B, & WATERHOUSE, D. Е, 1981,
"Butterflies of Australia." (Angus & Robertson:

Sydney),

DE BAAR, M. & HANCOCK, D. L. 1993, The Austra-
lian species of Elodina С. & R. Felder (Lepidop.
tera: Pieridae). Australian Entomologist 2
25-43.

DUNN, K. L, & DUNN, L, E. 1991. “Review af Aus-
tralian butterflies: distribution, life history and

~

taxonomy." 4 parts. (Privalely published: Mel-

bourne),

EDWARDS, E. D. 1991. The nomenclature of
Paralucia pyrodiscus (Lepidoptera: Lycaenidae}.
(Doubleday). Australian Entomological Maga-
zine 18: 27-30.

ELIOT, J. N. & KAWAZOE, A, 1983. "Blue butterflies
of the Lycaenopsis group". (British Museum of
Natural History: London).

MISKIN, W. H. 1871. Occurrence of Danais archippus
in Queensland. Entornologists Monthly Maga-
zine 8: 17.

1874a. Note on Mynes guerini, Wallace, Transac
tions of the Entomological Society of London
1874(2): 237-240.

1874b, Note on "A catalogue of the described diur-
nal Lepidoptera of Australia, by Mr. George Mas-
lers, of the Sydney Museum." Transactions of the
Болур Society of London 1874(2); 241-

46.

1874c. New Lycaenidae from Queensland.
Entomologist's Monthly Magazine 11: 165.

1876. Descriptions of new species of Australian
diurnal Lepidoptera, Transactions of the Entomo-
logical Society of London 1876(3): 451-457.

1883, On Ogvris genoveva, Hewitson, and its life
history. Transactions of the Entomological Soci-
ety of London 1883(4); 343-345, pl. XV.

1884a. Note on Tachyris melania of Fabricius.
Transactions of the Entomological Society of
London 1884(1): 91-92.

1884b. Descriptions of new Australian
Rhopalocera. Transactions of the Entomological
Society of London 1884(1): 93-96,

1888. Papilio parmatux (G. R. Gray) at Mackay.
Proceedings of the Royal Society of Queensland
4: 17.

18893, Descriptions of hitherto undescribed Austra-
lian Lepidoptera (Rhopalocera). Proceedings of
the Linnean Society of New South Wales 3(4):
1514-1520.

1889b. Note on Danais chrysippus (L.), and D.
petilia (Stoll). Proceedings of the Linncan Soci-
ety of New South Wales 4(1); 119-120.

1889c. Descriptions of some new species of Austra-
lian Hesperiidae. Proceedings of the Royal Soci-
ety of Queensland 6(4): 146-154.

1889d, Revision of the Australian species of the
Lepidopterous genus Terias, with descriptions of
some new species, Proceedings of the Royal Sa-
ciety of Queensland 6(5): 256-263.

1889c. Note on some undescribed Australian Lepi-
doptera (Rhopalocera). Proceedings of the Royal
Society of Queensland 6(5): 263-266.

1890a. A revision of the Australian species of Ew-
ploea, with synonymic notes, and descriptions of
new species. Proceedings of the Linnean Society
af New South Wales 4(4): 1037-1046.

189Db. A revision of the Australian genus Ogvris,
with description of a new species. Proceedings of

528

the Linnean Society of New South Wales 5(1):
23-28.

1890c. Descriptions of hitherto undescribed Austra-
lian Lepidoptera (Rhopalocera) principally
Lycaenidae. Proceedings of the Linnean Society
of New South Wales 5(1): 29-43.

1890d. Note on Danais petilia, Stoll. Proceedings
of the Linnean Society of New South Wales 5(1):
142.

1891. A synonymical catalogue of the Lepidoptera
Rhopalocera (butterflies) of Australia with full
bibliographical reference; including descriptions
of some new species. Annals of the Queensland
Museum 1: i-xx, 1-93, i-ix.

SANDS, D. P. A. 1980. A new genus, Acrodipsas, for
a group of Lycaenidae (Lepidoptera) previously
referred to Pseudodipsas C. & R. Felder, with
descriptions of two new species from northern
Queensland. Journal of the Australian Entomo-
logical Society 18: 251-265.

MEMOIRS OF THE QUEENSLAND MUSEUM

1986. A revision of the genus Hypochrysops C. &
R. Felder (Lepidoptera: Lycaenidae). Entomono-
graph 7: 1-116.

WATERHOUSE, G. A. 1903. Notes on Australian
Rhopalocera: Lycaenidae. Part Ш.- Revisional.
Proceedings of the Linnean Society of New South
Wales 28(1); 132-275, pls i-iii.

1933. Notes on the type specimens of Hesperiidae
(Lepidoptera) in the museums in Australia, with
special reference to those in the South Australian
Museum. Records of the South Australian Mu-
seum 5(1): 49-62.

1937. Australian Hesperiidae. VII. Notes on the
types and type localities. Proceedings of the
Linnean Society of New South Wales 62(34):
107-125.

WATERHOUSE G. A. & LYELL, G. 1914. "The but-
terflies of Australia. A monograph of the Austra-
lian Rhopalocera." (Angus & Robertson:
Sydney).

EFFECTS OF PRESCRIBED BURNING ON HERPTILES IN SOUTHEASTERN
QUEENSLAND

DAVID S, HANNAH AND GEOFFREY C. SMITH

Hannah D.S, & Smith, С.С. 1995 12 01; The effects of prescribed burning on herptiles in
Southeastern Queensland. Memoirs of the Queensland Museurn 38(2):529-531. Brisbane.

ISSN 0079-8835.

Study of herptile faunas within native forests and managed exotic pine plantations, subject
to different burning regimes, produced inconsistent results with respect to the effects of fire
Management on Vertebrate diversity and species richness. Abundance varied markedly
among burning regimes in native forest areas (SPAT). [] Herptiles, diversity, burning,

Species richness.

David S, Hannah and Geoffrey С, Smith, Queensland Forest Research Institute, DPT Forest
Service, 80 Meiers Rd, Indooroopilly, Queensland 4068, Australia; 19 June 1995.

Information on the long term effects of repeated
fires on fauna appears to be scant (Christensen &
Abbott, 1989). There are few studies on the ef-
fects of fire on reptiles (Lunney et al., 1991) and
fewer on amphibians (Bamford, 1992). The
Beerburrum Plantation Complex (centred on
Beerburrum Forestry Office: 26° 57° S, 152° 57’
E), some 50 km north of Brisbane, incorporates
relatively undisturbed native forest and managed
plantations of exotic pine. These have mainly
been burnt at 2-3 year intervals. There has been,
however, no assessment of the impact of this or
any other fire management regime on the fauna
of these forests.

"This study aimed to provide information on the
diversity and density of berptiles in forests sub-
ject to different burning regimes. The native veg-
etation study was located in Scientific Purposc
Arca ] (SPA1). Vegetation types in the SPAI
include heath, herblands, shrublands, and Scrib-
bly Gum/Bloodwood associations. Fire treatment
plots consisted of a control (unburnt since 1973),
plots burnt during autumn-winter on a 3 year
cycle (last burned 1991) and during winter-spring
on a 5 ycar cycle (last burned 1993). Each plot
was approximately 1.5 ha, with two replicate
plots per treatment. Data from these replicates
were pooled. The plantation study was located in
the Twins Plantation Area (TPA), established in
1947-49 with Pinus elliotii, P. carribaea var.
hondurensis, hybrids of these species, and rem-
nant riparian and swamp vegetation. Treatments
werc located in three separate compartments: a
control (unburnt since establishment) and two
treatments (without fire for 7 years and without
fire for 2 years). There was no regular burning, as
per the ЅРА1. Compartments averaged 25ha in

area, systematic sampling targeted three lha
quadrats per compartmenttreatment.

Systematic sampling involved pitfall trapping with
drift fences und active searching. Five pitfall traps,
400 mm deep by 240mm diameter, placed at five
metre intervals, connected by a 300mm high alumin-
ium fly wire drift fence, formed a pitfall line. There
were б and 3 pitfall lines per treatment in the SPA]
and TPA, respectively. The SPA] plots were sur-
veyed over 6 consecutive days each (11-28 January
1994) and the TPA over 5 consecutive days each (24
March-20 April 1994). There were 180 and 75 pitfall
trapping nights of effon per teatment in the SPAI
and TPA, respectively. A Diversity Index (Shannon
& Weaver, 1949) was calculated for each treatment
based on pit trapping data. Study areas were also
actively and systematically searched. Incidental ге»
cords were made in and around the study areas.

Systematic pit trapping and active searches re-
corded 133 individuals of 11 species of reptiles
and 263 individuals of 4 species of amphibians in
the SPA], and 54 individuals of 9 species of
reptiles and 124 individuals of 9 species of am-
phibians in the TPA (Table 1), Identifications
were made by DH. With the exception of
Rhinoplocephalus boschmai cf. and Litoria
brevipalmata, the species recorded were typical
for forests of the region. The Rhinoplocephalus
spp. was uncharacteristic of R, nigrescens and
was recorded as R. boschmai, L. brevipalinata
was unexpected from the pine plantation; but the
specimens caught and released in the TPA, con-
formed fully with the description by Cogger
(1992). Specimens from the same locality have
since been identified by G. Ingram (Qld Museum,
pers. comm.). L. brevipalmata 15 listed as rare in
Queensland (ОРЕН, 1993) along with Crinia
tinnula, the most commonly caught species m the

530 MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 1.

NL
|

| REPTILES
| Agamidae

Incidental

Diporiphora australis
Pogona barbata

{Cobptonsseuirovmom | |} |
[Cryptoblepharus virgaus | -2) | -2) | -w | |
[жегир EM PE TE a
Selanne at | Ea a
IL. HR —1
[Lampropholisamicua | | | J —— ]| 4 | .— [| 13 [| |
раат А
Lygisaurus s foliorum
a = г г ү E ЕЕ
Culte

IDendreaphspunclata | | | 0 — Др
|Tropidonophis mairi | — | | — | Prset | | | | Present |
[Demansiapsammophis | | 1 | | lol] | | |
empresas [йр {J г И
[Rhinoplocephalusboschmai | | — — | — — | Prem | | | | |
ЕЕ
ilisbutmis | — | J T T Present |
| AMPHIBIANS
Myobatrachidae
lAdelowsbrevis | J o To | J —[ x3 |, [ |
|Crmatmua — — 3$ | 3310 | soa | | 1 | — ]| J — |
immdymsesomaus | | J Ç Jof CT
Limnodynastesperoni | | 2% | зч» | [|n]
[Limnodynastes terraereginae | 20) | 5 |

Pseudophryne coriacea =
Hylidae
[Litoria brevipalmaa | | | | -(2)
lLitoriacaerulea | | | Present | | |
(Litoria falas | — | reset | | | [Pre
Mioralaopdmaa —— | — | | | —— | — | «m |] ë T ë
lLitorianasuta | oo | — |] — | |] do | — |
Bufonidae
[Bufo marinus — — | 40) | 1806) | 4703) | | 220) | по | 9 [ |
|TOTALNUMBERS — [6195| 7946 | 305| [|4200| 3409 | aay | |
|NMBEROFSPECES | 9 | 2 | wo | s | B | 9 | 9 | 3
[SHANNONINDEX — | 118 | 1.56 | 125 | — | 191 | 126 | 242 | |

EFFECTS OF BURNING ON HERPTILES

SPA1. The introduced Bufo marinus was com-
mon in both study areas. It associated signifi-
cantly with the treatment burned every 5 years
(fired 6 months previously) and avoided the con-
trol (Х2=41.8, df=2, Р<0.05) in the ЅРАІ. In
contrast, it associated significantly with the con-
trol in the TPA (X2=7.0, df=2, P«0.05).

The treatment burnt every 3 years (fired two
and a half years previously) within the SPA1 and
the treatment without fire for 7 years in the TPA
yielded the highest Diversity Indices. The 3 year
burnt treatment within the SPAT and the control
within the TPA returned the highest species num-
bers (pit-trapping and searching combined); there
were no significant differences among treatments
within either the SPAT or TPA. The study sug-
gests that fire management does not consistently
affect diversity and species richness of herptiles.
However, abundance did vary markedly among
burning regimes in the SPA] (X?=42.3, df=2,
P<0.05), with more than expected numbers oc-
curring within the treatment burnt every 5 years
(fired 6 months previously) and fewer in the
unburnt control; differences were partly ex-
plained by B. marinus captures, but also
Lampropholis delicata captures.

The numbers of species (excluding incidental
records) were comparable between the SPA 1 and
TPA (15 and 18 respectively), despite differences
in trapping and searching effort. Nevertheless,

531

species composition was different in the pine
plantation compared to native vegetation.

ACKNOWLEDGEMENTS

Thanks to Glenda Head and Peter Schellenber-
ger for logistical support; to Patrick Couper, Glen
Ingram and Greg Czechura for assistance with
identification. This project was supported by the
DPI Forest Service.

LITERATURE CITED

BAMFORD, M. 1992, The impact of fire and increasing
time after fire upon Helioporus eyrei,
Limnodynastes dorsalis and Myobatrachus
gouldii (Anura: Leptodactylidae) in Banksia
woodland near Perth, Western Australia. Wildlife
Research 19; 169-78,

COGGER, H.S. 1992, Reptiles and amphibians of Aus-
tralia. (Reed Books; Sydney).

LUNNEY, D., EBY, P, & O'CONNELL, M. 1991.
Effects of logging and fire on small mammals in
Mumbulla State Forest, near Bega, New South
Wales. Australian Wildlife Research 14: 163-81.

QUEENSLAND DEPARTMENT OF ENVIRON-
MENT AND HERITAGE 1993, Preliminary list
of rare and threatened vertebrates in Queensland,
(Department of Environment and Heritage: Bris-
bane).

SHANNON, C.E. & WEAVER, W. 1949, The mathe-
matical theory of communication. (University of
Illinois: Urbana).

TABLE 1. Numbers of herptiles captured in pitfall traps and by active searching (in parentheses) in control and
fire treatments in SPA1 and TPA, Other incidental records are provided for each study area. Shannon index for
pit trapping only.

532

RECORD OF THE SCORPION LIOCHELES
KARSCHII (KEYSERLING, 1855) FROM NORTH
EAST QUEENSLAND (SCORPIONIDA:
ISCHNURIDAE). Memoirs of the Queensland Museum
38(2):532. 1995:- The ischnurid scorpion Liocheles karschii
(Keyserling, 1855) is known from New Guinea and adjacent
islands. In Australia the species is reported only from islands
of Torres Strait (Koch, 1977), but in December 1990 many
specimens were observed in boulder caves in Black Mountain,
20km S of Cooktown, NE Queensland. A female specimen
was collected from the Black Mountain caves and has been
deposited in the Museum of Tropical Queensland, Townsville
QMS105138).

Liocheles karschii is distinguished from other species of
Liocheles in Australia by the large adult size (carapace length
greater than 13.6mm) and the presence of a smooth continuous
groove on the manus of the pedipalp in which trichobothria
dst, dsb and db occur (Koch, 1977).

Liocheles karschii is not a troglobite, although it was
abundant in the caves in Black Mountain. This species has
been collected under logs in New Guinea (Koch, 1977), and
may be found in the woodlands surrounding Black Mountain,
although this habitat has not yet been investigated. Black

MEMOIRS OF THE QUEENSLAND MUSEUM

Mountain is constructed of granite boulders and its caves are
interconnected gaps between the boulders. Unlike solution
caves and lava tubes in which entry and egress is limited to
one or a few points, the Black Mountain boulder caves are
accessible at many points, Nurnerous other arthropods includ-
ing opiliones, archaeognathids and centipedes were also ob-
served in these caves.

This record extends the known range of Liocheles karschii
by approximately 600km, an confirms the presence of this
species in mainland Australia.

Literature Cited

Koch, L.E. 1977. The taxonomy, geographic distribution and
evolutionary radiation of Australo-Papuan scorpions.
Records of the Western Australian Museum 5(2): 83-
367.

Jamie Seymour, Erich Volschenk & Bronwen Scott, Depart-
ment of Zoology, James Cook University of North Queens-
land, Townsville, Queensland 4811, Australia; 19 October
1995.

BRYOZOA FROM HERON ISLAND, GREAT BARRIER REEF. 2
P.J. HAYWARD & J.S. RYLAND

Hayward,P.J. & Ryland, J.S. 1995 12 01: Bryozoa from Heron Island, Great Barrier Reef.
2. Memoirs of the Queensland Museum. 38(2):533-573. Brisbane. ISSN 0079-8835.

Forty-six species of Bryozoa are reported from shallow reef habitats of Heron Island,
Queensland. Twenty new species are described: Chaperiopsis harmeri, Beania pectinata,
Thairopora calcarata, Exechonella ampullacea, Celleporaria hesperopacifica, С, sicaria,
Calyptotheca rupicola, Rabertsonidra praecipua, Cheiloporina campanula, Stylopoma
herodias, Nimba saxatilis, Cigclisula fruticosa, Smittoidea incucula, Pleurocodonellina
laciniosa, Fenestrulina epiphytica, Celleporina bellatula, C. cochlearia, C. fistulata,
lodictyum mamillatum and I. receptaculum. Five species are reported for the first time from
Australian waters. [ ] Heron Island, Great Barrier Reef, Bryozoa, taxonomy.

P.J- Hayward & J.S. Ryland, Marine and Environmental Research Group, School of
Biological Sciences, University of Wales, Swansea, Singleton Park. Swansea SA2 8PP,

United Kingdom; I September 1995.

Tropical bryozoan faunas are rich in species but
poorly known taxonomically, Current knowledge
is still largely founded on a very few mono-
graphic accounts (e.g., Canu & Bassler, 1929;
Harmer, 1915, 1926, 1934, 1957), the usefulness
of which is limited by the vast geographical areas
they cover, and by their outdated and conserva-
live taxonomy. Bryozoans are especially abun-
dant in coral reef communities where they are an
important part of cryptic, sessile faunas. Yet, as
Winston (1986) showed, a century of ecological
research into reef-associated faunas resulted in a
world-wide checklist of just 284 species of
Bryozoa. It had been assumed that bryozoans
were of little consequence in reef ecosystems,
largely through unfortunately biased collecting
(Winston, 1986), and it was not until reef cecolo-
gists began to examine sessile reef communities
that their true significance began to be revealed
{e.g., Jackson & Winston, 1982). Faunistic stud-
ies of limited areas of reef (e.g., Havward, 1988),
amd systematic revision of selected genera in
limited geographical areas (e.g., Soule & Soule,
1973) continue to emphasize the abundance and
taxonomic variety of coral reef-associated
Bryozoa, and the present inadequate level of svs-
tematic study devoted to them. The bryozoan
fauna of the Great Barrier Reef is especially rich.
It certainly exceeds several hundred species, but
the total cannot be estimated. In the first account
of the bryozoan fauna of Heron Island reef flat
(Ryland & Hayward, 1992), 81 species were de-
scribed, of which 24 were recorded for the first
time from Australian waters and 14 were new, A
further 43 species are described and illustrated in
this paper, together with three others noted or
incorrectly identified in the previous paper (Ry-

land & Hayward, 1992). Of this total, 20 are new
to science, while another five are reported for the
first time from Australia.

A general account of Heron Island reef and the
collecting sites worked were given in the first
paper, together with details af 14 of the 28 sta-
tions sampled in 1972 and the occurrence of
Sargassum sp. in 1988, Notes on the remaining
1972 stations, and on a further group of stations
worked in 1988 are given in Appendix 1. All
collecting sites are mapped in Fig. 1.

SYSTEMATICS

The taxonomic order adopted here follows that
advocated by Gordon (1984, 1986, 19892). Tax-
onomic diagnoses are presented at family and
penus level, except where these have already
been provided in the previous account (Ryland &
Hayward, 1992). Species descriptions and mea-
surements are based on Heron Island specimens.
The synonymies given for each species are delih-
erately selective, and unverified synonyms have
been excluded.

Class STENOLAEMATA Borg, 1926
Order CYCLOSTOMATIDA Busk, 1852a
Family FILISPARSIDAE Borg, 1944

DIAGNOSIS

Colony erect, branching, with flattencd
branches. Autozooids arranged in quincunx, or in
transverse rows. Gonozooid frontal, partially im-
mersed, often indistinct,

534

FIG. 1. Heron Island and reef, showing location of samples.

Nevianopora Borg, 1944

TYPE SPECIES
Idmonea milneana d'Orbigny, 1839.

DIAGNOSIS

Colony erect, branching dichotomously, with
flat basal surfaces. Autozooid orifices in regular
transverse rows, the median one slightly more
proximally situated than those on either side.
Gonozooid frontally situated at a branch dichot-
omy, irregular in outline, the ooeciostome shortly
tubular and flared.

Nevianipora pulcherrima (Kirkpatrick)
(Fig. 2A)

Idmonea pulcherrima Kirkpatrick, 18902: 22, pl. 4, fig.
6.

Tubulipora pulcherrima (Kirkpatrick) Harmer, 1915:
129, pl. 9, figs. 1-5.

Nevianopora pulcherrima (Kirkpatrick) Brood, 1976:
292, fig. 12H,L.

MATERIAL EXAMINED
Stn. 27.

MEMOIRS OF THE QUEENSLAND MUSEUM

REMARKS

A single colony was collected
from Stn. 27, encrusting Jodictyum
receptaculum sp. nov.

DISTRIBUTION

Described originally from the
Torres Straits, N. pulcherrima is
widely distributed through the
Indo-Malayan region, and west-
wards to the coasts of East Africa.
It will probably prove to be com-
mon throughout the northern GBR
province.

Family LICHENOPORIDAE
Smitt, 1867
Disporella Gray, 1848

TYPE SPECIES
Discopora hispida Fleming,
828.

DIAGNOSIS

Colony encrusting, discoidal or
cup-shaped, with a thin peripheral
lamina. Autozooids in radiating or
quincuncial series; linked by calci-
fied struts defining alveoli, ex-
trazooidal coelomic spaces. Alveoli progres-
sively reduced by centripetal calcification but not
closing completely, retaining a central foramen.
Embryos brooded in extrazooidal brood cham-
bers in the central region of the colony; embryos
released through a short, tubular, upwards-di-
rected ooeciostome.

Disporella buski (Harmer)
(Fig. 2B-D)

Lichenopora buski Harmer, 1915: 161, pl. 12, figs 4,5.
Disporella buski (Harmer) Brood, 1976; 299, figs 17D-
G.

DESCRIPTION

Colonies circular, developing a deep cup-
shape, up to 3.5mm diameter, First three or four
astogenetic generations of autozooids ordered in
radiating linear series; thereafter diverging and
adopting a regular quincuncial arrangement. Au-
tozooid tubes more or less oval in section, to
0.08mm wide; especially long in deepest part of
cup, the distal lip drawn out into two or three
slender, spinous processes. Short spines also

HERON ISLAND BRYOZOA 5

m
л

FIG. 2. A, Nevianipora pulcherrima, x34. B-D, Disporella buski. B, part of colony, with ooeciostome, x40. C,
entire colony in oblique lateral view, x14. D, entire colony viewed from above, x15.

present on the upper surface of the autozooid
tube. Brood chamber apparent as a finely perfo-
rated sheet underlying calcified struts linking the
bases of the central autozooid tubes; ooeciostome
short, round-sectioned, its aperture level with the
surface of the chamber.

REMARKS

Numerous cup-shaped colonies of this species
were present on Sargassum, sometimes closely
intergrown with encrusting cheilostomates. The
spinous peristomes and the finely perforated
brood chamber are characteristic for D. buski, as
is the short, round-sectioned ooeciostome.

536

DISTRIBUTION

Described originally from the Indo-Malayan
region, D. buski is widely distributed in the west-
ern Pacific, from New South Wales to Japan, and
has been recorded from a dozen localities on the
East African coast (Brood, 1976). It is common
in the Torres Straits, and probably throughout the
GBR province.

Family CRISIIDAE Johnston, 1838

DIAGNOSIS

Colony erect, branching, jointed; each inter-
node of one to many tubular autozooids, all fron-
tally facing, linked by chitinous, tubular nodes.
Gonozooid inflated, replacing an autozooid, with
densely punctate calcification; ooeciostome dis-
tally situated, slit-like, tubular or flared.

Crisia Lamouroux, 1812

DIAGNOSIS

Sterile internodes consist of at least three, to
more than 30 autozooids; fertile internodes (bear-
ing gonozooids) with at least five autozooids.
Basally jointed spines present in many species,

TYPE SPECIES
Sertularia eburnea Linnaeus, 1758.

Crisia elongata Milne Edwards
(Fig. 3A)

Crisia elongata Milne Edwards, 1838: 203.
Crisia elongata Milne Edwards, Harmer, 1915: 96, pl.
8, figs 1-8; Gordon, 1989b: 448, figs 1,2.

REMARKS

This widely distributed tropical crisiid is recog-
nized by its curled branches, black joints, and
broad, globular gonozooid, which has a narrow,
slit-like ooeciostome. It has been widely recorded
from reef habitats in the Indo-West Pacific re-
gion, as well as from Samoa, and East Africa.
Several colonies were found attached to Sargas-
sum leaflets.

Bicrisia sp.
(Fig. 3B)

REMARKS

Several small colonies of Bicrisia were present
in the sample of Sargassum. Unfortunately, none
bore gonozooids, without which the species'
identity cannot be established.

MEMOIRS OF THE QUEENSLAND MUSEUM

Class GYMNOLAEMATA Allman, 1856
Order CHEILOSTOMATIDA Busk, 1852a
Suborder INOVICELLINA Jullien, 1888
Superfamily AETEOIDEA Smitt, 1867
Family AETEIDAE Smitt, 1867

DIAGNOSIS

Colony encrusting, creeping; consisting of tu-
bular autozooids, each with a decumbent proxi-
mal portion, and an erect tubular portion bearing
distally the frontal membrane and operculum.
The adherent proximal portions constitute a ram-
ifying, stolon-like growth, filiform or monili-
form. The polypide occupies the whole of the
autozooid, and may be withdrawn into the en-
crusting, proximal part. The everted tentacle
sheath has a finely toothed collar. Embryos
brooded in membranous, evanescent ovisacs, at-
tached at the distal end of the autozooid.

Aetea Lamouroux, 1812

TYPE SPECIES
Sertularia anguina Linnaeus, 1758,

DIAGNOSIS
As for family.

Aetea anguina (Linnaeus)
(Fig. 3D)

Sertularia anguina Linnaeus, 1758: 816.
Aerea anguina (Linnaeus) Harmer, 1926: 194, pl.13,
figs. 3,4; Ryland & Hayward, 1977; 45, fig. 13.

REMARKS

Dense colonies of this species were abundant
on Sargassum, while more diffuse and smaller
colonies occurred in the interstices of coral rub-
ble. Aetea anguina appears to have a worldwide
distribution, except for polar seas. It is distin-
guished from other species of the genus by its
vanably flexed, spatulate distal end. The Heron
Island specimens differed from European speci-
mens in displaying variation in the length of the
closely ringed erect portion of the autozooid,
below the frontal membrane. This varied by as
much as 100% among autozooids of a single
colony. The shorter individuals, with reflexed,
spatulate distal ends could not be distinguished
from European A. anguina. Species of Aetea are
probably common in cryptic reef environments,
but their delicate colonies are usually damaged
beyond identification by collection.

HERON ISLAND BRYOZOA 53

Suborder MALACOSTEGINA Levinsen, 1909
Superfamily MEMBRANIPOROIDEA
Busk, 1854
Family MEMBRANIPORIDAE Busk, 1854

DIAGNOSIS

Colony an encrusting, unilaminar sheet; in
some species developing erect, foliaceous,
bilaminar sheets from an encrusting base. Au-
tozooids lightly calcified, with little or no gym-
nocystal calcification: the frontal surface almost
entirely membranous. A small area of cryptocys-
tal calcification present below the membrane in
some species. Tubercles typically present at the
proximo-lateral corners of autozooids, often well
developed; small marginal spinules may border
the opesia. No avicularia or ovicells. Interzooidal
communication achieved via multiporous
septula. Larva a planktotrophic cyphonautes.

Membranipora de Blainville, 1830

TYPE SPECIES
Flustra membranacea Linnacus, 1767.

DIAGNOSIS

Characters of the family. Most species are epi-
phytic. The larva gives rise to atwinned ancestru-
lar zooid.

Membranipora tuberculata (Bosc)
(Fig. 3C)

Flustra tuberculata Bose, 1802: 118.

Membranipora tuberculata (Bose) Osburn, 1950: 23,
pl. 2, figs 4-6; Mawatari, 1974: 23, fig. 2; pl. 2, figs
1-4,

REMARKS

Small colonies of this species were common on
Sargassum sp. At the growing edge the plate-like
gymnocyst of developing autozooids could be
seen enrolling on each side to form the rounded
tubercles characteristic of later ontogeny, in the
manner described and figured by Osburn (1950)
and Mawatari (1974). The narrow cryptocyst
bore a few marginal spinules in some autozooids.

DISTRIBUTION

M. tuberculata is “found wherever Sargassum
drifts over the warmer seas...." (Osburn, 1950). It
has not been previously noted from Australia, but
is certainly common throughout the western Pa-
cific,

=J

Family ELECTRIDAE Stach, 1937

Colony encrusting, forming a unilaminar sheet;
or developing erect, bilaminar lobes or sheets
from an encrusting base. Autozooids with well
developed gymnocyst, perforate or imperforate,
but only a minimal development of the
cryptocyst. Spines generally present, though
sometimes reduced: a single medio-proximal
spine, often very long, occurs in almost all spe-
cies; additional marginal spines bordering ihe
opesia variably developed. No avicularia or
ovicells. Interzooidal communication via multi-
porous septula, Larva a planktotrophic
cyphonautes, giving rise to a single ancestrular
zooid.

Electra Lamouroux, 1416

TYPE SPECIES
Flustra pilosa Linnaeus, 1767,

DIAGNOSIS
As for family.

Electra bellula (Hincks)
(Fig. 4A)

Membranipora bellula Hincks 1881, 145, pl. &, figs 4,
4a, 4b.
Electra bellula (Hincks) Cook 1985, 89.

REMARKS

Electra bellula does not seem to have been
recorded again from Australia since Hincks’
(1881) original account, although Livingstone
(1927) regarded Haswell's (1881)
Membranipora cervicornis as belonging to it,
Hincks (1881) described the nominate species
and var. mullicornis from unknown Australian
localities, and var. bicornis from Ceylon. He also
recorded the nominate species from Madagascar,
St. Vincent and the Cape Verde Islands, Hastings
(1930) described and figured Electra bellula var.
bicornis (Hincks) from the Galapagos Islands;
Cook (1985) described West African specimens
and reviewed the geographical distribution of the
species.

The autozooids are 0.4-0.5mm long, with à
smooth gymnocyst bearing medially a long,
horny, basally pointed spine, characteristic of the
genus. The oval opesia, which comprises half the
total autozooid length has two to four flat, van-
ably branched spines along its proximal edge.
which are curved over the frontal membrane,

un
a
с

forming a closely reticulate shield. There are four
or five thin, erect, reflected oral spines, and addi-
tional short, thin spines may occur elsewhere on
the gymnocyst or along the margins of the au-
tozooid. At Heron Island it occurred only on
Sargassum, although Cook (1985) reported it
encrusting hard substrata. Development of the
overarched opesial spines varies between cola-
nies and the varietal distinctions ‘bicornis’ and
‘multicornis’ are almost certainly without geo-
graphical or genetic justification.

Suborder NEOCHEILOSTOMINA
d'Hondt, 1985
Superfamily CALLOPOROIDEA
Norman, 1903
Family CALLOPORIDAE Norman, 1903

Parellisina albida (Hincks)

Membranipora albida Hincks, 1880; 81.

Parellisina albida (Hincks) Osbum, 1949: 4, fig. 8.

Parellisina curvirostris Ryland & Hayward, 1992:
229, fig. 2d.

REMARKS

This species was incorrectly identified as P.
curvirostris (Hincks) by Ryland & Hayward
(1992). P. albida is distinguished from P.
curvirostris by its avicularium, the mandible of
which has wide basal extensions, above which it
abruptly narrows. lt was common at Heron Is-
land, occurring in nine samples.

DISTRIBUTION

Presently known only from an unrecorded
Indo-Malayan locality ("Singapore or Philip-
pines" (Hincks 1880)). The P. albida described
and figured by Mawatari & Mawatari (1980)
from Japanese localities has an extremely large
avicularian chamber and the mandible lacks the
Jatero-basal lobes of Hincks’ species; it is almost
certainly distinct from P. albida (Hincks). This is
the first Australian record of the species,

Family CHAPERIIDAE Jullien, 1881

DIAGNOSIS

Colonies encrusting; or erect, foliaceous or
branching. Autozooids with variably developed
cryptocyst, opesia occupying one-fifth to two-
thirds frontal length. No gymnocyst. Occlusor
laminae. - calcified struts marking position of
opercular occlusor muscles - present within cay-

MEMOIRS OF THE QUEENSLAND MUSEUM

ity of zooid. Spines, ovicells and avicularia pres-
ent or absent,

Chaperia Jullien, 1881

TYPE SPECIES
Flustra acanthina Lamouroux, 1825.

DIAGNOSIS

Colony encrusting. Autozooids with membran-
ous frontal wall. partly underlain by a smooth or
granular cryptocyst; opesia subcircular, occupy-
ing up to two-thirds frontal length. No gym-
nocyst. Obliquely orientated occlusor laminae
present within opesia, extending from proximo-
lateral corners to distal wall of autozooid. Spmes
present bordering the lateral and distal rim of the
opesia. No avicularia. No ovicells, Multiporous
septula present in vertical walls.

Chaperia sp.
(Fig. 5A)

MATERIAL EXAMINED
À single small colony on Sargassum sp.

REMARKS

This is possibly the same species as that re-
corded from various Indo-Malayan localities as
Chaperia acanthina (Lamouroux), for example
by Harmer (1926). С. acanthina is a southern
hemisphere, cold temperate. species, distributed
from the Falkland Isles to Kerguelen and south-
em New Zealand (Gordon, 1986). It is character-
ized by a broadly oval, thick-rimmed opesia,
constituting about half the total length of the
autozooid, and a distal row of five to seven spines,
Its occlusor laminae are long, and almost parallel
to the disto-proximal axis of the autozooid,
widely spaced where they intersect the distal
wall. The Heron Island species has rather small
autozooids, commonly 0.45 x 0.3mm, with an
oval opesia occupying about two-thirds au-
tozooid length. There are 10-13 distal spines,
some borne on the rim of the opesia, and some
issuing from the distal wall. It closely resembles
the specimens figured by Harmer (1926, pl.14,
figs. 9,10) and Scholz (1991) as C. acanthina.
Chaperia judex (Kirkpatrick), recorded from
Mauritius and the Philippines (Hayward, 1988) is
another tropical species formerly confused with
C. acanthina. Jt has 15 or more stout spines
almost completely enclosing the opesia. The
Heron Island material is insufficient to give a
complete description of a new taxon; it is obvi-

HERON ISLAND BRYOZOA 539

FIG. 3. A, Crisia elongata, x50. B, Bicrisia sp., x50. C, Membranipora tuberculata, x70. D, Aetea anguina, x70.

ously different from C. acanthina, but a formal Chaperiopsis harmeri sp.nov.
description must await the discovery of further (Fig. 4B,C)
material.

Chaperia cervicornis: Harmer, 1926: 230, pl.14, fig. 8.

MATERIAL EXAMINED
HOLOTYPE: QMG304963, on Sargassum sp.

MEMOIRS OF THE QUEENSLAND MUSEUM

540

HERON ISLAND BRYOZOA

PARATYPES: QMG304968, on Sargassum sp.

DESCRIPTION

Colonies developing small, unilaminar
patches. Autozooids oval, small, commonly 0.45
x 0.3mm, with an almost circular opesia compris-
ing just less than half total length; proximal cal-
cification thin and finely granular. There are four
(rarely six) distal spines: proximal pair fairly
stout, bifurcating three or four times to develop
an open-branched antler shape, the tines cylindri-
cal; distal pair shorter and thinner, bifurcated
once or twice. At the growing edge some au-
tozooids have a third pair of very thin spines
projecting almost horizontally from the distal
wall. Occlusor laminae very short, and indistinct;
widely spread apart, passing between the bases of
the proximal and distal spine on each side.
Ovicell a shallow helmet shape, with a very large
aperture, above this is a narrow, transverse frontal
fenestra. The distal pair of spines curves along the
sides of the ovicell. Avicularia rare, shortly co-
lumnar, present on the gymnocyst of autozooids
succecding an ovicell, projecting above the
ovicell, with a short, proximally-directed mandi-
ble.

REMARKS

This is undoubtedly the species described and
figured by Harmer (1926) as C. cervicornis
(Busk). Gordon (1986) remarked that Harmer’s
specimens, from the Torres Straits, were proba-
bly distinct from C. cervicornis, which ranges
from the Bass Strait and Tasmania, south to New
Zealand, and west to the Magellan Strait, C
cervicornis has six to eight rather stout, cervicom
spines; the proximal pair are usually broadened
with the tines flexed medially and meeting above
the frontal membrane.

ETYMOLOGY
For S.F. Harmer.
Family QUADRICELLARIIDAE Gordon, 1984
Nellia Busk 18522
Nellia simplex Busk

(Fig. 4Е,Р)

Salicernaria marginata Busk, 18522: 367.

541

Nellia simplex Busk, 1852h: 19, pl. 65, fig 1; pl, 65
(bis), fig. 3.

Acanthodesia simplex (Busk) Harmer, 1926- 220,
pl.13, fig. 23.

MATERIAL EXAMINED
Stns. 25b, 28.

DESCRIPTION

Colony a dense, bushy tuli, 55mm high.
branching dichotomously at intervals of 1-4mm;
attached by a thick bundle of tubular rhizoids.
Branches square-sectioned, 0.3mm wide, coh-
sisting of four longitudinal autozooid series, in
two back-to-back pairs. Joints flexible; dichoto-
mies continuously calcified when first formed,
but fracturing across base of each new ramus, the
joints then formed from thick chitinous tubes.
Autozooids oblong, 0.5-0.65 x 0.3mm; frontal
surface largely membranous, bounded by narrow,
smoothly calcified rim of lateral wall, and with a
small, triangular area of gymnocystal calcifica-
tion on each proximo-lateral corner. Cryptocyst
smooth, imperforate, concave, underlying about
one-third total length of frontal membrane; a
similar-sized shelf of interior calcification occu-
pies the distal third of the autozooid cavity, but at
a lower, i.e., more basal, level than the cryptocyst.
Latero-basal walls meet in à pronounced suture
along the midline of the autozooid. Large, multi-
porous septula present on lateral walls. Ovicell
distinctive, wider than long, smoothly calcified
and imperforate; protruding into the cavity of the
distally succeeding autozooid, the vertical, lateral
walls of which enclose the ovicell on each side,
and extend as narrow tongues across the distal
edge of the ovicell, fusing and defining the opesia
of the distal autozooid. A vicularia absent. Small,
triangular foramina in one or both of the proximo-
lateral corners of some autozooids mark the on-
gins of the long, tubular rhizoids, which grow
basally, closely applied to the surfaces of the
branches, and gather to form the rooting bundle.

REMARKS

A single, large, profusely branched colony was
collected from Stn. 25b, and several small cola-
nies from Stn. 28. Nellia simplex was described
from the Torres Straits by Busk (1852b) and listed
by Kirkpatrick (1890b) in his report on the Torres

FIG. 4. A, Electra bellula, x 160. B,C, Chaperiopsis harmeéri; autozooids at the colony edge, x60. C, detail showing
ovicells and an avicularium, x93, D, Beania magellanica, x110. E,EF, Nellía simplex, E, autozaoids at a branch

dichotomy, x70. F, ovicelled autozooids, x40.

542

Straits collections made by A.C. Haddon in 1888-
1889. Harmer (1926) founded his account on the
fragmentary specimens described by these two
authors, and the species does not seem to have
been described subsequently.

Superfamily BUGULOIDEA Gray, 1848
Family BEANIIDAE Canu and Bassler, 1927

Beania magellanica (Busk)
(Fig. 4D)

Diachoris magellanica Busk, 1852b: 54, pl. 67, figs
1-3.

Beania magellanica (Busk) Ryland & Hayward, 1992:
235.

REMARKS

This widely distributed species was described
by Ryland & Hayward (1992) but not illustrated.
The specimen figured here is part of a large
colony from Station 23.

Beania ?plurispinosa Uttiey & Bullivant
(Fig. 5B)

Beania plurispinosa Uttley & Ваа, 1972: 28,
Gordon, 1984: 46, pl. 12, figs A-C.

MATERIAL EXAMINED
Stn. 88 ВІ.

REMARKS

A single, minute specimen of an especially
spiny Beania was collected at Stn. B1. The oper-
culum and frontal membrane are bordered by
very long, thin spines, and numerous simular
spines cover the basal walls of the autozooids.
The species is closely similar to the Kermadec
Ridge specimen figured by Gordon (1984, pl.12,
fig.B), but ihe Heron Island specimen consisted
ôf no more than 10 autozooids and a firm identi-
fication cannot be given.

Beania pectinata sp.nov.
(Fig. 5C,D)

MATERIAL EXAMINED
HOLOTYPE: QMG304961, on Sargassum sp.

DESCRIPTION

Colony diffuse, consisting of tapered, fusiform
autozooids, 0.6mm long, standing erect on an
adherent stolon, 0.05mm wide, which comprises
the proximal portions of the autozooid. Frontal

MEMOIRS OF THE QUEENSLAND MUSEUM

membrane of autozooid entirely concealed by an
almost flat shield formed from two comb-like
rows of lateral spines, touching but not fusing
along the midline of the autozooid. Operculum
terminal, as wide as long; four short, slender
processes spaced around the orifice rim: two dis-
tal and two lateral. Basal wall of autozooid
smooth; no avicularia. Each autozooid has four
basal connecting tubes: its own proximal portion,
а distal tube representing the proximal portion of
the next autozooid, and a short tube on each side,
either or both of which may develop into a lateral
autozooid. Ovicells were not present.

REMARKS

The four basal connecting tubes, comb-like
frontal shield, smooth basal wall and lack of
avicularia together distinguish this species from
all other described species of Beania,

ETYMOLOGY
Latin pectinatus, comblike.

Beania klugei Cook
(Fig. 6A,B)

Beania klugei Cook, 1968: 164, figs 2A.B.

MATERIAL EXAMINED
Stn. 29

DESCRIPTION

The colony forrns diffuse, branching, uniserial
chains of slender autozooids, creeping across the
surface of coral rubble. Autozooids are 0.8-0.9 x
about 0.25mm, more or less parallel-sided, but
strongly tapered in the proximal one-fifth; the
frontal surface is entirely membranous. Spines
are lacking, except for a pair of short, pointed
distal processes. Each zooid bears a pair of small
pedunculate avicularia attached laterally adjacent
to the operculum. New autozooids are budded
from a single disto-basal septulum, and paired
proximo-lateral septula. Ovicells were not pres-
ent.

REMARKS

Beania klugei was described from West Africa
(Cook, 1968) but shown to have a tropical/sub-
tropical distribution extending west to Panama,
and east to the Red Sea and Zanzibar. Cook
(1985) discusses the probability of its wider oc-
currence in the Indian Ocean. This is the first
record of the species for Australia, and the West
Pacific region.

HERON ISLAND BRYOZOA

Superfamily MICROPOROIDEA Gray, 1848
Family ONYCHOCHELLIDAE Jullien, 1882

DIAGNOSIS

Colony encrusting. Autozooids with an exten-
sive cryptocyst, but no gymnocyst; opesia consti-
tuting half or less of total autozooid length. Distal
edge of cryptocyst notched or indented to accom-
modate parietal and/or opercular muscles. Avi-
cularia vicarious, with a large mandible
consisting of a thick, longitudinal sclerite and
single or paired lateral laminae. No spines. Em-
bryos brooded internally, often in dimorphic fe-
male zooids.

Smittipora Jullien, 1882

ТҮРЕ SPECIES
Vincularia abyssicola Smitt, 1873.

DIAGNOSIS

Autozooid with an extensive opesia. Avicular-
ian mandible symmetrical, with two equally de-
veloped lateral laminae.

Smittipora cordiformis Harmer
(Fig. 6C)

Smittipora cordiformis Harmer, 1926; 260, pl. 16, figs
14-18,

DESCRIPTION

Colony an encrusting, unilaminar sheet. Au-
tozooids hexagonal to irregularly polygonal, con-
cave, separated by distinct sutures; 0.5-0.6 x
about 0.4mm. Frontal membrane light brown,
distinct, underlain by a thick, finely granular
cryptocyst; opesia longer than wide, 0.15 x
0.L3mm, situated in distal half of autozooid, its
proximal edge with an angular lip. Avicularia as
long as autozooids but much narrower (0.2mm),
the cryptocyst deeply concave; with an elongate
oval opesia, distinctly broadened distally; distal
end of avicularium narrowed and channelled for
the reception of the rachis.

DISTRIBUTION

The single colony collected, at Stn. 9, marks the
first record of this species from Australia. lt was
described by Harmer (1926) from two ‘Siboga’
stations in Indonesia, with additional material
from Singapore, Burma and the Amirante 15-
lands. D'Hondt (1986) recorded it from New
Caledonia.

543

Family THALAMOPORELLIDAE
Levinsen, 1909

DIAGNOSIS

Colony encrusting or erect. Frontal membrane
of autozooid entirely underlain by a granular,
concave cryptocyst; opesia usually coincident
with operculum; single or paired opesiules pres-
ent, large. No gymnocyst. Avicularia vicarious.
Ovicell large, bivalved; or absent. Vertical walls
with multiporous septula. Calcarcous spicules
present in cavity of autozooid.

Thairopora MacGillivray, 1882

TYPE SPECIES
Membranipora dispar MacGillivray, 1869.

DIAGNOSIS

Colony encrusting, unilaminar. Frontal surface
of autozooids almost entirely membranous, un-
derlain by a complete, porous cryptocyst; opesia
almost exactly coincident with operculum, with
large, paired opesiules. Opesia longer than wide.
oval or horseshoe-shaped, flanked by paired, hol-
low tubercles. Avicularia present, vicarious.
Ovicells absent.

Thairopora calcarata sp.nov.
(Figs 6D-F, 7A}

MATERIAL EXAMINED
HOLOTYPE: G304962, on Sargassum sp.
PARATYPE; G304967, on Sargassum sp.

DESCRIPTION

Colonies developing irregular, unilaminar
patches on Sargassum leaflets, adjacent lobes
occasionally fusing to form short, erect,
bilaminar plates. Autozooids more or less rectan-
gular, in regular radiating rows which bifurcate at
intervals; each new row commences with a vicar-
ious avicularium. Vertical walls of autozooid
form a thin, raised rim around the frontal mem-
brane; beneath it the cryptocyst is flat to concave,
and dips distally towards two large, irregularly
oval, unequal-sized opesiules. Cryptocyst ini-
tially thin and smooth, with about 30 minute
perforations; proximal end more distinctly nodu-
lar in later ontogeny. Opesia terminal, longer than
wide, broadly oval; distal two-thirds with a
smooth rim, terminating proximally, on each
side, with a sharp, upturned condyle; proximal
third with a broad rim of nodular, cryptocystal
calcification. On each side of the operculum the

544 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. A, Chaperia sp., x86. B, Beania ?plurispinosa, x80. C,D, Beania pectinata. C, part of colony, x50. D,
detail of a single autozooid, x150.

smooth gymnocystal calcification constituting Avicularia about half as long as autozooids; ros-
the distal opesial rim develops an erect tube, trum smoothly calcified, in the form of a rounded,
0.4mm high, which expands and divides at its tip — parallel-sided hood, projecting almost perpendic-
to form a branched, club-like process; these form ularly from frontal plane of zooids; proximal half
a close lattice over the surface of the colony. with a narrow band of cryptocyst underlying the

HERON ISLAND BRYOZOA

frontal membrane; mandible elongate oval, with
elongate sclentes converging at its rounded distal
end. Three types of spicule present: bow-shaped
compasses of two sizes (0.04mm, 0,08mm), and
very long, almost straight calipers (0.19mm).

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.55340.05;
width 0.28+0.03.

REMARKS

Thairopora calcarata is distinguished from
other species of the genus by the paired, branched
processes flanking the zooid opesiae. These inter-
lace above the colony surface forming a regular,
reticulate structure, with the meshes situated im-
mediately above the opercula, presumably allow-
ing lophophores to project through them.
Numerous colonies were found on Sargassum
leaflets; the epiphytic habit is characteristic of the
genus.

ETYMOLOGY
Latin, calcaris, a spur.

Superfamily CELLARIOIDEA Fleming, 1828
Family CELLARIIDAE Fleming, 1828

DIAGNOSIS

Colony erect, branching, jointed or unjointed,
attached by chitinous, tubular rhizoids. Branches
cylindrical or flat-sectioned, dividing dichoto-
mously, and/or producing lateral adventitious
shoots. Autozooids in alternating, longitudinal
series, in whorls around branch axis; gymnocyst
ahsent, cryptocyst extensive, opesia coincident
with operculum. Avicularia present or absent,
Ovicells immersed (endotoichal).

Cellaria Ellis and Solander, 1786

ТҮРЕ SPECIES
Farcimia sinuosa Hassall, 1840.

DIAGNOSIS

Branches cylindrical, dividing dichotomously
at regular intervals, and sometimes producing
additional, adventitious branches, Each branch
(internode) originating from a chitinous joint
(node). Autozooids typically hexagonal or dia-
mond-shaped. Opesia scarcely larger than oper-
culum; condyles usually present within straight
or convex proximal lip. Avicularia vicarious, in-
tercalated within an autozooid row and small, or
entirely replacing an autozooid, Ovicell with in-
dependent aperture distal to opesia.

545

Cellaria punctata (Busk)
(Fig. 7D)

Salicornaria punctata Busk, 18522: 366 (in part).

Salicornaria gracilis Busk, 1852b: 17, pl. 63, fig. 3; pl.
65 bis, fig. 2.

Cellaria punctata (Busk) Harmer, 1926: 337, pl. 21,
figs 14-16; text-fig. 13a.

MATERIAL
Stn. 28; one small colony attached to Pleurotoichis
clathratus.

DESCRIPTION

Internodes slender, to 2.5mm long in the pres-
ent specimen; diameter 0.25mm in non-fertile
portions, 0.35mm in fertile portions, Autozooids
arranged in alternating whorls of four, broadly
lozenge-shaped; rounded distally, pointed proxi-
mally, borders marked hy clear sutures; 0,3-0.35
x 0.2mm. Cryptocyst finely granular, deeply con-
cave between paired, curved, longitudinal ridges
which fuse with the terminal wall distal to the
opesia; extending proximally almost to the prox-
imal end of the autozooid, but not meeting. Ope-
sia about one-sixth total autozooid length;
proximal edge convex, rounded, projecting
slightly; distal edge finely denticulate; a thick,
rounded denticle present in each proximal corner.
Fertile portions of internode swollen, with three
whorls of autozenids bearing endotoichal
ovicells; aperture of ovicell transversely oval,
partially occluded by a rectangular plate. Avi-
culana sparse, as large as an autozooid; mandible
semicircular; rostrum resembling a cowl, project-
ing from the frontal plane of the internode, prox-
imal edge of opesia straight, with two deep
notches.

REMARKS

The characteristic avicularia are well figured by
Harmer (1926), who provided a synonymy for C.
punctata (Busk). Avicularia were present in the
Heron Island specimen but unfortunately were
not scanned. C. punctata was originally descnbed
from the Queensland coast; it was reported from
throughout the Indo-Malay region by Harmer
(1926), whose synonymy extended its range
westwards to $n Lanka and Zanzibar, and north-
wards to Japan.

MEMOIRS OF THE QUEENSLAND MUSEUM

546

HERON ISLAND BRYOZOA

Superfamily HIPPOTHOOIDEA Busk, 1859
Family HIPPOTHOIDAE Busk, 1859

Hippothoa flagellum Manzoni, 1870

MATERIAL
Stn. B24.

REMARKS

This inconspicuous, creeping bryozoan favours
biogenic carbonates as a substratum, Described
originally as a Pliocene fossil from Italy, it seems
to be one of the few genuinely cosmopolitan
species of Bryozoa, occurring even in Arctic and
Antarctic seas. The Heron Island colonies
showed no significant difference from European,
or indeed Antarctic, specimens.

Trypostega Levinsen, 1909

TYPE SPECIES
Lepralia venusta Norman, 1864.

DIAGNOSIS

Colony encrusting, unilaminar. Autozooids
with regularly perforated frontal shield; primary
orifice deeply sinuate, with prominent condyles,
appearing cleithridiate. Operculate heterozooids
(zooeciules) interspersed with autozooids.
Ovicell hyperstomial, prominent, regularly per-
forated, closed by autozooid operculum. Basal
pore chambers present.

Trypostega venusta (Norman)
(Fig. 7B,C)

Lepralia venusta Norman, 1864: 84, pl. 10, figs 2,3.
Trypostega venusta (Norman) Harmer, 1957: 953.

MATERIAL
Sin. B23,

REMARKS

Trypostega venusta is widely distributed in
warm temperate to tropical seas and has been
recorded from all of the oceans. It forms thin,
rather inconspicuous sheets on biogenic carbon-
ates.

547

Superfamily ARACHNOPUSIOIDEA
Jullien, 1888
Family EXECHONELLIDAE Harmer, 1957

Exechonella Duvergier, 1924

TYPE SPECIES
Cyclicopora? grandis Duvergier, 1921.

DIAGNOSIS

Colony encrusting. Autozooid frontal shield
formed by the fusion of centripetally developing
calcareous processes, with thickly rimmed or
tubaeform foramina remaining between. A tall.
imperforate peristome develops around the ori-
fice. Avicularia, ovicells and spines absent.
Multiporous septula present in vertical walls.

Exechonella ampullacea sp.nov.
(Fig. 7E)

MATERIAL EXAMINED
HoLorYPE: Stn. B27; QMG304975.
PARATYPE: Stn. B27: QMG304977.

DESCRIPTION

Colony encrusting, unilaminar; developing a
small, coherent patch, or dividing into short series
of two or three autozooid rows. Autozooids aval,
convex, separated by deep grooves, with a tall,
cylindrical, terminal peristome imparting overall
a bottle shape; 0.7-0.9 x 0.45-0.55mm. Frontal
shield. with 30-40 round foramina, each with a
thick, raised, more or less circular rim; peristome
finely nodular, imperforate, cylindrical, with à
slightly flared rim, up to 0.3mm long when un-
damaged. Primary orifice transversely oval, with-
out conspicuous condyles. No avicularia present.

REMARKS

This species differs from other described spe-
cies of Exechonella in the small size of its au-
tozooids and in the proportionately smaller, more
densely distributed foramina. It most resembles
E. tuberculata (MacGillivray) but lacks the spi-
nous frontal processes described in that species
by Harmer (1957), and is very much smaller, with
smaller frontal foramina, than the specimen
illustrated by Gordon (19894; pl.23, fig.D).

FIG. 6. A,B, Beania klugei. A, part of colony, x50, B, distal end of an autozooid, with avicularia, x240. C.
Smittipora cordiformis, x45. D-F, Thairopora calcarata. D, edge of colony, with spines in place, x45, E, cleaned
portion of colony, with two avicularia, x55. F, an avicularium in lateral view, x95.

548

ETYMOLOGY
Latin, ampullaceus, flask-like.

Superfamily LEPRALIELLOIDEA
Vigneaux, 1949

REMARKS

Gordon (19932) published the justification for
the unfortunate necessity of using this taxon for
the now familiar Umbonuloidea Canu, 1904.

Family EXOCHELLIDAE Bassler, 1935
Escharoides Milne Edwards, 1836

TYPE SPECIES
Cellepora coccinea Abildgaard, 1806.

DIAGNOSIS

Colony encrusting. Autozooids with an um-
bonuloid frontal shield bordered by large mar-
ginal pores. Orifice with a prominent, distal,
subapertural plate; bordered proximally by a
thickened peristome bearing denticles on its inner
face. Oral spines present. Avicularia adventi-
tious, typically lateral to orifice. Ovicell promi-
nent, hyperstomial, imperforate, Basal pore
chambers present.

Escharoides longirostris Dumont
(Fig. 8A)

Escharoides longirostris Dumont, 1981: 630, fig. 1A.
Escharoides longirostris, Dumont, Hayward, 1988;
293.

MATERIAL EXAMINED
Stns, 14, 15.

DESCRIPTION

Colony a small, inconspicuous patch. Au-
tozooids broadly hexagonal, convex, separated
hy deep grooves; 0.45 x 0.40mm. Frontal calcifi-
cation smooth, with scattered, round nodules;
marginal pores large, round and distinct. Orifice
with a low peristome proximally, developed me-
dially as a prominent, rounded mucro bearing on
its inner face an angular denticle; paired proximo-
lateral denticles also present on the inner face. Six
slender, distal oral spines present, bordering a
broad, smooth oral plate. Avicularia single or
paired, lateral to orifice, laterally directed; ros-
trum slender, tapered, 0.2mm long, hooked dis-
tally. Ovicell inconspicuous, recumbent on
distally succeeding autozooid and partially im-

MEMOIRS OF THE QUEENSLAND MUSEUM

mersed; its aperture is marked by an angular lip
extending between the bases of the distalmost
spine pair.

REMARKS

This species was described from the Sudanese
Red Sea coast by Dumont (1981) and subse-
quently (Hayward, 1988) recorded from Maurit-
ian reefs. Its colonies are always small, rarely
exceeding 2-3mm?, and seem restricted to cryptic
habitats among coral rubble. This is the first
occurrence of the species in Australian waters.

Family LEPRALIELLIDAE Vigneaux, 1949

REMARKS

Gordon (1993a) has demonstrated that
Celleporaria Lamouroux, 1821 and Lepraliella
Levinsen, 1917 should be accommodated in the
single family Lepraliellidae, which has taxo-
nomic priority over Celleporariidae Harmer,
1957

Celleporaria columnaris (Busk)
(Fig, 8B-D)

Celleporacolumnaris Busk, 1881: 343, 348, pl. 26, fig.
4; Busk, 1884: 194, pl. 29, fig. 11; pl. 35, fig.16.
Holoporella intermedia (Busk) Hastings, 1932: 446,
text-fig. 20

Celleporaria columnaris (Busk) Harmer, 1957: 677,
pl, 42, figs 18-21, 23; Gordon, 1993a; 335, fig.15-
a-d.

MATERIAL EXAMINED
Sins. 16, 25,

DESCRIPTION

Colony a small, multilaminar patch. Au-
tozooids oval, convex, 0.4-0.5 x 0.3-0.35mm,
with tinely nodular calcification; frontal marginal
pores few in number, widely spaced. Primary
orifice wider than long, commonly 0.1 x 014mm;
proximal edge almost straight; condyles rounded,
conspicuous, no oral spines, no peristome. Subo-
ral avicularium medially situated, perpendicular
to orificé plane; rostnim facing laterally, 0.05mm
long, the distal end rounded and coarsely toothed;
frequently, the cystid is developed as a siout,
spike-like umbo, apparently calcifying in later
ontogeny and obliterating the rostrum, which var-
ies little in size. Vicarious avicularia infrequent,
smaller than autozooids; rostrum about 0.2mm
long, slender, almost parallel-sided, its distal end
deeply cupped, with a coarsely toothed rim.

HERON ISLAND BRYOZOA

549

FIG. 7. A, Thairopora calcarata, detail to show spicules, x260. B,C, Trypostega venusta. B, primary orifice,
x450. C, group of autozooids, x90. D, Cellaria punctata, x60. E, Exechonella ampullacea, x55.

Ovicell slightly wider than long, prominent, with
coarsely nodular calcification.

REMARKS

This small species is readily distinguished by
its conspicuous condyles and its lack of a peri-
stome. Although the cystid of the suboral avicu-
larium varies in size, often being developed as a
prominent columnar umbo, the rostrum is essen-
tially monomorphic and shows little variation in
size. C. columnaris was originally described from

the Bass Strait (Busk, 1881); Harmer (1957) de-
scribed material from Indonesia, and his synon-
ymy implied a wide geographical distribution, to
Ceylon, the Seychelles and Zanzibar, and north-
wards to Japan. However, as with all species in
this difficult genus, the synonymy of C. col-
umnaris needs to be reviewed following re-exam-
ination of the specimens on which it is founded.
Hasting's (1932) specimens of "Holoporella in-
termedia" from the GBR are referable to C. col-
umnaris, and the micrographs of C. columnaris

550 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 8. A, Escharoides longirostris, x85. B-D, Celleporaria columnaris. B, primary orifice, x340. C, ovicelled
autozooids, x60. D, autozooids and an enlarged avicularium, x100. E,F, Celleporaria granulosa. E, portion of
colony, with enlarged avicularium, x22. F, detail of three autozooids, showing primary orifice, x50.

HERON ISLAND BRYOZOA

from New Caledonia published hy Gordon
(1993b) show the same species as that described

and figured here.

Celleporaria granulosa (Haswell)
(Fig. 8E,F)

Cellepora granulosa Haswell, 1881: 4Q.
Celleporaria granulosa (Haswell) Harmer, 1957: 688,
pl, 43, figs 14-19,

MATERIAL EXAMINED
Stn. B27,

DESCRIPTION

Colony multilaminar, nodular. Autozooids
large, convex, with distinct boundaries marked
by sharp sutures; 1.0-1.1 x 0.8-0.9mm. Primary
orifice slightly wider than long, 0.25 x 0.3mm;
proximal border shallowly concave, no condyles,
no oral spines. Peristomial rim low and thickened.
Suboral avicularium medially situated, perpen-
dicular to plane of orifice, facing laterally; ros-
trum oval, distally rounded, with a smooth nm.
Vicarious aviculana as large as autozooids, ros-
trum 0.8mm long, broadest across the thickened
crossbar, spatulate and deeply cupped distally,
Ovicell prominent, spherical. Frontal calcifica-
tion thick, nodular; with a single series of closely
spaced. small, marginal pores, which become
increasingly accentuated as calcification thick-
ens.

REMARKS

This distinctive species is recognized by the
large size of its autozooids, and the correspond-
ingly large primary orifice, and by its thick,
coarsely nodular calcification, The Heron Island
specimens haye been compared with the paratype
specimens described by Harmer (1957) and con-
served in the Natural History Museum, London
(BMNH reg.nos. 1883.11.29.50, 1910.6.16.6),
and there is no question that the two are identical.
C. granulosa was described from the Queensland
coast, and certainly occurs in the Torres Straits.
Thornely’s specimens of "Holoporella simplex"
from Cargados Shoal, Indian Ocean, were re-
ferredto C. granulosa by Harmer, but should now
be re-examined before the implied geographical
distribution can be accepted.

Celleporaria hesperopacifica sp.nov,
(Fig. 9A,B)

Celleporaria mamillata Gordon, 1993b: 338, fig. Loe.

551

MATERIAL EXAMINED
HOLOTYPE: QMG304974, on Sargassum sp.
PARATYPE: QMG304970, on Sargassum sp.

DESCRIPTION

Colony an encrusting, multilaminar sheet;
thickening and becoming distinctly mamillate
with increasing size. Autozooids tumid, convex,
smoothly calcified, with few, indistinct, marginal
pores; boundaries distinct àt the growing edge,
elsewhere only newly budded autozooids clear.
Primary orifice orbicular, the proximal border
deeply concave between small, rounded con-
dyles. Two long, widely spaced, distal oral spines
present in earliest ontogeny only. Peristome well
developed, deep and thick, incorporating a verti-
cally-orientated, median suboral avicularium, of
widely varying size, the cystid produced apically
as a tapered, round-sectioned umbo. Adjacent to
the orifice the peristome rim is deeply notched;
with ontogenetic thickening the inner edges of
this notch close slightly, to define a narrow chan-
nel. The avicularium may be very small,
«0.05mm, with a spine-like mucro; or much
larger, to about 0.2mm, with a thick, columnar
mucro, with basal diameter about O. lmm. In all
cases the rostrum is oval, rounded and slightly
hooded distally, with fine denticulations on the
distal rim. Similar aviculana, equally variable in
length, occur on the interstices between àu-
tozooids. Ovicell hemispherical, smoothly calci-
fied, except for occasional short spines on the rim
above the aperture. In some autozooids a second,
slender, spine-like mucro develops on the lateral
peristome rim.

REMARKS

This species is most similar to C. fusca
(Busk)(see Ryland & Hayward, 1992) but differs
in its orbicular primary orifice, with deeply can-
cave proximal rim, in its smooth calcification,
and in the range of avicularia present. The large
vicarious avicularia of С. hesperopacifica have a
broad, scaphoid, or asymmetrically tapered ros-
trum, and the crossbar is slender, lacking a colu-
mella. Those of C. fusca tend to have narrow
spatulate rostra and the crossbar bears à stout
columella. The suboral avicularia of C. fusca
show a wider range of size than those of C.
hesperopacifica, and in the largest the rostral rim
is deeply and coarsely corrugate.

ETYMOLOGY
Latin, hesperus, west; pacificus, referring to the
ocean.

552 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 9. A,B, Celleporaria hesperopacifica. A, group of autozooids and a variety of avicularia, with the broken
bases of columnar processes, x130. B, detail showing ovicells, avicularia and columnar processes, x75. C,D,
Celleporaria sicaria. C, primary orifice, spine bases and characteristic avicularia, x 130. D, portion of colony

with enlarged avicularium, x40. E, Pleurotoichus clathratus, x70. F, Calyptotheca rupicola, primary orifice,
x250.

HERON ISLAND BRYOZOA

Celleporaria sicaria sp.nov.
(Fig. 9C,D)

MATERIAL EXAMINED
HOLOTYPE: QMG304965, Stn. 25.

DESCRIPTION

Colony encrusting, multilaminar. Autozooids
broadly oval, convex, separated by distinct su-
tures; 0.6-1.0 x 0.4-0.6mm. Frontal shield calci-
fication finely nodular, with a single series of
indistinct marginal pores, and a more conspicu-
ous series of larger pores traversing the shield just
proximal to the suboral avicularium, Primary or-
ifice wider than long, 0.12 x 0.22mm, with
straight proximal edge; two to four slender, distal
oral spines present, peristome developed as a low,
projecting lip along proximal edge of orifice.
Suboral avicularium medially situated, with sub-
stantial cystid; rostrum 0.15-0.2 mm, elongate tri-
angular, acute to frontal plane, facing laterally.
Vicarious avicularia as large as autozooids, with
dagger-like rostrum, 0.5-0.7mm long, pointed
distally and with a coarsely toothed rim. Numer-
ous small adventitious avicularia present, devel-
oped along one or both of the autozooid frontal
pore series; mostly smaller than the suboral avic-
ularium, with elongate oval rostrum. Ovicells not
present.

REMARKS

A single colony lem? was collected from Stn.
25. The distinctive orifice and suboral avicular-
jum, and striking vicarious avicularium, together
distinguish this species from all others presently
assigned to Celleporaria.

ETYMOLOGY
Latin, sica, a dagger.

Superfamily EUTHYRISELLOIDEA
Bassler, 1935
Family EUTHYRISELLIDAE Bassler, 1935

DIAGNOSIS

Colony encrusting; or erect, flexible and
rooted. Autozooids with interior cryptocystidean
calcification, bounded by an extrazooidal, col-
ony-wide, cuticle, above hypostegal and ex-
trazooidal coelomic space, Avicularia present or
absent. Embryos brooded in ovicells or dimor-
phic female zooids.

„л
л
ы

Pleurotoichus Levinsen, 1909

TYPE SPECIES
Euthyris clalhrata Harmer, 1902.

DIAGNOSIS

Colony erect, flustriform: with broad, branch-
ing, unilaminar fronds. Autozooids with
cryptocystidean frontal shields consisting of ir-
regular, cuticle-bounded bars of calcite: bounded
frontally with exterior cuticular wall, above а
hypostegal coelom, Common extrazooidal coe-
lomic space along the margins of the frond, and
over the entire basal surface. Multiporous septula
present between autozooids. Embryos brooded in
dimorphic female zooids with enlarged orifices.

Pleurotoichus clathratus (Harmer)
(Fig. 9E)

Euthyriy clathratus Harmer, 1902; 266, pl. 16, figs
18-3].

Pleurotoichus clarhratus (Harmer) Levinsen, 1909;
270; Cook and Chimonides, 1981: 63, figs 8, 34,

MATERIAL EXAMINED
Numerous colonies attached 10 dead shell and coral-
line algae, Stn. 28.

DESCRIPTION

Colony erect, flustrine, unilaminar; horny and
flexible, present specimens to 110mm high.
Branches concavo-convex, almost parallel-sided,
but broadening slightly towards distal ends; di-
viding dichotomously at irregular intervals, but
also developing adventitious branches at differ-
ent levels of the colony; commonly 2-3mm wide
in youngest colonies, but up to 8mm in oldest,
basal parts of large colonies. Autozooids ar-
ranged in regular, alternating longitudinal series,
0.8-0.9 x 0.35mm, with a large, bell-shaped aper-
ture comprising one-third total length. Frontal
shield cuticular, underlain by few, irregular cal-
cified skeletal elements, also bounded by thick
cuticle. No spines or avicularia; ovicells brooded
in dimorphic autozooids.

REMARKS

The colonies are strong and flexible, presum-
ably adapted to withstand considerable water
flow and turbulence. They are richly overgrown
with epizoites, including numerous hydroids and
bryozoans. Pleurotaichus clathratus 15 widely
distributed in the Indo-West Pacific region; it has
been reported from the coasts of South Australia,

MEMOIRS OF THE QUEENSLAND MUSEUM

554

ефе
A

*

SENT

HERON ISLAND BRYOZOA 35

New South Wales and Queensland, and a spect-
men from Heron Island was described by Cook
& Chimonides (1981).

Superfamily SCHIZOPORELLOIDEA
Jullien, 1883
Family PARMULARIIDAE Canu & Bassler,
1927

Calyptotheca rupicola sp.nov-
(Figs 9F, 10A,B)

TYPE MATERIAL
HOLOTYPE: QMG304959, Stn.7
PARATYPE: QMG304966, Stn.B21

DESCRIPTION

Colony a spreading, unilaminar sheet. Au-
tozooids rectangular to irregularly polygonal,
flat, separated by thick, raised sutures. Primary
orifice broader than long; proximal border shal-
lowly concave between short, rounded condyles.
Frontal shield densely perforated, the perfora-
tions interspaced with smooth, rounded tubercles,
which thicken in later ontogeny. A curved band
of especially large tubercles extends around the
proximal and lateral borders of the orifice, con-
stituting a distinct peristomial rim. Avicularia
rather rare; when present, proximo-lateral to ori-
fice, on a slightly inflated cystid; rostrum 0.2mm
long, bluntly triangular, directed medially.
Ovicell recumbent on distally succeeding au-
tozooid, not crossed by sutures; slightly broader
than long, distinctly flattened; perforation and
tuberculation uniform with frontal shield.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.65+0.06;
width 0.41+0.06; orifice length 0.130.007;
width 0.19+0.007.

REMARKS

The shape of the primary orifice, and the orien-
tation of the adventitious avicularium in relation
to it, together distinguish this species from other
described species of Calyptotheca.

ETYMOLOGY
Latin, rupes, rock.

сл

Family CHEILOPORINIDAE Bassler, 1936

Cheiloporina campanula sp. iov.
(Fig. 10E,F)

MATERIAL EXAMINED
HOLOTYPE: QMG304978, Stn. 25.
PARATYPE: QMG304972, Stn. 25.

DESCRIPTION

Colony encrusting, forming a thick, spreading,
unilaminar sheet. Autozooids broadly hexagonal,
convex, separated by distinct sutures. Frontal
shield thick, coarsely nodular, closely and
densely perforated by small round pores, the mar-
ginal series larger and more irregular than the
rest; cuticle clearly visible above the calcified
shield, pinkish brown in dried material. Primary
orifice distinctly wider than long, proximal mar-
gin appearing straight but actually slightly up-
curved in the frontal plane; prominent, rounded,
downcurved condyles impart a bell-like outline
to the orifice. Avicularia lateral to orifice, single
or paired, missing in some autozooids; oval prox-
imally, the rostrum abruptly narrowed distal to
the crossbar, accommodating a slender, needle-
like mandible; size variable, commonly 0.3mm
long, but as small as 0. 1mm, directed distally or
medio-distally. Fertile autozooids not observed.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 1.02+0.10;
width 0.70-90.11; orifice length 0.24+0.002;
width 0.30+0.001; avicularium length
0.32--0.005.

REMARKS

In species of Cheiloparina the ovicell is re-
duced and immersed, and brooding autozooids
are characterized by dimorphic orifices, which
are shorter than those of non-reproductive au-
tozooids and typically trilobed. In C. сатрапија
all autozooids have a slightly trilobed orifice with
a distinct bell-shaped outline. No dimorphic ori-
fices could be discerned, and the morphology of
brooding autozooids remains unknown.

ETYMOLOGY
Latin, campanula, a bell.

FIG. 10. А,В, Calyprotheca rupicola. A, autozooids and an ovicell, x50. B, distal end of an autazooid, with an
avicularium, x150. C,D, Robertsonidra praecipua. C, portion of colony, with ovicells and avicularia, x40. D.
detail of a single autozooid, with avicularium, x150. E,F, Cheiloporina campanula. E, group of autozooids, x55.

F. detail of a single autozooid, with avicularium, x70.

556 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 11. A, Gigantopora pupa, x50. B,C, Stylopoma herodias. B, Group of autozooids, x60. C, a vicarious
avicularium, x100. D,E, Nimba saxatilis. D, group of autozooids, x40. E, autozooids in lateral view, to show
peristomes, x45. F, Cigclisula fruricosa, x85.

HERON ISLAND BRYOZOA

Family GIGANTOPORIDAE Bassler, 1935

Colony encrusting or erect. Autozooids with
regularly perforated frontal shield. Primary ori-
fice broadly concave proximally, below promi-
nent lateral condyles. Avicularia adventitious,
lateral to orifice, paired; extending medially
across the frontal rim of the peristome, above a
large, lower foramen, Ovicells hyperstomial or
immersed.

Gigantopora Ridley, 1881

TYPE SPECIES
Gigantopora lyncoides Ridley, 1881.

DIAGNOSIS

Colony encrusting; or erect, foliaccous or
vinculariiform. Frontal shield nodular, closely
perforated. Ovicells hyperstomial, nodular,
densely perforated. Uniporous septula present in
vertical walls.

Gigantopora pupa (Jullien)
(Fig. 11A)

Galeopsis pupa Jullien, in Jullien and Calvet, 1903; 95,
pl. 12, fig. 1.

Gigantopora pupa (Jullicn) Harmer, 1957: 880, pl. 60,
figs 3,9; Gordon, 1984: 79, pl, 26, fig. F

MATERIAL EXAMINED
Stn. 14, a single colony,

REMARKS

This distinctive species was described origi-
nally from the Tuamotu Archipelago. It has been
recorded since from the Philippine Islands and
the Torres Straits (Harmer, 1957) and from the
Kermadec Ridge (Gordon, 1984). This is its first
occurrence from the GBR,

Family SCHIZOPORELLIDAE Jullien, 1883

Robertsonidra praecipua sp.nov.
(Fig. 10C,D)

TYPE MATERIAL
HOLOTYPE: OMG304955, Sin. 5.

DESCRIPTION

Colony an encrusting, unilaminar sheet. Au-
tozooids oval to hexagonal, convex, separated by
distinct sutures. Primary orifice wider than long;
with a short, U-shaped sinus occupying about

one-half its proximal width, flanked by short,
sharply pointed condyles. Two delicate evanes-
cent spines on distal border, persisting tn
ovicelled autozooids. Frontal shield with glisten-
ing, smoothly nodular calcification typical of the
genus, bordered by a single series of large, round,
marginal pores. An adventitious avicularium
present on a minority (<10%) of autozooids, lat-
eral suboral in position, directed proximo-later-
ally; opesia triangular, rostrum strongly tapered
to a rounded distal tip; crossbar slender, com-
plete. Ovicell recumbent on distally succeeding
autozooid, about as wide as long, globular; calci-
fication more finely nodular than that of frontal
shield.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0,65+0.05;
width 0.43+0.04; orifice length 0.140.006;
width 0.1840.009; avicularium length 0.373-0.04.

REMARKS

This species is very similar to R. novella Ryland
& Hayward 1992, a colony of which was growing
adjacent to the unique holotype of А. praecipua.
Apart from an obvious difference in size, the
autozooids also differ in the size and shape of the
primary orifice, which in R. novella is almost as
long as wide (R. novella: autozooid length
0.54+0.06; width 0.38+0.03; orifice length
0.130.002; width 0.14+0.003; avicularium
length 0.23+0.04). R. praecipua lacks the suboril
umbo seen in R. novella and also the disto-later-
ally directed, lateral suboral avicularium charac-
teristic of that species, The enlarged,
proximo-laterally directed avicularium which oc-
curs in some autozooids of Ж, novella is very
much smaller than that of R. praecipua.

ETYMOLOGY

Latin, praecipuus, extraordinary.

Stylopoma herodias sp.nov,
(Fig. 11B,C)

MATERIAL EXAMINED
HOLOTYPE: QMG304953, Stn. B24
PARATYPE: QMG304954, Stn. B24.

DESCRIPTION

Colony an extensive, multilaminar, encrusting
sheet. Autozooids hexagonal to irregularly po-
lygonal, flat or slightly convex, separated by dis-
tinct sutures. Primary orifice wider than long;
sinus deep, U-shaped, occupying almost whole of
proximal border, and constituting one-third of

558

total orifice length; condyles small and rounded,

inconspicuous. No oral spines; peristome devel-
oped as a low ridge of finely granular calcifica-
tion around distal and lateral borders of orifice. A
single, lateral oral, adventitious avicularium pres-
ent; rostrum bluntly triangular, slightly acute to
frontal plane, directed disto-laterally. Frontal
shield evenly and densely perforated by large
round pores, each surrounded by a rim of thick-
ened calcification. Vicarious avicularia sporadic,
larger than autozooids, with gigantic spatulate
rostrum; lacking a palate, but with a complete,
slender crossbar, Ovicells not observed.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.57+0.08;
width 0.34+0.03; orifice length 0.100.005;
width (,123:0,005.

REMARKS

This species is most similar to S. thornelyae
Livingstone, but differs in its smaller orifice and
proportionately larger, more broadly U-shaped,
sinus. In S. thornelyae the sinus occupies less than
half the total width of the orifice and is flanked
by rather prominent knobbed condyles. Enlarged
spatulate avicularia occur in S. rhornelyae, hut are
adventitious, each developing on the frontal
shield of an autozooid.

ETYMOLOGY
Greek, herodias, a heron.

Nimba Jullien in Jullien & Calvet, 1903

TYPE SPECLES
emis praetexta Jullien in Jullien & Calvet
1903,

DIAGNOSIS

Colony encrusting, uniserial. Autozooid frontal
shield with marginal pores, Orifice sinuate, en-
closed by a tall peristome. Adventitious avicula-
ria present, or absent. Ovicell hyperstornial, not
closed by autozooid operculum.

Nimba saxatilis sp.nov.
(Fig. 11D,E)

MATERIAL EXAMINED
HOLOTYPE: QMG304951.

DESCRIPTION

Colony encrusting, the autozooids in creeping,
uniserial chains, branching at irregular intervals.
Autozootds clongate, oval to distinctly club-

MEMOIRS OF THE QUEENSLAND MUSEUM

shaped, broadest at the distal end. Primary orifice
about as wide as Jong, proximal edge broadly
concave, condyles small and knob-like, Peri-
stome well developed, forming an erect tube
completely encircling orifice, its rim thin and
evenly flared. Frontal shield convex, thick, nod-
ular; bordered by a single or double series of
small marginal pores, separated by a well marked
groove from the vertical walls, No spines or avi-
cularia. Ovicells not found.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.63+0.07;
width 0.37+0.03; orifice length 0.13+0.006;
width 0.14+0.008.

REMARKS

Despite the moderately large size of its au-
tozooids, colonies of this species were especially
inconspicuous among the epifauna of coral rub-
ble; frequently, only the deep, flared peristomes
were immediately evident. This feature, and the
nodular frontal shield, with marginal pores only,
govern the inclusion of this species in Nimba. It
is distinguished from other members of the genus
(Gordon, 1984, 19892) by its broad sinus and
almost tubular peristome.

ETYMOLOGY
Latin, saxatilis, found among rocks.

Family STOMACHETOSELLIDAE Canu and
Bassler, 1917

Cigclisula fruticosa sp.nov,
(Figs 11Р, 12A,B)

MATERIAL EXAMINED
HOLOTYPE: QMG304950, Stn. 27.
PARATYPE; QMG304979, Stn, 27,

DESCRIPTION

Colony erect, branching, with broad, lobed,
regularly dividing branches up to 8mm wide;
developing a compact, rigid, three-dimensional
structure, 70 x 50mm, 45mm high. Autozooids
hexagonal to irregularly polygonal, large, thickly
calcified, strongly convex, separated by distinct
sutures; 0.7-0.9 x 0.4-0.5mm. Frontal shield
finely granular, with large, widely spaced mar-
ginal pores, and a few large, irregularly distrib-
uted frontal pores; calcification thickens through
ontogeny, developing irregular ridges and knobs
between the pores. Primary orifice longer than
wide, proximal border almost straight, with a
short, U-shaped median sinus occupying about

HERON ISLAND BRYOZOA 559

FIG. 12. A,B, Cigclisula fruticosa. ^, ovicelled autozooid, x90. B, primary orifice, x250. C,D, Smittoidea
incucula. C, autozooids at the colony edge, x80. D, ovicelled autozooids, x70. E,F, Pleurocodonellina laciniosa.
E, primary orifice and avicularia, x180. F, ovicelled autozooids and an enlarged avicularium, x55.

one quarter its total width; large, blunt condyles
conspicuous in proximo-lateral corners; no oral
spines. A low thickened peristome developing in
early ontogeny, obscuring but not completely
hiding the orifice, its rim variously produced into
low, blunt processes. Adventitious avicularia di-
morphic: characteristically small, oval, 0.05mm
long, with finely toothed rostrum, situated lateral
to snus, acute to frontal plane and laterally di-
rected; rarely larger, 0.2mm long, with elongate
oval rostrum. Large vicarious avicularia occur
sporadically, the cystid almost as large as an
autozooid, bearing a broadly spatulate rostrum,
up to 0.4mm long, Additional small adventitious
avicularia may be present around the peristomial
rim of the autozooid, or elsewhere on the frontal
shicld. Ovicell longer than wide, convex, con-
spicuous, bordered by large marginal pores and
with a medio-frontal group of about six large,
cribrate pores.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0,78::0.06;
width 0.4240.04; orifice length 0.180.006;
width 0.15+0.005.

REMARKS

This species is most similar to C. осс/иха
(Busk), which has been accorded a wide, and
perhaps spurious, geographical distribution from
the Torres Straits to the Philippines, and west to
the Red Sea. C. früricosa is distinguished im-
mediately from C, occlusa by its elongate, nar-
rowly sinuate orifice, and massive condyles.
Ryland and Hayward (1992) recorded С.
areolata (Kirkpatrick) from Heron Island; both
С. areolata and C. cautium Hastings have been
reported previously from the GBR (Hastings,
1932).

ETYMOLOGY
Latin, fruticosa, bushy.

Family SMITTINIDAE Levinsen, L909

Smittoidea incucula sp.nov.
(Fig. 12C,D)

MATERIAL EXAMINED
HOLOTYPE: QMG304952, Stn. 16,

DESCRIPTION

Colony an encrusting, multilaminar sheet. Au-
tozooids polygonal to irregular in outline, con-
vex, separated by thin, raised sutures. Primary
onfice slightly wider than long; distal edge with

MEMOIRS OF THE QUEENSLAND MUSEUM

indistinct denticulation: proximal edge with a
conspicuous, thickened lyrula, with convex edge
and rounded corners, occupying about half its
width; condyles narrow, downcurved, with finely
toothed edges. Six distal oral spines present in
early ontogeny, obscured by the development of
athin, raised peristome. Frontal shield irregularly
nodular, with large, round marginal pores in sin-
gle or double series. Avicularium almost perpen-
dicular to lyrula, enclosed within peristome;
wedge-shaped, broadest across the pivot bar, ta-
pered distally; a thick columella present. Ovicell
as broad as long, flattened frontally, with numer-
ous irregularly sized pores; obscured by a sutured
ooecial cover.

Measurements (means and standard deviations
of 10 values, mm): autozooid length 0.520,07;
width 0,33+0.04; orifice length 0.12320.01; width
0.120.005.

REMARKS

There are relatively few tropical species of
Simittoidea, and in reef habitats the genus is usu-
ally overshadowed by a diverse fauna of
Parasmittina. species, Smittoidea incucula is
readily distinguished from other described spe-
cies by its large, anvil-shaped lyrula and
downcurved condyles, and the six distal oral
spines. Tt was not common on the Heron Island
reef flat, occurring in just two samples,

ETYMOLOGY
Latin, diminutive of incus, an anvil.

Pleurocodonellina laciniosa sp.nov.
(Fig. I2E,F)

TYPE MATERIAL
HoroTyPE: QMG304956, Stn. B30.

DESCRIPTION

Colony a broad, multilaminar sheet. Au-
tozooids hexagonal to regularly polygonal,
slightly convex, separated by distinct raised su-
tures. Primary orifice as wide as long, more or
less flush with frontal plane of autozooid; proxi-
mal edge with a short, rounded-tnangular median
projection, variably developed; condyles con-
spicuous, oval, downcurved, with finely serrated
edges revealed by SEM. Two very small distal
oral spines present in earliest ontogeny, obscured
by the development of a low peristomial rim.
Frontal shield coarsely nodular, with a single
series of large marginal pores. Avicularia lateral
suboral, single or paired (or absent), dimorphic:

HERON ISLAND BRYOZOA

narrowly elliptical, 0.1mm long, with a slender
crossbar, large palatal foramen and oval opesia,
proximo-laterally directed; less frequently, en-
larged, to 0,3mm long, parallel-sided, with
cupped, narrowly spatulate distal end and exten-
sive palatal foramen, crossbar thick, opesia oval.
Ovicell about as wide as long, flattened frontally,
with a ring of large frontal pores; initially smooth,
but developing a nodular ooecial cover.

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.55+40.07;
width 0.37+0.04; orifice length 0.1230.006:
width 0.123-0.004.

REMARKS

The open orifice, which is scarcely immersed,
and parallel-sided avicularia, with oval proximal
opesiae seem characteristic of this genus, as do
the downcurved condyles. P. clavicula Ryland &
Hayward (1992) has a very similar ovicell to P.
laciniosa, but is distinguished by its primary ori-
fice, which is deeply concave proximally, with
much narrower condyles.

Family MICROPORELLIDAE Hincks, 1877

Microporella lunifera (Haswell)
(Fig. 134)

Lepralia lunifera Haswell, 1881: 40.
Microporella lunifera (Haswell) Harmer, 1957: 965

MATERIAL EXAMINED
Stn. 25.

DESCRIPTION

Colony an encrusting, unilaminar sheet. Au-
tozooids elongate oval, convex, separated by
deep grooves; 0.5-0.7 x 0.4-0.5mm. Frontal
shield finely nodular, punctured by numerous
tiny pseudopores, and with a few slightly larger
marginal pores. Primary orifice longer than wide,
with seven oral spines; ascopore close to proxi-
mal border of orifice, separated from it by a
distance equivalent to less than the orifice length;
with a thickened, broadly oval rim and a broad,
finely toothed, reniform lumen. Avicularia
paired, lateral, situated mid-way between as-
copore and orifice, directed disto-laterally; the
rostrum is short and shovel-shaped, to accommo-
date a long, setiform mandible. Ovicell recum-
bent on distally succeeding autozooid; small,
spherical, the calcification densely nodular ex-
cept for a smooth area above the aperture; there
is a peripheral ring of large pores.

561

REMARKS

This species was introduced cursorily, and
without illustration, by Haswell (1881) in his
account of Queensland Bryozoa. Harmer (1957)
described a specimen from Holborn Island, Tor-
res Straits. This record from Heron Island is only
the third occurrence of the species; the specimen
figured here has been compared with that de-
scribed by Harmer (1957) and found to match it
exactly.

Fenestrulina epiphytica sp.nov.
(Fig. I3B,C)

MATERIAL EXAMINED
HOLOTYPE: QMG304960, on Sargassum sp.
PARATYPE: QMG304964, on Sargassum sp.

DESCRIPTION

Colonies developing small, rounded, uni-
laminar patches. Autozooids oval, small, sepa-
rated by distinct grooves; boundaries of incurved
vertical walls marked by a low ridge around the
frontal shield. Primary orifice wider than long; no
distal oral spines. Ascopore situated almost сх-
actly in the middle of the frontal shield, with a
thick, transversely oval mm; lumen cresceniic,
without denticulations. A single series of large,
round pores borders the frontal shield, and con-
tinves around the distal edge of the onfice; a
double series of pores extends between the as-
copore and tbe primary orifice. Ovicell as wide
as long. rather flat, smooth-surfaced; ectooecial
calcification limited to an indistinct ridge on the
frontal shield of the succeeding autozooid, with a
series of small, irregular pores between it and the
entooecium. The orifice of brooding autozooids
is noticeably larger than that of sterile autozooids,

Measurements (means and standard deviations
of 20 values, mm): autozooid length 0.41+0.04;
width 0.300.03; отсе length 0.090.005;
width 0.120.007.

REMARKS

This small Fenestrulina is especially
characterised by its incurved gymnocystal, lateral
walls, and the distinct rim bordering the frontal
shield. The ascopore is also unusually large in
relation to the extent of the frontal shield. It was
found only on Sargassum leaflets.

ETYMOLOGY
Greek, epi-, upon, phyton, plant.

562 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 13. A, Microporella lunifera, x70. B,C, Fenestrulina epiphytica. A, portion of a colony, x110. B, detail of
an ovicelled autozooid, x140. D-F, Celleporina bellatula. D, portion of colony, including ovicelled autozooids,
x60. E, primary orifice, x450. F, detail showing peristomes in profile, x 110.

HERON ISLAND BRYOZOA

Superfamily CELLEPOROIDEA
Johnston, 1838
Family CELLEPORIDAE Johnston, 1838

Celleporina bellatula sp.nov.
(Fig. 13D-F)

MATERIAL EXAMINED
HOLOTYPE: QMG304971, Stn. 28.
PARATYPES: QMG304969, Stn. 28.

DESCRIPTION

Colony pisiform, minute, 1-2mm diameter in
the present material. Autozooids small, fusiform,
smoothly calcified, closely packed so that the
orifice and peristome are terminal in position and
little of the frontal shield can be seen in later
ontogeny; each has about five small marginal
pores. Primary orifice slightly longer than wide,
about 0.09 x 0.085mm; proximal edge with a
short V-shaped sinus occupying about one-third
itstotal width, condyles narrow, indistinct, except
for a short peak disto-laterally. Peristome well
developed, erect, slightly flared, with an indis-
tinct notch medio-proximally, adjacent to a single
columnar avicularian cystid; rostrum 0.05mm
long, oval, terminal, acute to plane of orifice and
directed obliquely laterally. Large vicarious avi-
cularia were not found. Ovicell slightly wider
than long, not encroached upon by the peristome;
tabula large, with an almost straight proximal
edge; crossed by finely calcified, slender struts of
ectooecial calcification, fusing medially with the
underlying entooecium.

REMARKS

Numerous colonies of this tiny species were
collected from Station 28, encrusting a large tuft
of Nellia simplex. In its single, laterally directed
suboral avicularium it resembles C. rosrellata
Harmer (1957). That species, however, is
characterised by a very broad primary orifice,
with a wide sinus occupying most of its proximal
width, and a proportionately larger avicularium.
C. rostellata has numerous vicarious avicularia,
with a broad, rounded, scaphoid rostrum.

ETYMOLOGY
Latin, bellatula, diminutive of bellus, beautiful.

Celleporina cochlearia sp.nov.
(Fig. 14A,B)

MATERIAL EXAMINED
HOLOTYPE; QMG304976, Heron Ida, on Sargassum
sp.

563

DESCRIPTION

Colony a small, domed patch, 2-3mm diameter.
Autozooids small, tumid, smoothly calcified,
with few, large marginal pores. Primary orifice
terminal, 0. 10mm long, about as wide as long;
rather angular, the proximal edge concave, be-
tween indistinct condyles. Peristome perdos
from a pair of large septula flanking the orifice;
short and thick laterally, thickened and umbonate
proximally; incorporating a single median subo-
ral avicularium, with short oval rostrum, acute to
orifice plane and proximally directed. Ovicell
more or less hemispherical; tabula flat and occu-
pying most of frontal surface, with irregular slits
around its periphery. Vicarious avicularium dis-
tinctive, 0.3mm long, broadest across the slender
crossbar; proximal portion more or less semicir-
cular, rostrum long and slender, only half width
across condyles, almost parallel-sided with a
smoothly rounded tip.

REMARKS

This small Celleporina occurred only on Sar-
gassum, but was abundant in the samples col-
lected. It is especially characterized by its single,
median suboral avicularium, by the broad frontal
tabula to the ovicell, and by the long, slender
avicularium, which resembles a cook's ladle.

ETYMOLOGY
Latin, cochlear, a ladle.

Celleporina fistulata sp.nov.
(Fig. 14C-B)

MATERIAL EXAMINED
HOLOTYPE: QMG304973, Sin. 28,

DESCRIPTION

Colony pisiform, or cylindrical. Autozooids
oval and convex when newly budded; calcifica-
tion smooth, with one or two proximal marginal
pores, and a triple series of closely spaced pores
around the distal half of the autozooid: 0.4-0.45
x ¢.0.3mm. Primary orifice pear-shaped, longer
than wide, 0.12 x 0.08mm, the proximal edge
deeply sinuate between inconspicuous condyles;
no oral spines. Peristome developed as an erect
tube, 0.4-0.5mm high, completely hiding orifice,
smoothly calcified, with tubular extensions of the
autozooid pores around its basal half, A single,
tubular avicularium incorporated within the peri-
stome, its rostrum triangular, 0.06mm long, situ-
ated on the proximo-lateral rim, slightly acute to
secondary aperture, directed outwards, Other avi-

564 MEMOIRS OF THE QUEENSLAND MUSEUM

HERON ISLAND BRYOZOA

cularia not found. Ovicell wider than long, irreg-
ularly oval, situated at base of peristome; entooe-
cial tabula occupying most of frontal surface,
finely granular, bordered by thickened struts of
ectooecium.

REMARKS

This small Celleporina is especially
characterised by its pear-shaped primary orifice
and long, tubular peristome, bearing just a single
avicularium on its rim. Colonies were encrusted
on Pleurotoichus clathratus from Station 28.

ETYMOLOGY
Latin, fistula, a pipe.

Celleporina rostellata Harmer
(Fig. 14F, 15A,B)

Celleporina rastellata Harmer, 1957: 907, pl. 62, figs
18,19,

MATERIAL EXAMINED
Numerous colonies on beached Sargassum sp.

DESCRIPTION

Colonies developing small patches, nodules or
cylinders, 2-3mm long. Autozooids smoothly
calcified, closely packed; frontal shield, visible
only in marginal autozooids, with few, relatively
large, marginal pores. Primary orifice slightly
longer than wide, 0.15 x 0.12mm; proximal edge
with a deep, U-shaped sinus occupying half its
total width; condyles low and rounded. Peristome
developed in early ontogeny, characteristic: com-
pletely surrounding orifice and incorporating a
proximo-lateral, suboral avicularium, with oval,
vertically orientated rostrum, the distal rim finely
denticulate; medio-proximal rim deeply notched,
opposite proximo-lateral edge developed as a
thickened umbo. The avicularium faces laterally;
its proximal edge is produced as a sub-triangular
process projecting above the peristomial notch.
Ovicell hemispherical with a large frontal tabula,
perforated by two rows of irregular pores. Vicar-
ious avicularia frequent, varying in size, up to
0.27mm long; rostrum slightly broadened dis-
tally, and deeply cupped; crossbar complete, with
an indistinct median thickening; palate with an
extensive foramen.

565

REMARKS

The specimens described by Harmer (1957),
from the Torres Straits (Fig. 15b), had grown as
slender cylinders, probably investing hydroid
stems. They bear a close resemblance to the
Heron Island specimens. This appears to be just
the second report of the species, and its first
occurrence in the GBR province.

Family PHIDOLOPORIDAE Gabh & Hom
1862

&

Plesiocleidochasma Soule, Soule & Chaney,
199]

TYPE SPECIES
Lepralia porcellana var. Livingstone, 1926.

DIAGNOSIS

Colony encrusting, unilaminar to multilamimar.
Autozooid frontal wall with few marginal perfo-
rations; primary orifice orbicular, smooth-
rimmed, with sharp. downcurved condyles above
a broad, arcuate sinus. Avicularia adventitious,
typically latero-proximal to orifice, distally or
laterally directed. Ovicell hyperstomial, im-
perforate, with well-developed labellum, not
closed by autozooid orifice.

Piesiocleidochasma normani (Livingstone)

Lepralia porcellana var. normani Livingstone, 1926;
92, pl. 8, fig. 1.

Plesiocleidochasma normani (Livingstone) Soule.
Soule & Chaney, 1991: 474, pl. 4. fig. 4, pl. 3, fig.
3,

Schedocleidochasma porcellanum (Busk) Ryland &
Hayward, 1992: 287, fig. 27е.

REMARKS

This species was incorrectly attributed to
Schedocleidochasma porcellanum by Ryland &
Hayward (1992). It has proved to be common al
Heron Island, occurring at 21 ofthe sites sampled,

DISTRIBUTION

Plesiocleidochasma normani seems to bc
widely distributed in the western Pacific. Soule er
al. (1991) record it from Indonesia, the Solomon

FIG. 14, А,В, Celleporina cochlearia. A, primary orifice, x320. B, ovicelled antozooids with characteristic
peristome, and vicarious avicularia, x110, C-E, Celleporina fistulata. C, primary orifice, x270. D, ovicelled
anlozooids, x70, E, peristomes in lateral view, x70. Е, Celleporina rostellata, part of colony, with ovicelled

autozooid and vicarious avicularium, x90.

566 MEMOIRS OF THE QUEENSLAND MUSEUM

HERON ISLAND BRYOZOA

Islands, Fiji and Vanuatu, and eastwards to
Tonga, French Polynesia and Hawaii,

Reteporellina denticulata (Busk)
(Fig. 15C.D)

Retepora denticulata Busk, 1894: 109, pl. 26, fig. 1a-d.

Reteporellina denticulata (Busk) Harmer, 1934: 581,
pl. 35, figs 21-23, pl. 38, figs 27-32, text-figs 25D,
33,

DESCRIPTION

Colony architecture indeterminate, generally
arising from a narrow basal stalk, developing
curved, spreading or twisted lobes of narrow,
irregularly dividing branches; fusing irregularly
to give an uneven reticulate structure, or partly
non-anastomosed; may exceed 40mm height,
with equivalent spread. Trabeculae consist of five
to nine alternating. longitudinal series of au-
tozooids, each more or less hexagonal, gently
convex, separated by distinct raised sutures; com-
monly 0.5 x 0.3mm. Frontal shield of autozooid
smooth, with just two or three large pores mar-
ginally, towards its proximal end; thickening
steadily through ontogeny, with orifice becoming
deeply immersed, sutures more indistinct, hut
pores remaining visible. Primary orifice wider
than long, transversely oval, with small rounded
condyles; no oral spines. Peristome with а syn-
metrical, medio-proximal notch, flanked by
prominent, rounded Knobs; as this deepens
through ontogeny, a deep channel develops on the
inner face of the peristome. Avicularia sporadic,
suboral, obliquely transversely onentated; the
rostrum about 0.2mm long, sharply hooked api-
cally, the tip typically forked, Ovicell pyriform,
with a short frontal fissure and a narrow labellum.

DISTRIBUTION
Keteporellina denticulata is widely distributed
throughout the tropical Indo-west Pacific region.

Iodictyum mamillatum sp.nov-
(Figs 15E,F, 16A)

MATERIAL EXAMINED
HOLOTYPE: QMG304957, Stn. 27.
PARATYPE: QMG304958, Stn. 27.

567

DESCRIPTION

Colony an irregular, repent fan shape, sup.
ported two or three mm above the substratum by
columnar processes developed on its basal sur-
face; area exceeding 20 x 30mm, the edge irreg-
ularly folded but not enrolled. Colour dull white.
Fenestrulae small, oval, commonly 1.0 x 0.5mm;
trabeculae stout, consisting of three or four alter-
nating, longitudinal autozooid series, doubled at
points of trabecular fusion. Autozooids hexago-
nal, becoming irregularly polygonal in later on-
togeny, rather flat, separated by distinct raised
sutures; 0.3-0.4 x 0.15-0.2mm. Frontal shield
finely granular, with two to four, rarely more,
large and distinct pores close to its proximal
margins; typically, each autozooid bears a pair of
prominent, smoothly conical umbones proximo-
lateral to the peristome. Primary orifice orbicular,
as wide as long. the proximal border shallowlv
concave; condyles bluntly rounded, conspicuous;
distal rim with relatively large denticulations. No
nral spines. A mid-proximal pseudospiramen
forms in carly ontogeny, developing as a clearly
defined tube as the peristome forms; peristome
encircling and obscuring primary orifice, with an
orbicular secondary aperture, ils rim produced
into about ten short, blunt, spikes, but without
clear internal ridges. Frontal avicularia sporadic,
shoe-shaped, normal to frontal plane, with vary-
ing orientation; O.lmm long. Tess frequently,
slightly larger avicularia occur, 0,1-0.15mm
long, with narrow triangular rostrum, acute to
frontal plane. Larger avicularia with irregularly
spatulate rostrum, 0,2-0.25mm long, occasional,
usually borne by autozooids on the margins of the
fenestrulac. Ovicells not developed in the present
materia].

REMARKS

Two colonics only were found. Both had devel-
oped as irregular spreading fans with their frontal
surfaces more or less parallel to the substratum
surface, and attached to it by the thickened base
of the colony, and by short, stoutly calcified pro-
cesses developed from the basal surfaces.

ETYMOLOGY
Latin, mamillatus, breast-shaped,

FIG. 15. A,B, Celleporina rostellata. A, primary orifice, Heron Island specimen, x330. B, part of the holotype,
Torres Straiis, x55. C,D, Reteporellina denticulata. C, group of avtozooids with characteristic avicularia, x 120.
D, oblique view to show peristomes, x 130, E,F, Jodictyum mamillatum, E, primary orifice, x550, F, group of
autozooids with characteristic umbones and ауісиапа, x90,

568

MEMOIRS OF THE QUEENSLAND MUSEUM

Fig. 16. A, lodictyum mamillatum, autozooids at the growing edge, showing development of peristome, x130.
B-D, lodictyum receptaculum. B, ovicelled autozooids, x110, C, autozooids at a fenestrula, with a peristomial
avicularium, and a vicarious avicularium, x70. D, primary orifice, x350.

lodictyum receptaculum sp.nov.
(Fig. 16B-D)

MATERIAL EXAMINED
HOLOTYPE: QMG304949, Sin. 27.

DESCRIPTION

Colony 35mm high, with horizontal spread of
50mm, its basal portion forming a short peduncle,
the free edge lobed and folded, the lobes fused at
one or two points, resembling a flower basket.
Colour deep magenta, Fenestrulae broadly oval,
rather small, commonly 1,5 x 1.0mm; trabeculae
consisting of four alternating longitudinal series
of autozooids, doubled at points of trabecular

fusion. Autozooids hexagonal to polygonal, con-
vex, separated by distinct raised sutures; 0.4-0.45
x 0.2-0.25mm. Frontal wall smooth, with four to
six very large marginal pores. Primary orifice
pyriform, broadest distally, tapered proximally to
a deep U-shaped sinus which occupies whole of
proximal edge; condyles small and rounded, dis-
tal denticulations comparatively large, distinct.
No oral spines. Peristome projecting when newly
developed, later submerged by thickening calci-
fication and opening flush with autozooid sur-
face; with a symmetrical, semicircular notch
mid-proximally, continued within the peristome
as a defined groove; rim produced into four or
five indistinct peaks on each side, not markedly

HERON ISLAND BRYOZOA

100

% total species

5 1 15 20 25 30 35

No, of occurrences
Fig. 17. Species richness at 54 hard-substratum stations,

ribbed on its inner surface. Avicularia sporadic,
on frontal shield of autozooids, 0.1-0.15mm long,
rostrum elongate triangular, slightly acute to
frontal plane, with variable orientation, Rarely, a
suboral avicularium present, with inflated cystid
proximo-lateral to peristome and slightly curved,
slender rostrum, 0.2mm Jong, orientated trans-
versely across proximal edge of peristome.
Within the proximal edge of each fenestrula a
larger avicularium, with slender, lanceolate ros-
trum, 0.3mm long. Ovicell pyriform, with a short
median fissure and a very long, narrow labellum.

REMARKS

This species differs from most of the known
pigmented species of /odictyum in its smooth
peristome. It is most similar to /. buchneri
Harmer, but is distinguished from that species by
its deeper coloration, pvriform primary orifice,
and slender infrafenestral avicularium, Jodicryum
buchneri does not seem to develop the sporadic
suboral avicularia seen in /. receptaculum.

ETYMOLOGY
Latin, receptaculum, à vessel,

569

CONCLUSION

Bryozoan material reported here and in Ryland
& Hayward (1992) was collected at 54 sampling
stations on the reef flat at Heron Island. Addition-
ally, three species of Bugula were collected at
Blue Pools and a sample of Sargassum, probably
originating from a bed of the seaweed between
Heron and Sykes reefs, was collected from the
beach of Heron Cay. A total of 124 species has
now been described from these collections, of
which 34 were new to science and 32 were not
previously recorded for Australian seas. It is
worth restating that of the remaining 58 species
few have been adequately described or illustrated
in recent decades, and only a very small number
of the species described in previous accounts of
the GBR Bryozoa (notably Livingstone, 1927;
Hastings, 1932) have been collected again. Tax-
onomic research on Australian Bryozoa remains
slow and difficult. In part this is because thc
taxonomic diversity of Australian bryozoan fau-
nas has always been underestimated,-and any
faunistic survey will produce a significant num-
ber of undescribed species. However, another
major problem is the taxonomic confusion sur-
rounding the identity of many species. The
lengthy synonymies, and lists of specimens, as-
sembled by Harmer (1957) cannot be relied upon,
and species records by earlier authors, unsup-
ported by descriptions or figures, have to be con-
tinually reviewed, In all instances it is almost
always necessary to re-examine type and other
published materials before specimens may be
assigned to little-reported or long-unused taxa.
Taxonomic precision is greatly aided by scanning
electron microscopy, and taxonomic research on
Australian Bryozoa will become progressively
easier as the fauna is redescribed, and illustrated
with high quality SEM micrographs. Bock's
(1982) account of the South Australian fauna and
Gordon's (1984, 1986, 19893) works on the New
Zealand Bryozoa are models for this type of
study.

Some preliminary ecological conclusions may
be drawn from this survey of Heron Island reef
flat bryozoans. Most obvious is the difference
between the hard substratum fauna and that of the
single, and perhaps unrepresentative, sample uf
Sargassum, Fifteen species were found growing
on Sargassum leaflets, but only two, Aetea an-
guina and Rhynchozoon tubulosum, were also
recorded on coral rubble. The 13 species found
only on Sargassum included two known epi-
phytes, Membranipora tuberculata and Electra

570

beliula, but no less than six new species, together
with another, Chaperia sp., which could not be
confidently assigned to any described species.
The fifty-four hard substratum samples thus
yielded 111 species of Bryozoa; the most diverse
sample (Stn,25) contained 41 species, while the
mean number of species per station was 10.77.
The standard deviation (8.49) is high, most sta-
tions yielded just 1-5 species, and a simple plot
of the data suggests a non-random distribution
(Fig.17). The most abundant species was
Hippopodina feegeensis, a common reef flat spe-
cies in the Indo-West Pacific, which was present
as broad spreading sheets in 32 of the samples.
Seven other species together account for more
than half the total species occurrences: Srylopoma
duboisii (29), Parasmittina hastingsae (22),
Pleurocodonellina signata (20), Plesio-
cleidochasma normani (21), Rhynchozoon com-
pactum (20), R. xplendens (22) and R, tubulosum
(23). The first four were generally present as
spreading unilaminar or multilaminar sheets,
while the three species of Rhynchozoon formed
small patches or nodules.

ACKNOWLEDGEMENTS

We are grateful to the Nuffield Foundation for
the provision of a Travelling Fellowship to JSR
(1971/72), to the Royal Society for 4 travel grant
in 1988, and to the Australian Biological Re-
sources Survey for generous financial support.
Thanks are also due to Dr, Тап Lawn, director of
the Heron Island Research Station, and to Mary
E. Spencer Jones (Natural History Museum, Lon-
don). Dr, M.R, Fordy (SBS, University of Wales,
Swansea) is thanked for the scanning electron
micrography.

LITERATURE CITED

ABILDGAARD, P.D, 1806, [Descripsit сі tabulas].
Pp.1-36 ін Müller, O.F.: ‘Zoologica Danica, seu
Animalium Daniae ei Norvegiae rariorum ас
minus notorum descriptiones ct historia, ete’.
Edn.3, Vol.4. Hafniae.

ALLMAN, G.J, 1956, ‘A monograph of the freshwater
Polyzoa, including all the known species, hath
British and foreign. (The Ray Society: London).

BASSLER, R.S. 1935. Bryozoa. Gencrum et
Genotyporum, Index et Bihliographica. In
Quenstedt, W. (ed.), "Fossilium Catalogus 1:
Animalia’. Рт.67, 1-229. (W. Junk; ‘є Gravenb-

ape),
1936, Nomenclatorial notes on fossil and recent
Bryozoa, Journal of the Washington Academy of
Science 26: 156-162.

MEMOIRS OF THE QUEENSLAND MUSEUM

BLAINVILLE, H.M_D.DE, 1830, ‘Zoophytes’. Vol.60
in Cuvier, G.F.: "Dictionnaire des sciences
naturelles, dans lequel on traite méthodiquement
des différents étres de la nature..,.par plusicurs
professeurs du Muséum National d'Histoire
Naturelle et des autres principales écoles de Paris,"
(F.G. Levrault: Paris).

BOCK, РЕ. e Pry eri Phyl Bryozoa).
Pp. 319-394 1н Shepherd. S.A. & Thomas, LM.:

“Marine inveriebrates of Shes Australia’. Part
1, (Government Printer: Adelaide].

BORG, F. 1926. Studies on Recent Cyclostamatous
Bryozoa, Zoologiska Bidrag fran Uppsala 10:
181-507.

1944. The stenolaematous Bryozoa, Further zoolog-
ical Results of the Swedish Antarctic Expedition,
1901-1903, 3(5): 1-276.

BOSC, L.A.G. 1802. "Histoire naturelle des vers."
Vol.3 in Buffon, G.L.L.de: "Histoire naturelle de
Buffon, classée: d'après le système de
Linné....par R.R. Castel....nouvelle édition."
(Déterville; Paris).

BROOD, К, 1976, Cyclostomatous Bryozoa from the
coastal waters of East Africa. Zoologica Scripta 5:
2771-300.

BROWN, D,A, 1952, ‘The Tertiary Cheilostomatous
Polyzoa of New Zealand’, (Trustees of the British
Museum (Natural History): London).

BUSK, G. 18522. An account of the Polyzoa and Ser-
tularian Zoophytes, collected in the voyage of the
"Rattlesnake' on the coast of Australia and the
Louisiade Archipelago, etc, Appendix no.IV.
Pp.343-402. In MacGillivray, J., “Narrative of the
Voyage of H.M.S. Rattlesnake’. (T. & W. Boone;
London), Vol.1, 402pp

1852b. ‘Catalogue of marine Polyzna in the collec-
пол of the British Museum’. 1: 1-54, (Trustees of
the British Museum: London).

1854. ‘Catalogue of marine Polyzoa in the collection
of the British Museum’, 2; 55-120, (Trustees of
the British Museum; London),

1859. ʻA monograph of the fossil Polyzoa of jhe
Crag’, (Palaeontographical Society: London).

1881. Descriptive catalogue of the species of
Cellepora collected on the "Challenger" Expedi-
поп. Journal of the Linnean Society, Zoology 15:
341-356.

1884. Report on the Polyzoa collected by H.M.S.
Challenger during the years 1873-76, Part 1, The
Cheilostomata. Report on the Scientific Results
of the Voyage of H.M.S, Challenger, Zoalogy
10(30): 1-216,

CANU, F. 1904, Les Bryozoaires du Palagonien,
Echelle des Bryozoaires pours les terrains ter-
aires. Mémoires de la Société Geologique de
France, Paléontologie 12(3) 1-30.

CANU, F. & BASSLER, R.S. 1917. A synopsis of
American Early Tertiary cheilostome Bryozoa,
Bulletin of the United States National Museum 96:
1-87,

HERON ISLAND BRYOZOA

1927. Classification of the cheilostomatous
ryozoa. ings of the United States Na-

tional Museum 69(14): 1-42.
1929, Bryozoa of the Philippine Region. Bulletin
E 16 United States National Museum 100: 1-
COOK, P.L. 1968. Bryozoa (Polyzoa) from the coasts
Е tropical West Africa, Atlantide Report 10; 115-

1985, Bryozoa from Ghana: a preliminary survey,
Annales du Musée royale de l'Afrique Central
(Sciences Zoologiques) 238; 1-315.

COOK, P.L. & CHIMONIDES, Р.Ј. 1981. Morphology
and systematics of some interior-walled
cheilostome Bryozoa, Bulletin of the British Mu-
seum (Natural History), Zoology 41(2): 53-89.

DUMONT, J.P.C. 1981. A report on the cheilostomc
Bryozoa of the Sudanese Red Sea. Journal of
Natural History 15: 623-637,

DUVERGIER, J, 1921, Note sur les Bryozoaires du
Néogéne de l'Aquitaine. Actes de la Société
linnéenne de Bordeaux 72; 5-41,

1924, Deuxiéme note sur les Bryozoaires du
Néogéne de l'Aquitaine. Actes de la Société
linnéenne de Bordeaux 75: 145-190.

ELLIS, J. & SOLANDER, D.D. 1786. ‘The natural
history of many curious and uncommon
zoophytes". (Benjamin White & Son; Landon),

FLEMING, J. 1828, A history of British animals, exhib-
iting their descriptive characters and systematical
arrangement of the genera and species of quadru-
peds, birds, reptiles, fishes, Mollusca, and Radiata
of on United Kingdom. (Bell & Eradfute: Edin-

urgh).

GABB, М.М. & HORN, О.Н. 1862, The fossil Polyzoa
of the Secondary and Tertiary Formations of
North America. Journal af the Academy of Natu-
ral Sciences of Philadelphia 5(2): 111-179.

GORDON, D.P, 1984. The marine fauna of New Zea-
land; Bryozoa! Gymnolaemata from the
Kermadec Ridge. Memoirs of the New Zealand
Oceanographic Institute 91; 1-198.

1986. The marine fauna of New Zealand: Bryozoa:
Gymnolaemata (Ctenostomata and
Cheilostomata Anasca) from the western South
Island Continental Shelf and Slope. Memoirs af
rx Nw Zealand Oceanographic Institute, 95:

19892. The marine fauna of New Zealand: Bryozoa:
Gymnolaemata (Cheilostomida Ascophonna)
fram the western South Island Continental Shelf
and Slope. Memoirs of the New Zealand Ocean-
ographic Institute. 97: 1-158.

1989h. Intertidal bryozoans from coral reef-flat ub-
ble, Sa'aga, Western Samoa. New Zealand Jour-
nal of Zoology 16: 447-463.

1993a, Bryozoan frontal shields studies om wm-
bonulomorphs and impacts on classification.
Zoologica Scripta 22; 203-221,

1993b. Bryozoa: The ascophorine infraorders
Cribriomorpha, Hippothoomorpha and Um-

571

banulomorpha mainly from New Caledonian wa-
ters. In Crosnier, A. (ed.), ‘Résultats des
Campagnes Musorstom', Урі. 11, Mémoires du
Маам national d'Histoire Naturelle 158: 299-
347,

GRAY, LE. (848. ‘List of the specimens of British
animals in the collection of the British Museum’.
Part 1, Centronize, or radiated animals, (Trustees
of the British Museum: London).

HARMER, S.F. 1902. On the morphology of the
Cheilostamata, Quarterly Journal of Microscopi-
cal Science 46: 263-350.

1915, The Polyzoa of the Siboga Expedition, Pari 1,
Entoprocta, Cienostemáta and Cyclostomata.
Siboga Expedition Reports 28А: 1-180,

1926. The Polyzoa of tlie Siboga Expedition. Part 2,
Cheilostomata Anasca, Ibid, 28B: 183-501,

1934, The Polyzoa of the Siboga Expedition. Part 3,
Cheilostomata Ascophora, Family Rewporidae.
Ibid, 28C: 503-640,

1957. The Polyzoa of the Siboga Expedition. Part 4,
Picton Ascophora П. Ibid. 28D; 641-
1147.

HASSALL, А.Н. 1840. Catalogue of Inish Zoophytes,
rà and Magazine of natural History 6: 166-

HASTINGS, A,B. 1930, Cheilostomatous Polyzoa
from the vicinity of the Panama Canal collected
by Dr, С, Crossland on the cruise of the S. Y. "St.
George". Proceedings of the Zoological Society of
London 1929, No.47; 697-740,

1932. The Polyzoa, with a note on an associated
hydroid, Scientific Reports of the Great Barrier
Reef Expedition 4(12): 399-458.

HASWELL, W.A. 1881. On some Polyzoa from the
Queensland coast, Proceedings of the Linnean
Society of New South Wales 5: 33-44,

HAYWARD, PJ, 1988, Mauritian Cheilostome
Fave. Joumal of Zoology, London 215: 269-
356,

HINCKS, Т, 1877, On British Polyzoa. Part Il, Classi-
fication. Annals and Magazine af natural History
(4)20; 520-532,

1880. Contributions towards a general history of the
marine Polyzoa. П, Foreign Membraniporina.
Ibid, (5)6; 81-92, 376-381,

1881, Op. cit. IV, Foreign Membraniporina (second
series). Ibid. (5)7: 147-155.

HONDT, LL.D 1985, Contribution à la systématique
des Bryozoaires Eurvstomes. Apports récents et
nouvelles propositions. Annales des Sciences
Naturelles, Zoobogie, série 13, 7: 1-12,

1986, Bryozoaires de Nouvelle-Calédonie et du pla-
leau des Chesterfield. Bulletin du Muséum Na-
tional d'Histoire naturelle, Paris (4)8; 697-756,

JACKSON, J.B.C. & WINSTON, J.E. 1982. Ecology
of cryptic coral reef communibies. 1, Distribution
and abundance of major groups of encnishng
organisms. Journal of € tal marine Biol-
ogy and Ecology 57: [35-147,

$72

JOHNSTON, G. 1838. 'A history of the British

ytes’. (W.H. Lizars: London).

JULLIEN, J. 1881. Remarques sur quelques espèces des
Bryozoaires Cheilostomiens, Bulletin de la
Société Zoologique de France 6: 163-168.

1882, Note sur une nouvelle division des
Bryozoaires Cheilostomiens, Bulletin de la
Société Zoologique de France б: 271-285,

1883. Bryozoaires. Espéces draguées dans l'océan
Atlantique en 1881. Bulletin de la Société
Zoologique de France 7: 497-529,

1888, Bryozoaires. Mission scientifique du Cap
Hom 1832-1883, 6 (Zoologie part 3): 1-92.
JULLIEN, J. & CALVET, L. 1903. Bryozoaires pro-
venani des c s de l'Hirondelle. Résultats
des campagnes scientifiques accompli par le

prince Albert 1, 23: 1-188.

KIRKPATRICK, R. 1890. Report upon ће Hydrozoa
and Polyzoa collected by P.W. Bassett-Smith,
Esq., Surgeon R.N. during the survey of the Tizard
and Macclesfield Banks, in the China Sea, by
Н.М.5. "Rambler", Commander У.У, Moore,
АН and Magazine of natural History (6)5:
11-24.

1890b. Reports on the zoological collections made
in Torres Straits by Professor А.С. Haddon,
1888-1889, Hydroida and Polyzoa, Scientific
Proceedings of the Royal Dublin Society ns.
6(10); 603-626.

LAMOUROUX, Ј.У,Е, 1812, Extrait d'un mémojre sur
la classification des Polypiers coralligénes non
entieerement pierreux. Nouveau bulletin
емие de la Société Philosophique 3: 181-

1816, ‘Histoire des Polypiers Coralligenes Flexi-
bles, vulgairement nominés Zoophytes’, (Е, Pois-
son; Caen).

1821, ‘Exposition méthodique des genres de l'ordre
des Polypiers" (V. Agasse: Paris).

1824-26, Description des Polypiers flexibles,
Pp.603-643 in Quoy, JRC. & Gaimard, J.P.
"Voyage autour du Monde...exécutée
sur.,,l' Uranie et Ia Physicienne...'. (Pillét Aine:
Paris).

LEVINSEN, G.M.R, 1909. "Morphological and sys-
tematic studies on the cheilostomatous Bryozoa",
(Nationale Forfatteres Forlag: Copenhagen).

1917, Danmarks Expeditionen til Groenlands
nordostkyst 1906-1908. Meddelelser om Grn-
land 43 (16): 432-472.

LINNAEUS, C, 1758, 'Systema naturac per regna triae
naturae, secundum classes, ordines, genera, spe-
cies, cum characteribus, differentiis, synanymis,
locis’, 10th Edn. (Laurentii Salyii: Holmiae).

1767, ‘Systema naturae', [2th Edn, (Laurentii
Salvii: Holmiae).

LIVINGSTONE, A.A. 1926. Studies on Australian
Bryozoa, No.3. Report on the Bryozna collected
on the Great Barrier Reef, Queensland, in 1925,
by W.EJ. Paradice, Lieutenant Surgeon on

MEMOIRS OF THE QUEENSLAND MUSEUM

H.M.A.S. "Geraniam", Recoeds of the Australian
Museum 15: 79-99.

1927. Studies on Australian Bryozoa, No.5. A check
list of the marine Bryozoa of Queensland. Re-
cords of the Australian Museum 16: 50-69.

MACGILLIVRAY, P.H. 1869. Descriptions of somc
new genera and species of Australian Polyzoa; to
which is added a list of species found in Victoria.
Transactions and Proceedings of the Royal Soci-
ety of Victoria 9: 126-148.

1882. Descriptions of new, or little known, Polyzaa.
Transactions and Proceedings of the Royal Soci-
ety of Victoria 18: 115-121,

MANZONI, A. 1870. Вгуоғоі Pliocenici Italiani.
Quarta contribuzione, Sitzungsberichte der
Akademie der Wissenschaften in Wien (Abt, 1)61>
323-349,

MAWATARI, S. 1974, Studies on Japanese anascan
Bryozoa 3, Division malacostega (1). Bulletin of
the national Science Museum, Tokyo 17(1): 17-
52,

MAWATARL S. & MAWATARI, S.F. 1980, Studies
on Japanese anascan Bryozoa 5. Division
Malacostege. Bulletin of the Liberal Arts and Sci-
ences Course, School of Medicine, Nihon Univer-
sity 8; 21-114,

MILNE EDWARDS, H. 1836. Observations sur les
Polypiers fossiles du genre Eschare. Annales des
Sciences naturelles, Zoologie (2)6: 321-345,

1838, Mémoire sur les Crisies, les Hornéres et
plusieurs autres Polypes. Annales des Sciences
naturclles, Zoologie (2)9: 193-238.

NORMAN, A.M, 1864. On undescribed British
Hydrozoa, Actinozoa and Polyzoa. Annals and
Magazine of natural History (3713: 82-90,

1303, Notes on the natural History of East Finmark,
Annals and Magazine of natural History (7)11-
567-598.

ORBIGNY, A.D.’ 1835-1847. "Voyage dans
l'Amérique méridionale". (P. Bertrand: Paris; V.
Levrault: Strasbourg).

OSBURN, R.C. 1949. The genus Parellisina
{Cheilostomata Anasca, Bryózoa). Occasional pa-
pers of the Allan Hancock Foundation 10; 1-9,

1950, Bryozoa of the Pacific coast of America, Part
1, Cheilostomata Anasca, Allan Hancock Pacific
Expeditions 14(1): 1-265,

RIDLEY, S.O. 1881. Polyzoa, Coelenterata and
Sponges of Franz-Joseph Land. Annals and Mag-
azine of natural History (5)7. 442-457.

RYLAND, 1.5, & HAYWARD, P.J, 1977. British an-
ascan Bryozoans. Linnean Society Synopses of
the British Fauna n.s, 10: 1-188.

1992, Bryozoa from Heron Island, Great Barner
Reef, Memoirs of the Queensland Museum 32:
223-301.

SCHOLZ, J. 1991. Die Bryozoenfauna der
philippinischen Riffregion Cebu, Mitteilungen
Géologisch-Paláontologisches Institut der
Universität Hamburg 71: 253-403.

HERON ISLAND BRYOZOA

SMITT, F.A. 1867. Kritisk forteckning ófver
Skandinaviens Hafs-Bryozoer, Il. Ofversigt af
Kungliga Vetenskaps-Akademiens
Förhandlingar 23 (Supplement): 395-534.

1873. Floridan Bryozoa, collected by Count L.F. de
Pourtalés. Part II. Kungliga Svenska Vetenskaps-
Akademiens Handlingar 11(4): 1-83,

SOULE, D.F, & SOULE, J.D. 1973. Morphology and
speciation of Hawaiian and eastern Pacific
Smittinidae (Bryozoa, Ectoprocta). Bulletin of the
American Museum of Natural History 152: 365-
440.

SOULE, J.D., SOULE, D.F. & CHANEY, Н.Н, 1991.
New tropical Pacific and Indian Ocean
Cleidochasmatidae (Cheilostomata: Ascophora).
Bulletin de la Société des Sciences naturelles de
l'Ouest de la France, Mémoire HS 1: 465-486.

APPENDIX
List of additional collecting stations.

573

STACH, L.W. 1937. Reports of the McCoy Society for
Field Investigation and Rescarch. Lady Julia
Percy Island. 13. Bryozoa. Proceedings of the
Royal Society of Victoria n.s. 49(2): 373-384.

UTTLEY, G.H. & BULLIVANT, J.S. 1972. Biological
results of the Chatham Islands 1954 Expedition.
Part 7. Bryozoa Cheilostomata. New Zealand
Oceanographic Institute Memoir 57: 1-61.

VIGNEAUX, M. 1949. Révision des Bryozoaires
néogénes du Bassin d' Aquitaine et essai de clas-
sification. Mémoires de la Societé géologique de
France (n.s.) 28: 1-153.

WINSTON, J.E. 1986. An annotated check list of coral-
associated Bryozoa. American Museum Novitates
2859: 1-39.

1.9 Apr, 1972. Reef flat, Research Station side. thin, plate-like corals with areas of sand.

3. 10 Apr, 1972. Reef flat, Wistari Reef side. Collections from tubular Acropora, either dead or not in vigorous
growth, attached or loose but not obviously thrown up by cyclone *Emily' (Apr, 1972).

7. 11 Apr, 1972. Reef flat, western side. Bryozoans from the lower side of an Acropora, finger-like rather than

tubular.

9. 11 Apr, 1972. Reef flat, western side. Bryozoans from the underside of a detached tabular Acropora, growing
around the pedestal together with Lithothamnion, a thick red sponge, Margaretta and lace-corals,

10. 11 Apr, 1972. Reef flat, western side. Bryozoans from the underside of an attached, living Acropora.

15. 13 Apr, 1972. Bryozoans on coral rock, from a rapids area flowing through a cleft in the reef edge.

19. 14 Apr, 1972. Bryozoans from coral boulders on the reef crest.

27. 21 Apr, 1972. Bryozoans collected at about 20 feet depth along the reef edge.

28. 22 Apr, 1972. North side of reef, at 20-30 feet depth along the reef edge.

B1-B30: refer to individual boulders deposited by cyclone ‘Emily’ (Apr, 1972) between Blue Pool and the Coral
Cascades, which were sampled during the period 27 July - 2 August, 1988.

574

THE HOLOTYPE AND ADDITIONAL RECORDS OF
POGONA HENRYLAWSONI WELLS & WELLING-
TON, 1985. Memoirs of the Queensland Museum 38(2):574
1995— Two names haye been proposed for a species of
bearded dragon (Pogona) from black-soil plains, central and
western Queensland. Pogona henzylawsoni Wells & Welling-
lon 1983 was described. from à single specimen with field
sigttings and one other specimen (QMJ1051) also cited.
Description of the holotype was based on metric and color-
ation data, with limited diagnostic comparisons, Witten
(1994a) claimed that the holotype could not be found among
specimens in the Australian Museum(AM), citing a personal
communication by R. Sadlier noting the lost holotype, Witten
further claimed that the diagnosis was inadequate to differen-
иге P, henrylawsoni from the sympatric P. virticeps (АМ,
1926), nominated as neotype for P. henrylawsoni a specimen
of P. vitticeps (AMR143896), and described the "P.
henrylawsoni" аз Pogona brevis. The description of Pogona
brevis was largely of scalation, although metric characters
(short tail and limbs) were noted from Witten (1994b).

Witten's actions were invalid for two reasons. Wells &
Wellington (1985) compared P. henrylawsoni to P. vitticeps,
although one of the claimed diagnostic characters, smooth
ventral scales, was shown to be unreliable (Witten, 19942),
Wells & Wellington also noted the small size, short tail and
rounded head of their species, The holotype of P.
henrylawsoni, an ovigerous female, (SVL 130mm), (Wells &
Wellington, 1985) is smaller than the minimum mature size
for populations of P, virticeps outside Victoria (Witten &
Coventry, 1990), A rounded head and short tail Were also
reported as diagnostic for P. brevis (Witten, 19943). Secondly,
the holotype of P. henrylawsoni has been in the Australian
Museum collection since 1985, ‘The holotype (АМК 116984;
AM Field No, 16814) was registered on 6.1x, 1985, along with
other Wells & Wellington types. Witten misinterpreted à letter
from R. Sadlier, which noted two other specimens of P.
henrylawsont had not been lodged in the collection, but did
not mention the holotype (R. Sadlier, pers. comm.).

The holotype of P. henrylzwsoni (Fig. |) matches the
description of P. brevis. Wells & Wellington (1985) did not
report on scalation described by Witten ( 19942). The holotype
has 14 supralabials, 15 infralabials, 5 prenasals, 5 subnasals,
13 intemasals, 17 scales between rostral and panctal, 105
midbody scales, 17L/18R subdigital lamellae and 4L/4R lem-
oral + preanal pores. Witten (1994) reported that the tail
length given by Wells and Wellington (1985) was shorter than
im the series he examined. I confirm the short tail of the
hodotype; I measure SVLz 126mm and tail length=1 16mm.

MEMOIRS OF THE QUEENSLAND MUSEUM

Under the ICZN, existence of both a peotype and holotype
for P. henrylawsoni requires formal resolution, Application
for a ruling is in preparation (Witten, pers. comm.). P.
henrylawsoniis in common usage (Greer, 1989; de Vosjoli &
Mailloux, 1993), Thus stability is served by suppression of the
neotype, and retention of P. henrylawsoni in its onginal
application.

Details of distribution and ecology of P. henrylawsoni are few,
Five additional specimens are reported: AMR 143864, 5.8km W
Landsborough Hwy on Boulia rd, К. Sadlier & С. Shea 3.iv. 1994;
R144528, 35km by пй 5 Winton, W. van Devender 23,1,1994;
Northern Territory Museum К 11415, 19km W. Winton, Р. Horner
191.1984; R11416, 25km W. Winton, P. Horner 19.1.1984;
QMJ57178, S. of Beryl Sin, 80km E. Winton, A, Emmott
9.1ii.1993, Locality data for QMJ38760-62 (Witten, 19943) are
incomplete. These records are from $1.8km N Muttaburra on
Hughenden rd, 54.2km S Hughenden on Muttaburra rd and 21 Акт
N Hughenden on Hann Hwy respectively. Distribution of this
species thus extends from Downs to Lo h and
Aramac. AMR144528 (SVL= 120mm) and QMIS57178 (SVL=
117mm) are ovigerous females with 5 and 9 eggs, Ovipgerous
females have previously been reported in January (Wells & Wel-
lington, 1985), northern. hemisphere captives have oviposited
mostly between March and June (de Vosjoli & Mailloux, 1993),
AMER 143864 regurgitated by the elapid Pseudechis collent; is the
first record of predation for the species.

Literature Cited

de Vosjoli, P. & Mailloux, R. 1993. The general care and
maintenance of bearded dragons. (Advanced Vivarium
Systems; Lakeside USA).

Greer, А.Е. 1989, The biology and evolution of Australian
lizards, (Surrey Beatty & Sons: Chipping Norton).

Wells, R.W. & Wellington, C.R. 1985. A classification of the
Amphibia and Reptilia of Australia. Australian Journal
of Herpetology (Suppl. Ser.) (1): 1-61.

Witten, G.J. & Coventry, A.J. 1990, Small Pogona viniceps
(Reptilia: Agamidae) from the Big Desert, Victoria,
with notes on other Pogona populations, Proceedings
of the Royal Society of Victoria 102(2): 117-120.

Witten, G.J, 1994a. Taxonomy of Pogona (Reptilia:
Lacertilia: Agamidae). Memoirs of the Queensland
Museum 37(1); 329-343.

19945, Relative growth in Pogona (Reptile
Lacertilia;Agamidae). Memoirs of the Queensland Ms-
seum 37(1): 345-356,

Glenn М. Shea, Department of Veterinary Anatomy, Univers
airy af Sydney, NSW 2006,

FIG. |. Holotype of Hugona henzylawroni,

NEW SPECIES AND A NEW GENUS OF EARTHWORMS IN THE COLLECTIONS OF

THE QUEENSLAND MUSEUM (MEGASCOLECIDAE: OLIGOCHAETA)
B.G.M. JAMIBSON

Jamieson, B.G.M. 1995 1201: New species and a new genus of earthworms in the collections
of the Queensland Museum (Megascolecidae: Oligochaeta). Memoirs of the Queensiand
Museum 38(2): 515-596. Brisbane, ISSN 0079-8835.

The present study adds to the endemic earthworm fauna of Australia (Megascolecidae), one
species of the subfamily Acanthodrilinae (Rhododrilus glandifera sp. nov.) and nine species
of the subfamily Megascolecinae, in the tribes Perionychini (Heteroporodrilus
montiserralae, Terrisswalkerius windsori, Cryptodrilus bunyaensis spp, nov.);
Dichogastrini (Digaster lingi, D. moretonensis spp. nov.); and Megascolecini (Oreoscolex
retrocystis sp. nov., Propheretima eungella gen. et sp. nov., Propheretima hugalli sp, nov.,
and Spenceriella conondalei sp, nov.). Rhododrilus glandifera is the second species of this
primarily New Zealand genus to be described from Australia (both from Queensland). lt is
possible that it has acquired the microscolecin arrangement of male pores (prostates ane pair
discharging with the vasa deferentia in XVII) from à precursor with the acanthodrilin
arrangement (prostate pores 2 pairs, in XVII and XIX; male pores in XVIII) independently
of the New Zealand species. Type localities of Heteroporodrilus montiserratae, Ter-
risswalkerius windsori, Digaster lingi, D. moretonensis, Oreoscolex retrocystis and
Spenceriella conondalei are well within the ranges of their respective genera, However, the
insular locality of D. moretonensis, on Moreton Island, is of zoogeographic interest and
Cryptodrilus bunyaensis represents a northern extension for its genus, The new genus
Propheretima, has been erected for species which resemble the Oriental and Australo-Papuan
Pheretima assemblage of genera in having setae between the male pores but which are
жер рг in retaining the gizzard in segment V, not in the apomorphic location of VIII
seen in all pheretimas. It endorses the view that the ancestry of the Pheretima assemblage
was Australian. [ ]Rhododrilus, Heteroporodrilus, Terrisswalkerius, Cryptodrilus, Digas-
ter, Oreoscolex, Propheretima, Spenceriella, Megascolecidae.

B.G.M. Jamieson, Zoology Department, University of Queensland, Brisbane, Queensland

4072, Australia; 1 September 1995,

Revisionary work on megascolecid earth-
worms of Australia has led to recognition of ten
new species in collections newly and formerly
lodged іп the collections of the Queensland Mu-
seum. Three of these species, in the genera
Rhododrilus, Terrisswalkerius, Digaster and a
new genus of the Megascolecini were fixed and
preserved in ethanol by Mr. Keith McDonald, E.
Ling and A, Hugall for cladistic studies on mito-
chondrial DNA which are in progress. A further
species assignable to the new genus was collected
by Dr. G. Dyne and Mr. A. Postle and an addi-
tional species of Digaster, and a species of
Spenceriella and of Heteroporodrilus were
lodged in the Museum collections by Dr, R.
Raven and à museum group. Two species, in the
genera Oreoscolex and Cryptodrilus, come from
the author's collection. The type localities for the
ten species range from the north of Sydney, New
South Wales, to the Palmerston National Park on
the Atherton Tableland in north Queensland.

Subfamily ACANTHODRILINAE
Rhododrilus glandifera sp. nov.
(Figs 1; 2A-C; 3A-F)

MATERIAL EXAMINED

HoLorTYrE: QMG211465 (Undissected).

PARATYPE! QMG211478 (P1); QMG211511 (P2; in-
cludes slide of anterior end). All specimens from
17°35'5,145°45'Е., altitude 400-440m, Palmerston
National Park, along track to Nandrooya Falls, Qld.
riam K. McDonald, J. Ledger, D, Ledger, Sep.

DESCRIPTION

Length 58mm, width, midclitellum, 3.2mm
(Paratype 2), Segments 154 (Holotype, posterior
amputees}, 195 (P2). Colour pigmentless in etha-
ñol, Prostomium not determinable, First dorsal
pore 9/10 (Н). Setae Iumbricin, closely paired;
the pairs widely separated; in XXVIII:
ag.ab:be;cd:idd =4.7:1:4.7:1.4:17; or
13,0:2,8:13:3.7:47,596. Clitellum in XI-XVII,
strongly tumid, saddle-shaped with ventral mar-
gins shortly lateral of b lines. A pair of combined

prostamium

mala
porophore

imm

FIG. |. Rhododrilus glandifera sp. nov., Holotype,
QMG211465. Forebody and clitellar region.

male and prostatic pores in XVII, on indistinct
oval porophores, each pore directly behind two
strongly protuberant penial setae; a pair of tubular
prostates visible through the body wall, winding
from XVII to XXIX. Genital markings: a pair of
distinct papillae in intersegmental furrow 11/12,
in be (holotype, paratypes aclitellate, genital
markings undeveloped though penial setae protu-
berant to exterior), Female pores not recogniz-
able. Spermathecal pores a single pair, in
intersegmental furrow 8/9, on conspicuous papil-
lae which extend into VIII and IX (H).

Dorsal blood vessel single, continuous onto the
pharynx. Last hearts in XIII. Gizzard large.
strongly muscular, in V; preceded by a wide

MEMOIRS OF THE QUEENSLAND MUSEUM

proventriculus which is not constricted off from
it. Calciferous glands: in each of the six segments
X-XV there is a conspicuous pair of large, white,
nacreous elongate sacs which curve upwards
from the ventrolateral aspect of the oesophagus
which they invest on each side to its dorsal aspect:
in a cleared preparation of the anterior region of
paratype 2, each is seen to have longitudinal
plications in its proximal half; the glands are not
effervescent in acid alcohol. Intestinal origin XVI
(P1). Typhlosole not observable. Nephridia
avesiculate holonephridia. Iridescent sperm fun-
nels in X and XI; racemose seminal vesicles in IX
and XII. Ovaries, with large oocytes, in XIII,
Prostates a single pair, long tortuous and slen-
derly tubular; each with a slender, poorly demar-
cated duct. Penial setae present, two (H) to four
(P2) projecting at each male porophore; the setae
strongly curved, through at least 90°; with a
pointed tip; the ectal region of the shaft with
numerous palmate ectally directed scales, closely
applied to the surface; length of a seta 1.9mm,
measured around the curvature, or 1.3mm in а
straight line from base to tip (P1). Spermathecae
1 pair, discharging anteriorly in IX; with an ovoid
or somewhat clubbed ampulla and a broadly dig-
itiform diverticulum almost as long (left) or about

immature
penial seta

3 nacreous gland

A Sá spermathecal
spermathecal diverticulum
duct

spermathecal ^X:
ampulla p"

FIG. 2. Rhododrilus glandifera sp, nov., Paratype,
QMG211478. A, nacreous gland of segment XV. B,
right spermatheca; C, right prostate gland.

NEW EARTHWORMS 571

WOO ani

FIG. 3. Rhododrilus glandifera sp. nov., Paratype, QMG211478. Scanning electron micrographs of penial setae.
A, a whole seta. B, ectal end. C, ectal tip. D,E, scale-like sculpturing of ectal region. F, detail of a scale.

half the length (right), converging at the pore ETYMOLOGY
without intervention of an appreciable duct (РІ). For the highly developed nacreous glands.

FIG. 4. Heteroporodrilus montiserratae sp. nov., Ho-
lotype, QMG211443, ventral view of genital field,

REMARKS

Rhododrilus was erected by Beddard (1889)
for a New Zealand species, R. minutus. The genus
was then, as it is now, definable by the combina-
tion of the microscolecin arrangement of male
pores (a single pair of prostates discharging with
the vasa deferentia on segment ХУП) with a well
developed gizzard. Later workers (Michaelsen,
1916; Lee, 1959; Dyne, 1980) raised the generic
total to 31 species, all but one, R. gueenslandicus
Michaelsen, 1916. from New Zealand and
neighbouring islands, The new species here de-
scribed is therefore the second endemic species
of Rhododrilus to be described from Australia,
and from Queensland.

MEMOIRS OF THE QUEENSLAND MUSEUM

Rhododrilus glandifera differs from R.
queenslandicus Michaelsen, 1916, in the follow-
ing respects: the spermathecal pores are in inter-
segmental furrow 8/9, not in 7/8; and dorsal
pores, accessory genital markings and nacreous
(calciferous) glands are present.

Rhododrilus kermadecensis, a euryhaline spe-
cies described from Tasmania by Jamieson
(1974) as R. littoralis, differs notably from К.
glandifera in having 2 pairs of spermathecal
pores, in 7/8 and 8/9; in the different distribution
of genital markings and in lacking nacreous
glands; and is intermediate in the weak develop-
ment of the dorsal pores.

Subfamily MEGASCOLECINAE
Tribe PERIONYCHINI
Heteroporodrilus montiserratae sp. nov.
(Figs 4, 5, 6)

MATERIAL EXAMINED

HoLoTvPE: QMG211443 (EX G10311).

PARATYPE: QMG211444 (EX С10311). All speci-
mens from 27°57°5, 152°21'E,, Mistake Mtns, N. of
Cunningham's Gap, Qld., under rocks in complex
notophyll vine forest. Collected R. Raven, 6 Oct. 1976.

DESCRIPTION

Length 76mm in the paratype (P); the holotype
(Н) is a posterior amputee. Width 5.0 (P), 6.4mm
(H), segments 217 (P). Pigmentless in ethanol.
Prostomium tanylobous, with a transverse furrow
at midlength; neither it nor the body canaliculate;
peristomium short. First dorsal pore 5/6. Setae

aaab:be:cd:dd = 2.4;1:2.0;1,7:4.6; ог
14.5:6.1:12:10.6:27.996 (Н); in regular longitudi-
nal rows throughout. Nephropores visible as
mostly conspicuous dimpling of the interseg-
ments, in the holotype in d lines in II-IV: in c lines
in V and VI, in c lines (left) and d lines (right) in
VII, in VII-X alternating from d to c (left) or the
reverse (right); in XI and posteriorly, alternation
is regular between d and b, though out of phase
on the two sides. Clitellum annular, embracing
the posterior 2/3 of XIII-XVII, and dorsally the
anterior 1/4 of XVIII; strongly tumid in XIV-
XVII. Combined male and prostatic pores each a
small dark point at the site of the absent setae b
of ХУШ, on indistinct round papillae which fill
most of the posterior two thirds of the segment
(Н); the male field is damaged and not clearly
discernible in the paratype. Anterior genital
markings in X and XI (H, P); consisting of a low,
midventral mostly presetal papilla with pore-like
centre in X, and a transverse midventral pad in
the setal arc of XI, bearing a pair of pore-like

NEW EARTHWORMS

va : <]
Ec ra

X genital

XI markings

хіі SS 0,

XII — secu

XIV - female
pores

FIG. 5. Heteroporodrilus montiserratae sp. nov., Para-
rype 1, QMG211444, showing genital markings dif-
fering from those of the holotype though in the same
segmental location. Scale as Fig.4.

markings near its lateral borders and thrce minute
indistinct ‘pores’ near its posterior border (Н); or
(P) a pair of transversely elliptical papillae in ab
of X and XI but less far apart in XI, each pair
almost filling its segment longitudinally and with
a pore-like centre or (left posterior papilla) with
two smaller pore-like markings. Posterior genital
markings (Н) a narrow, slightly tumid transverse
band in the setal arc median to each male papillae,
each band with a row of four elliptical pore-like
markings, in XVIII; in XIX a tumid area on each
side which includes the seta Б of its side but does
not reach the anterior and posterior borders of the
segment (H). Female pores a pair of minute aper-
tures in a transverse furrow midway between the
setal arc and the anterior border of XTV, almost
1/2aa, 1.0mm, apart. S athecal pores 3 pairs,
not externally visible, ioni internal examination
in 6/7, 7/8 and 8/9, the last pair in ab.

Septa 8/9-10/11 the strongest, greatly thick-

. Dorsal blood vessel single, continuous onto

the pharynx. Last hearts in XIII, those in X-XIII
latero-oesophageal. Gizzard very large, in V, en-
sheathed by the very thin septum 5/6 and deflect-
ing several septa posteriorly. Calciferous glands
4 pairs, in Х-ХШ, sessile ventrolaterally on the
oesophagus and almost meeting midventrally. In-
testine commencing with abrupt expansion in
XV; with no appreciable typhlosole (Н, Р),
Nephridia stomate vesiculate holonephridia; pre-
lesticular nephridia have wide convoluted ducts
cach with a terminal swelling constituting à small
but definite bladder; by XII the bladder, laterally

579

Situated, is very large and depressed sub-
spheroidal; in the antenor intestinal region, e.g.
XVII, the bladder is slightly smaller and is trans-
versely elliptical; further posteriorly (XXXV) the
bladder is drawn oul medianly in the direction of
its supplying duct and projects laterally beyond
its excretory pore so that a short broad diverticu-
lum may be considered to exist. Holandric; con-
voluted sperm funnels, lacking spermatozoal
iridescence, in X and XI. Seminal vesicles in TX
and XII; those in XII more strongly multiloculate,
including a greater development of minute bead-
like loculi than those in IX. Ovaries a few short
egg strings, in XIII. Prostates racemose and
deeply incised; in the holotype with some appear-
ance of a compressed S-shape; limited to ХУШ;
duc! long and muscular, although only a small
fraction of the length of the gland; twisted in à
loop; entering the body wall in ab but nearer b (H,
P). Spermathecae 3 pairs, in VII, VIII and IX;
each with a large thin walled arnpulla, which is
widest distal to its equator and is circumferenti-
ally pleated, and a very small, narrow muscular
duct which is joined at its junction with the am-

vnlt

ША
\ М

prostate duct

prostate
gland "S
A 1mm
spermathecal spermathecal
duct diverticulum

spermathecal
ampulla

B 1mm

FIG. 6. Heteroporodrilux montiserratae sp. nov., Holo-
type, QMG211443. A, right prostate gland, B, night
spermatheca of segment IX.

pulla by a digitiform uniloculate diverticulum
containing many iridescent sperm masses (H, P).

ETYMOLOGY
Refers to the type locality Mistake Mountains.

REMARKS

Heteroporodrilus montiserratae is distin-
guished from other species of the genus in pos-
sessing four pairs of calciferous glands and three
pairs of spermathecae.

Terrisswalkerius Jamieson, 1994

The key to species given by Jamieson (1994) is
emended below to accommodate the new species,
T. windsori, and to acknowledge the presence of
five pairs of spermathecae and their pores in T.
millaamillaa which has been established in the
present study in a re-examination of the holotype.

KEY TO THE SPECIES OF
TERRISSWALKERIUS

1, Spermathecal pores 5 pairs, in or shortly behind
intersegmental furrows 4/5-8/9. . ...... . 2
Spermathecal pores 1 to 4 pairs (rarely unpaired,
midventral), in or shortly behind some or all of
intersegmental furrows 4/5-B/9 . ........ 5

Spermathecal pores 5 pairs

2. Spermathecal pores in a lines, Last hearts in XIII.
(Spermathecal duct very short.) . . . T. grandis
Spermathecal pores median to a lines. Last hearts

in XII. (Spermathecal duct at least one fourth the
length of the ampulla)

3. Nephridial bladders absent... ......... 4
Nephridial bladders present

4. Male pores well median of a lines. Moderatel
small worms «3mm wide - T. millaamillaa

Male pores in setal lines a or ab. Fairly large
worms >6mm wide. ....... T. phalacrus

5, Spermathecal pores in 4 intersegments, 5/6-8/9 or
4/5-7/8, paired or single
Spermathecal pores 3 pairs or fewer ...... 9

Spermathecal pores in 4 intersegments

6, Spermathecal pores in 5/6-8/9, unpaired, midvent-
ne T. macdonaldi
Spermathecal Pores 4 РА in 4/5-7/8 or 5/6-8/9

к о4о 46, n mE у 11667

7. Spermathecal pores 4 pairs, in 4/5-7/8.. 2...
sor oss T. ferrareginae
Spermathecal pores 4pairs, m 5/6-8/9 ,., . 8

MEMOIRS OF THE QUEENSLAND MUSEUM

8. Prostomium tanylobous, with wide dorsal tongue.
(Peristomium short). Nephropores in a straight
series on each side T. erici

Prostomium epilobous or proepilobous; with lon-
gitüdinal grooves which continue to the hind
margin of the peristomium. Nephropores in an ir-
regularly sinuous series, varying from far dor-
sally to far veritrally, on eac ach side

\ я о = + жу жү» у» ү »

9. Spermathecal pores 3 pairs, in or in or shortly be-
hind 4/5-6/7, 5/6-7/8, or 6/7-8/9 10
Spermathecal pores | or 2 pairs, in some. of 4/5-
MO sos узр Ура re 16

б, ж

Spermathecal pores 3 pairs

10. Spermathecal pores 3 pairs, in 4/5-6/7 . .. . 11
Spermathecal pores 3 pairs, in or shortly behind
5/6-7/8 or 6/7-8/9 , ... „а ‚13

11. Spermathecal diverticulum long and tortuous.
Spermathecal pores in the vicinity of d to e lines

Spermathecal diverticulum short and clavate
Spermathecal pores in or between a and b tines
ified ein eies T. montislewisi (part)
2. Spermathecal diverticulum longer than ampulla
plus duct, (Spermathecal pores in d lines)
Pian. he Pee Bs Т. barronensis
Spermathecal diverticulum shorter than ampulla
plus duct. (Spermathecal pores in d-e lines)
Tc iE ES SACRE YE T. raveni
13. Spermathecal pores immediately behind 5/6-7/8
“Ж-ге a i asa ri als T. covacevichae
Spermathecal poresin 6/7-8/9 ......., 14
14. Male pores very close together, almost contigu-
ous midventrally, on a common field or papilla
T. athertonensis
Male pores well separated, between sctal lines b
and f, on a pair оЁрарі ае... ....... 15
15, Spermathecal diverticulum simple, shortly cla-
vate T. canaliculaius
Spermathecal diverticulum composite, consist-
ing of as many as 4 parallel conjoined tubes
with terminal, knoblike seminal chambers; ap-
proximately as long as the sperrnatheca
i уур x руй een m) d 9 T. oculalus
16. Spermathecal pores 2 pairs, in 4/5 and 5/6 or 5/6
and 6/7
Spermathecal pores 1 pair, in 6/7 or 7/8 or 8/9
"d eT Ie mede re yg RISE 18

wo om o9 m t n] 0] 0 8 8 39 n 5 g n 3 on

m IU

Spermathecal pores 2 pairs

17. Spermathecal pores 2 pairs, in 5/6 and 6/7 ,
(^p EA. a ere pep ee «s. T. montislewisi (part)
Spermathecal pores 2 pairs, in 4/5 and 5/6, . ,
а isa dae i-i Pam reni ә PS i wet T. windsori

NEW EARTHWORMS

Spermathecal pores 1 pair

18, Spermathecal pores 1 pair, in or shortly posterior
to 8/9, Spermathecal diverticulum long and tortu-
ous

Spermathecal pores | pair, in 6/7 or 7/8. . , 20

19. Spermathecal pores іп setal linesc tod ....

аднаго [бэ ч эз ho LOTH eT Т. nashi
Spermathecal pores well median of setal lines а

eae GPE ae dum LIE T. liber

20. Spermathecal pores 1 раігіп 6/7 . . . T. crateris

Spermathecal pores in 7/8 . . .... T, blounti

Terrisswalkerius windsori sp. nov.
(Figs 7. 8)

MATERIAL EXAMINED

HoLorvPe: QMG211466 (Clitellate, posterior ampu-
tec).

PARATYPES: QMG211457 (PI, aclitellate, intact; P2-4,
clitellate posterior amputees). All specimens from
16°15°S, 145"02" E, Windsor Tableland, hut site alti-
tude 1060-1080m, Qld. Collected K. McDonald, 27
Mar 1993.

DESCRIPTION

Length 155mm. Width (midclitellar) 7mm,
Segments 266 (Pl). Pigmentless in alcohol. Pro-
stomium pro-epilobous, strongly canaliculate.
Peristomium very short, bisected dorsally and
ventrally. Body weakly canaliculate dorsally.
First dorsal pore 5/6 (H, Pl). Setae minute and
difficult to discern; dorsal interval (zz) narrow
and similar to adjacent intervals; ventral interval
(aa) about 3-4 ab, throughout; approximately 36
or 48 setae in XII (H, Pl), Nephropores in a
straight series on each side, slightly lateral of
middorsum in the forebody, though near mid-
ventral in segment II; dorsolateral in the hind
body. Clitellum annular, XIV-1/2XVIII but ex-
tending to 18/19 immediately lateral of the male
porophores. Male pores (H, P2-4) minute orifices
on small elliptical porophores in the setal arc of
XVIII in setal lines d-e, each preceded and suc-
ceeded by a large crescentic swelling which con-
stitute a larger porophore which protrudes
slightly into the adjacent segments, XVII and
XIX. Genital markings absent. Female pores not
visible but presumably paired in the transverse
furrow situated anteriorly in XIV. Spermathecal
pores 2 pairs, in 4/5 and 5/6, dorsolateral, in setal
lines 11-12 (H).

Septa 7/8 - 10/11 greatly thickened and funnel-
shaped. Dorsal blood vessel single, continuous
onto the pharynx. Last hearts in XIII, latero-oe-
sophageal, like the preceding hearts, slender. Giz-
zard in Vl, large, strongly muscular, almost

581

A imm

XVII

XVIII

FIG. 7. Terrisswalkerius windsori sp. nov., Holotype,
QMG21 1466. A, dorsolateral view of anterior end. B,
ventral view of male genital field.

582

spermathecal
diverticulum

spermathecal
ampulla

FIG. 8. Terrisswalkerius windsori sp. nov., Holotype,
QMG211466. A, right prostate. B, right posterior
spermatheca.

globose but its anterior limit straight sided. Cal-
ciferous glands and oesophageal dilatations ab-
sent. Intestinal origin XVII. Typhlosole absent.
Nephridia stomate, vesiculate holonephridia;
Those in П large and very complexly tufted and
with elongate bladder opening anteriorly near the
midventral line; those in HI and IV with bladders
apposed to those of II but progressively more
dorsal and, though tufted, smaller; those in V and
VI with very long ectally tapenng bladders, lack-
ing diverticula, discharging approximately in line
with the spermathecae and compactly tufted; the
duct joins the bladder slightly subterminally in
anterior segments but with no true diverticulum.
Nephridia not visible in УП - XT; in XII with very

MEMOIRS OF THE QUEENSLAND MUSEUM

elongate bladder; by XV bladders are sessile sub-
spheroidal, with duct entering subterminally.
Large, nacreous, free male funnels in X and XI.
Seminal vesicles large and racemose on the ante-
rior face of XII and small and smoothly sub-
spheroidal (pseudovesicles?) on anterior face of
X. Ovaries delicate laminae of oocytes, in XIII.
Prostates tubuloracemose; in the holotype dorso-
ventrally depressed thick plates which are resolv-
able into a tortuous structure; duct coiled and
muscular, joined entally by the vasa deferentia;
inone paratype, however, the prostates projecting
at the amputated end are coiled tubes much as in
T. nashi. Spermathecae 2 pairs, each with elon-
gate ovoid ampulla the tapering distal portion of
which is joined near its ectal end by a clavate
diverticulum which is about one fourth of the
length of the spermatheca and ectal to which is a
very short common duct.

ETYMOLOGY
For the Windsor Tableland.

REMARKS

Only Terrisswalkerius montislewisi resembles
T. windsori in having (as a variant in the former)
two pairs of spermathecal pores but T. windsori
is unique in the genus in the dorsolateral location
of the pores and in presence of the first pair in
intersegmental furrow 4/5 rather than 5/6.

In view of variation from two to three pairs of
spermathecae in T. lewisi, it is appropriate to
make comparison between T. windsori and spe-
cies with three pairs of pores. Of these, both T.
raveni and T. barronensis have spermathecal
pores far separated from the ventral midline (in d
or d-e lines) but only T. raveni has short sperma-
thecal diverticula as in T. windsori. It also has
large male porophores and the intestinal origin in
XVII as in T. windsori but it differs in having
more numerous setae, in location of the last hearts
in XII (not XIT), in possessing true nephridial
diverticula, and four pairs of seminal vesicles.
The male porphores of T. windsori are also dis-
unctive in form.

Observed variation in the apperance of the
prostates, from a plate-like, albeit composite,
mass to a coiled tubular form is unusual and
suggests that the appearance of prostate glands is
considerably influenced by the conditions of fix-
ation.

NEW EARTHWORMS

Tribe DICHOGASTRINI
Cryptodrilus bunyaensis sp. nov.
(Figs 9, 10)

MATERIAL EXAMINED

HOLOTYPE: QMG211508.

PARATYPES: 10 examined for distribution of genital
markings, QMG211509; 63 additional paratypes, QM
G211510. All types clitellate. All specimens from
26'5T' S, 151735'E, red soil under leaf litter in Arau-
caria bidwilli forest, Bunya Mts, National Park, Qld.
Collected B.G.M. Jamieson, E. Bradbury, 17 Mar.
1971.

DESCRIPTION

Length 76mm, width (midclitellar) 3.5mm.
Segments 185 (Holotype). Colour pigmented
light brown with dark reddish brown clitellum, in
ethanol. Prostomium very small, tanylobous,
though crossed by a transverse furrow behind its
midlength; peristomitim very short and prosto-
mium not or only partly visible dorsally; peri-
stamium not demarcated from first setigerous
segment (II) ventrally and only by an indistinct
transverse groove dorsally to which the dorsal
tongue of the prostomium extends. First dorsal
pore 9/10 but dorsal pores indistinct. Setae 8 per
segment, black and distinctly visible, in regular
longitudinal rows; ab moderately closely paired
but cd widely spaced, c being lateral and d far
dorsal; aa:ab:be:cd:dd = 2.7: 1.0: 3.4: 2.3: 5.1; or
12.7:4.7:16.0:10.8:24%. Clitellum saddle-
shaped, in XI-XVII: intersegmental furrows
12/13 and 18/19 well developed but the interven-
ing intersegments indicated weakly and ventrally
only. Male pores in ХУШ; minute pores with thin
lips, in ab, these setae being absent, each pore in
a glandular depression the slightly raised oval
margins of which extend median of a lines and
lateral of Б lines. Genital markings paired "eye-
like" medianly conjoined oval markings with pre-
setal porelike depression in ab, the markings
situated in XI, XII, XV-XVII, XX and XXI; the
posterior limit of each marking in ХІ, XII XX and
XXI shortly behind setae ab; but in XV, XVI and
XVII, like the field in XVIII, reaching the poste-
rior border of the segment. Ten paratypes were
examined for distribution of genital markings;
they resembled the holotype exactly with the
exception that one specimen lacked the markings
in XI and XII; one lacked those in XII; and all but
three specimens lacked the markings in XXI; of
the 10 paratypes, in one the marking in XT was
unilateral left, and in another the marking in ХХІ
was unilateral right. Female pores minute each
with a weak oval field, far apart, shortly an-

583

prostomium

ÎS

y {used peristomium (11
& first setigerous
segment (11)

spermathecal
роге 2

genital
markings

paired female
pore

genital
markings

XVI /
VIE
male pore
хіх minute papillas
ХХ f= г genital
xxi markings
XXII

FIG. 9, Cryptedrilus bunyaensis sp, nov,, Holotype,
QMG211508, Ventral view of forcbody and clitellar
region,

teromedian of setae a of XIV. Spermathecal pores

2 pairs, minute and indistinct, on small oval pa-

pillae at the anterior margins of УШ апа IX which

deflect intersegmental furrows 7/8 and 8/9 ante-
riorly in their vicinity.

Dorsal blood vessel continuous onto the phar-
ynx. Last hearts in XII. Gizzard large, broad,
cylindroid in V, with conspicuous anterior rim
which 3s continuous with a thinner walled pro-
ventriculus lying in IV. Oesophagus with paired
lateral dilatations in. VIII-XIV; those in XXIV

584

spermathecal
diverticulum

spermathecal
duct

spermathecal
ampulla

FIG. 10. Cryptodrilus bunyaensis sp. nov., Holotype,
QMG211508. A, ventral view of right prostate gland.
B, ventral view of right spermatheca.

forming true extramural calciferous glands with
internal lamellae; but only those of XH and XIII
constricted off from the oesophagus; best devel-
oped jn ХШ where they are reniform. Intestinal
origin abrupt, in XVI. Typhlosole absent. Nephri-
dia in the anteriormost segments several con-
loried, and sometimes almost tufted, large
meronephridia on each side but none found to be
enteronephric. In the anterior intestinal region,
shortly behind the prostates, three large
megameronephridia on each side with slender
exonephric ducts entering the body wall in setal
lines b, bc(nearer c) and in d; caudally, the largest
nephridium, discharging in b line, has a large
preseptal funnel and is thus à stomate
megameronephridium; no enteronephry demon-

MEMOIRS OF THE QUEENSLAND MUSEUM

strable. Holandric; testes and funnels not cer-
tainly demonstrated but free coagulum (putative
sperm masses) in X and XT; and seminal vesicles,
racemose with many berry-like loculi, in IX, XI
and XII. Ovaries large masses of many oocytes,
in ХП. Prostates elongated through several seg-
ments, broad and racemose; each with a slender
convoluted duct about one fifth the length of the
gland. Spermathecae 2 pairs, in VIII and IX; each
with a large ovoid ampulla, a slender duct a little
more than half the length of the ampulla, and a
short, digitiform diverticulum, with sper-
rmatozoal iridescence, sessile at the junction of
the two.

ETYMOLOGY
For the Bunya Mountains.

REMARKS

Cryptodrilus bunyanensis resembles C. medi-
ocris Fletcher, from New South Wales, in loss of
the intersegmental furrow between the peristom-
ium and the first setigerous segment. Although
this feature occurs in other genera (e.g. Digaster),
itis rare and its presence in the two Cryptodrilus
species gives some additonal support to inclusion
of the new species in the genus. It differs from C.
mediocris in lacking nephridial bladders and
from ali species of the genus in the distinctive
arrangement of genital markings.

Digaster lingi sp. nov.
(Figs 11, 12)

MATERIAL EXAMINED

HoLoTrYPE: QMG211484 (clitellate). From са.
28'12',8, 153 11 E., Binna Burra, Qld., in rainforest.
Collected E. Ling, M. Cunningham,

DESCRIPTION

Length (posterior amputee). Width (midclitel-
lar) 7.2mm. Pigmentless in life. Prostomium tan-
ylobous, with broad, posteriorly narrowing dorsal
tongue. Peristomium short; not canaliculate.
Forebody segments, at least from VI posteriorly,
strongly biannulate; traversed by a presetal
groove which is almost as pronounced as the deep
intersegmental furrow: postclitellar segments at
first biannulate, becoming weakly triannulate.
First dorsal pore 4/5. Setae small and, especially
the lateral pair, difficult to discern; both pairs
moderately closely paired; in XIV, aa:abzbc:cd =
2:1:2,8:1. Clitellum annular, strongly protuber-
ant, but weaker midventrally: dorsally XIII-
XVIII, ventrally XIII-2/3XVIIT; setae visible,
intersegmental furrows largely obscured; dorsal

NEW EARTHWORMS

pores suppressed. Male pores a pair of thin, trans-
versely extensive slits in XVIII on a pair of low
but large, transversely oval papillae which extend
from a lines into bc (relative to adjacent seg-
ments; setae a and b absent in this segment.
Genital markings: a slightly tumid elliptical area
fills the region between the male porophores from
the posterior limit of the setal annulus of ХҮП to
the anterior limit of that of XIX interrupting the
clitellum ventrally; it bears a pair of well defined
oval depressions at the site of intersegmental
furrow 18/19 which is suppressed on the tumid
area. Female pores inconspicuous, anteromedian
to setae a of XIV, in a transverse slit. Spermathe-
cal pores 2 pairs, in intersegmental furrows 7/8
and 8/9, in a lines, small, and visible only when
the intersegment is parted,

Septa 7/8-11/12 very strongly thickened, espe-
cially 8/9-10/1 1. Dorsal blood vessel single, con-
tinuous onto the pharynx. Last hearts very large,
in ХП; those in Х-ХП latero-oesophageal but
only those in XII shown to have (thin) connec-
tives to the dorsal vessel. Two moderately large,
subspheroidal muscular gizzards, in VI and VII,
separated by a thin-walled proventriculus-like
region. Oesophagus vascular but lacking calcif-
erous glands; strongly dilated and intestine-like
in XIV, Intestinal origin XVIII. Nephridia (cau-
dal region of body not available) stomate,
avesiculate micromeronephridia; in the vicinity
of the prostates on each side there is a ventral
convoluted nephridial mass lateral to which are
about 10 small nephridia in transverse single file,
each with a long, thin ectal duct, egress of the
ducts at the body wall scattered between the
anterior and posterior limits of the segment. Tes-
125? and nacreous funnels in X and XI, the poste-
rior pair better developed. Seminal vesicles
racemose, in IX and XII, the latter pair the larger.
Ovaries represented by many thin, empty strands,
Prostates large racemose, flattened discs, each
with a short muscular ectal duct, in XVIII. Sper-
mathecae 2 pairs, in VIII and IX, each with an
ovoid ampulla and a shorter duct which is joined
ectally by a small, rounded and apparently
pluriloculate diverticulum.

ETYMOLOGY
For Edmund Ling who coilected the specimen,

REMARKS

Three species of Digaster, D. gwongorellae,
D. minor, and D. moretonensis, in addition to D.
lingi, have a midventral circular, oval or almost
bifid pad or tumid area on XVIII between or

585

4— male pore

genital
marking

FIG. 11. Digasier lingi sp. nov., Holotype,
QMG211484. A, dorsal view of anterior end showing
prostomium. B, ventral view of forebody and clitellar
region.

including the male pores. D. lingi differs fram àll
three in having genital markings at 18/19 and
differs further from D. gwongorellae and D,
moretonensis in having paired female pores. D.
ling: also differs from D. minor in having the
lateral setal couples about as wide as, rather than
twice às wide as, the ventral couples. D. lingi
further differs from D. moretonensis, among
other respects, in the absence of genital markings.
on segments XIX to XXIII and the much shorter

spermathecal
diverticulum

spermathecal
ampulla

FIG, 12. Digaster lingi sp. nov., Holotype,
QMG211484. A, left prostate gland. B, right posterior
spermatheca,

clitellum which extends to XX in D. moretonen-
sis,

Digaster moretonensis sp. nov.
(Figs 13)

MATERIAL EXAMINED

HOLOTYPE: QMGH 1445,

PARATYPE: QMGH1444. Both from 27°11’S,
15324" E, Moreton Island, Eager's Swamp. pitfall trap,
ae Collected Queensland Museum Party, 22 Sep
1982.

DESCRIPTION

Length 190 (P)-205 (Н) mm, width 9.8mm (H)-
11mm (P). Segments 218(H). Pigmentless in al-
cohol with light chocolate brown clitellum. Pro-
stomium prolobous; it and the peristomium with
several parallel longitudinal dorsal furrows. First
dorsal pore well developed, at 5/6 (H, P). Setae
minute; in holotype and paratype arrangement
not certainly determinable in the forebody; but
shortly behind the male pores four pairs, ab

ULIM OG TUE CO TEENCI ANPMITSETTAE

MEMOIRS OF THE QUEENSLAND MUSEUM

closely paired; cd slightly more widely paired; c
and especially d lines irregular; dd = 0.5u.
Clitellum from presetal XIII-XX ventrally, to
XXI dorsally; a posteriorly widening median
strip is aclitellate in XVII posteriad and is pale
and possibly interrupted in XV-XVI, and paler,
though complete, in XIV, annular but weaker and
less pigmented in XIII. Male pores a pair, minute,
lacking definite porophores, equatorial in XVIII,
in ab, in a non-glandular patch which extends
laterally beyond P lines relative to adjacent seg-
ments, setae ab being absent in this segment.
Accessory genital markings (H only) midventral
pale brown transverse oval patches, including the
ventral pairs of setae, in XX-XXIII, best devel-
oped in their postsetal portions. Fernale pore sin-
gle. slightly displaced to the right (H) or
midventral (P). A midventral pore presetal in ХШ
(H) also, in a glandular craterlike elliptical field
(ndicated as a genital marking in Fig. 13A).
Spermathecal pores 2 pairs, in 7/8 and 8/9, very
slightly median of a lines; those in 8/9 2.0mm
apart.

Dorsal blood vessel single, continuous onto the
pharynx. Last hearts in XII; those in X-XII large,
tortuous and latero-oesophageal, Two very large
globular gizzards, in VI and VII, with a very short
non-muscular zone between; calciferous glands
absent; intestine commencing with abrupt expan-
sion in XVIII but septum 17/18 pressed forward
into XVII. Small, nacreous, free sperm funnels in
X and XI; small, compact racemose seminal ves-
iclesin XI and XII. Nephridia numerous avesicul-
ate micromeronephridia observed in midbody in
segments in the vicinity of prostates, No
buccopharyngeal enteronephry observed. Cau-
dally, the medianmost nephridium greatly en-
larged as à megameronephridium with preseptal
funnel. Ovaries with many chains of small oo-
cytes, and large funnels, in XIII. Ovisacs absent,
Prostates racemose, each a rectangular lobe in
XVIII with its anterior (ental) end reaching pos-
terior XVI. Duct almost as long as the gland,
curved mediad and widened ectally. The duct
with three median diverticula of which the largest
is about half the length of the duct, the shortest
(anterior) almost subspheroidal, and the third in-
termediate in length. Vasa deferentia not ob-
served. Penial setae absent. Spermathecae 2 pairs,
in VIII and IX, each with an ovoid ampulla,
broad, poorly demarcated ectally tapering duct,
and multiloculate diverticulum on the dorsoanter-
ior aspect of the ectal end of the duct.

NEW EARTHWORMS

dorsal pore
spermathecal pore 1

FIG. 13. Digaster moretonensis sp. nov., Holotype.
QMGH144$. A, ventral view of forebody and clitellar
region. B, dorsa! view of anterior end. C, right prostate
gland. D, right posterior spermatheca.

ETYMOLOGY
Named after Moreton Island, the type-locality.

REMARKS
D. moretonensis is distinguished from other
species of the genus by a combination of charac-

587

ters: the presence of midventra! segmental genital
markings in XVIII to XXIII, the long clitellum,
extending to XX, and the unpaired, midventral or
unilateral female pore. The diverticula of the
prostate ducts are unknown in other oligochaetes.
They possibly represent branches of the duct typ-
ical of racemose prostates, which have not devel-
oped glandular coats.

Tribe MEGASCOLECINI
Oreoscolex retrocystis sp, nov.
(Figs 14A, 15)

MATERIAL EXAMINED

HoLorTvrE: QMG211416.

PARATYPES: QMG211417-211421, all types from
S.E., Galston Gorge near Sydney, N.S.W., in dark
sandy soil bordered above and at the sides by large
rocks, about 20 feet above the river shortly below the
үес bridge, Collected В.С.М, Jamieson, 23 Мау
1962.

DESCRIPTION

Lengths (3 intact paratypes) 75-86mm, mean
79.7mm. Width (segment XIV in holotype and 3
paratypes) 2.4 -3.1, mean 2.6mm. Segments 162-
172, mean of 3-168. Pigmentless in ethanol. Pro-
stomium very small, closed, epilobous 1/4 to «1/2
but may be retracted into the mouth so as to
appear prolobous. First dorsal pore 10/11 (holo-
type and 3 paratypes) or 11/12 (1 paratype). Setae
8 per segment, in regular longitudinal rows; small
but distinct on well demarcated setal annuli; in
XII (Н) aa:ab:bc:cd:dd-3.5:1:3,1:2.3:9.4; or
13.5:3.9:12.2:8.9:36,6%. Clitellum not deyel-
oped in the holotype; in three paratypes, repre-
sented by slight annular tumescence and partial
occlusion of dorsal pores in XIV-XVII, and dor-
sally in XIII; intersegmental furrows clearly de-
fined and, in view of the maturity of the
specimens, probably never obliterated by the
clitellum, Male pores minute on a pair of very
small papillae on XVIII, at the site of the absent
setae a. In the holotype and the, fully mature,
paratypes (QMG211417-21), accessory genital
markings are very large sharply defined, raised
subcircular intersegmental markings, with pore-
like centres in ab in 16/17 and 19/20. In 16/17
they extend to the setal annuli of the two adjacent
segments, where they are paired or (H and Para-
type QMG211417) unilateral, right. In 19/20,
they are paired (H and paratype QMG211417) or
(paratypes QMG211418, 21) unilateral, right. In
two specimens they are absent from 19/20
(paratypes QMG21 1419 and 20), The male papil-
lae lie at the lateral extremity of a deep depres-

588

genital
markings
male pore

genital
markings

FIG. 14. Oreoscolex retrocystis sp. nov., Holotype,
QMG211416. A, ventral surface, from the spermathe-
cal to the male genital regions.

sion, with the form of a wide partly open mouth,
the tumid lips of which are sharply defined at the
inner margin, which forms parentheses closely
lateral to the male papillae, and in the midline
circumscribes a large median papilla and (visible
in the holotype only) in front of this a small
median papilla; the tumid lips extending in the
midline to the setal annuli of segments XVII and
XIX. Female pores minute, less than one third aa
apart, in an inconspicuous transverse slit which
coincides with the furrow anteriorly limiting the
setal annulus of XIV (holotype, 1 paratype) or

MEMOIRS OF THE QUEENSLAND MUSEUM

small gaping slits slightly anterior to the setal
annulus on a common oval tumid area (3
paratypes). Spermathecal pores 2 pairs, on well
marked small, hemispheroidal papillae anteriorly
in VIII and IX, nearer 7/8 and 8/9 than they are
to the setal annulus, in a lines (holotype and 4
paratypes).

The holotype and paratype QMG211421 were
dissected for internal anatomy. Dorsal blood ves-
sel single, continuous onto the pharynx. Last
hearts in XIII; hearts in X-XIII each arising in
common with a latero-oesophageal vessel from a
short common lateral vessel from the dorsal ves-
sel. No hearts recognized in VIII and IX in which
the latero-oesophageal vessel encircling the cal-
ciferous glands is also much less developed than
in X-XIII. Gizzard very large, firmly muscular,
in V, cylindrical but slightly tapering posteriad,
deflecting septa 5/6 and 6/7 posteriorly. Calcifer-
ous glands very large, paired in VIII-XIII (6
pairs), each reniform and well demarcated from
the oesophagus which it joins only at its hilus.
Intestine commencing with abrupt expansion in
XVI. Typhlosole dorsal, low in XVI, well devel-
oped in ХУП and by XVIII becoming a very large
inverted T-shaped structure. Nephridia
meronephridia; large tufts anterior to the gizzard
but lying in V discharge by wide ducts into the
pharynx; in the oesophageal region and anterior
intestinal region many stomate avesiculate
micromeronephridia; caudally with several pre-
septal nephrostomes and a very conspicuous lon-
gitudinal ureter which is situated dorsally to the
dorsal blood vessel and continues posteriorly; it
is not traceable to its termination in the holotype
which is a posterior amputee but in the paratype
appears to enter the intestine about 25 segments
from the anus. Iridescent sperm funnels in X and
XI; seminal vesicles compact and racemose, in
XII only. Ovaries small, palmate, in XIV. Pros-
tates elongate racemose, the gland in XVIII and
XIX, with distally widening muscular curved or
straight duct; vas deferens running up the duct to
join the junction of gland and duct. A large sub-
spheroidal firm mass, presumably glandular or
glandulomuscular, corresponding with each of
the paired accessory genital markings and with
the large midventral marking. Spermathecae 2
pairs, in VIII and IX; each with a narrow pyriform
ampulla, a short, digitiform diverticulum (unin-
seminated in the holotype; with iridescent sperm
masses in the paratype), and a moderately slender
fairly muscular duct.

NEW EARTHWORMS

genital marking gland
at 16/17

"X prostate
prostate 7%, gland

duct

genital marking
gland at 19/20

spermathecal-£^ ^»
duct л

spermathecal
diverticulum

spermathecal
ampulla

FIG. 15. Oreoscolex reirocystis sp. nov., Holotype,
QMG211416. A, right prostate. В, ventral view of
right spermatheca of IX.

ETYMOLOGY

From retro, backwards, and cyst, a thin-walled
hollow organ, alluding to the location of the sper-
mathecal pores behind rather than in the inter-
segmental furrows.

REMARKS

Oreoscolex retrocystis resembles O. saccarius,
in addition to their general Oreoscolex characters,
in having the first dorsal pore in 10/11; inter-
segmental accessory genital markings; 2 pairs of
spermathecal pores on small protuberances; 6
pairs of reniform calciferous glands, in VIII-XTIT;
and last hearts in XIII. It agrees with the type
species, O. imparicystis, in having segmentally

589

located spermathecal pores (there posterior in VII
and VIII) and in having seminal vesicles in XII
only. It differs from all other Oreoscolex species,
and, indeed, all known oligochaetes, in the con-
figuration of the genital markings.

Propheretima gen. nov.

TYPE SPECIES
Propheretima eungella sp. nov.

DIAGNOSIS

Perichaetine; setae numerous per segment; a
few present between the male pores. Female pore
single, midventral (or sometimes, P. hugalli,
paired but very close together?). Clitellum lim-
ited to XIV-XVI. Gizzard in V. Calciferous
glands present or absent. Intestinal caeca absent.
Testes in X and XI.

D ESCRIPTION

Peristomium ventrally bisected or not. Dorsal
pores present. Setae numerous in setigerous seg-
ments; a few present between the male pores.
Clitellum annular, XIV-XVI. Male pores in
XVIII; porophores typically but not always pre-
ceded and succeeded by an intrasegmental cres-
centic groove. Intrasegmental accessory genital
markings present or (if immature?) absent. Fe-
male pore unpaired (typically) or paired. Dorsal
blood vessel single. Last hearts in XII or XIII.
Gizzard in V. Calciferous glands present or ab-
sent. Meronephric, typically with intrasegmental
funnels. Enteronephry unknown. Holandric;
seminal vesicles in IX and XII or IX, X, XI and
XII. Prostates tubuloracemose or racemose (sin-
gle lobed). Ovaries in XIII. Spermathecae each
with a single diverticulum.

ETYM OLO GY

Propheretima, referring to an apparent
plesiomorphic relationship to Pheretima sensu
lato.

OTHER SPECIES
Propheretima (= Megascolex)
(Michaelsen, 1916). Queensland.
Propheretima hugalli sp. nov. New South
Wales.

fuscus

SPECIES INCERTAE SEDIS

Propheretima (=Megascolex) heterochaeta.
Queensland.

Propheretima (= Perichaeta) newcombei
(Beddard, 1887). Queensland.

REMARKS

Jarnieson (1977) drew attention to the likeli-
hood that the great oriental assemblage of species
in the genus Pheretima sensu lato, which now
includes the Australo-Papuan genus Begemius
Easton, had its ancestry in Australia. It was sug-
gested that the Australian genus Spenceriella was
the nearest known extant relative of Pheretima,
The species here placed in the new genus Pro-
pheretima fall within the definition of
Spenceriella given by Jamieson (1972) and
Jamieson & Wampler (1979). However, they are
distinguished from other species placed in
Spenceriella by a combination of characters:
presence of setae between the male pores, restric-
tion of the clitellum to segments XIV-XVI, and,
typically, a single female pore, all of which are
features of Pheretima sensu lato. They differ
from Pheretima sensu lato in having the gizzard
in segment V, not the apomorphic location in
VIII, and in at least as many features as separate
the various genera now recognized within that
assemblage, Within Propheretima, the type spe-
cies, Propheretima eungella and P. fuscus appear
to be closely related, sharing crescentic grooves
which bound the male porophores anteriorly and
posteriorly,

P. hugalli is placed in Propheretima because
it has the clitellum restricted to XIV-XVI and
sctae, albeit only setae a and 5, between the male
pores. It differs from P. eungella in apparent
pairing of the female pores, though that they are
not medianly fused requires confirmation from
further material.

Placement of P. heterochaera and P. new-
combei in Prapheretima is uncertain. Р. new-
Combel is included because it has a single female
pore and appears from an illustration by Beddard
(1887) to have setae between the male pores.
However, if Beddard's account js correct, it is
unique in the Megascolecidae in location of the
gizzard jn IV, V and VI which retain their inter-
vening, rather delicate septa. The species is inad-
equately described, the types have been lost and
the author has been unable to collect it in the type
or other localities. Gates (1965) synonymy of the
New Zealand Megascolex laingi Benham in M,
newcombet is not here accepted.

P. heterochaeta is here included in Pro-
pheretima chiefly because Michaelsen (1916)
considered this species to be near Megascolex
(here Propheretima) newcombei which differed
in the form of the accessory genilal markings.
Inclusion in Spenceriella is contraindicated by
the large setal numbers (68 in segment 50), intes-

MEMOIRS OF THE QUEENSLAND MUSEUM

linc beginning in XVII (XV or XVI in
Spenceriella), and location of seminal vesicles in
X and XI (in addition to IX and XII). Details of
the meronephridia are not known. Because of the
immature state of the types, the generic place-
ment of this species, and indeed its specific char-
acteristics, remain uncertain.

KEY TO THE SPECIES OF PROPHERETIMA

1. Each male porophore immediately preceded and
succeeded by an intrasegmental crescentic
STOVE L^. php аја mip co. uno fmf дүе рб on 2

Each male porophore not immediately preceded
and succeeded by an intrasegmenial crescentic
grove lc ke СЕ Саге К

2. Genital markings on X, XVII-XX. Three pairs of
well developed sessile calciferous glands in X-
XII P, eungella

No preclitellar genital markings (immature?), [A

slightly raised transverse ficld may be present
midventrally in XVIII, and also posteriorly in
XVII and anteriorly in XTX], Calciferous glands
ЖЛ, es. ele te ee +. + P. fuscus

3. Each male porophore preceded and succeeded, al
the intersegment and slightly more medianly, by
a puckered pit, A pair of transverse pads
equatorially in X with a small presctal, pore-like
depression, .. . . P, hugalli

Each male porophore not preceded and suc-

ceeded by a puckered intersegmental pit. Genital
markings not as 3(1) 4

4. Prostomium epilobous. Genital markings not defi-

nitely developed. |. 5... P. heterochaeta
Prostomium prolobous. Preclitellar genital mark-

ings present to as far forward as VIL [Genital
markings very variable, When most numerous:
segment ХШ with a single, midventral papilla;
XI and ХИ each with three papillae close to-
gether, one median and one on cach side of this;
X with four papillae, of which the middle ones
correspond in position to the median papillae of
the two succeeding segments; TX with a single
papilla, corresponding in position to the outer-
most right hand one of segment X, the others
being indistinct; VII and УШ cach with a single
median papilla] Р. newcomber

3

оа оаа ааа а

ei e n ay ano

б жж ж ж ж s t od on n on ng

Propheretima eungella sp. nov.
(Figs 16, 17)

MATERIAL EXAMINED

HOLOTYPE! QMG211434 (clitellate). All specimens
from 21"08'S, 148°30'E, Bungella National Park, Palm
circuit, Qld., in a rotting log. Collectors A. Postle and
G, Dyne, 13.Jun,1974-Qid.

NEW EARTHWORMS

markings

FIG. 16. Propheretima eungella sp. nov., Holotype.

QMG211434. Forebody and clitellar region
DESCRIPTION

Length 61mm, width 3.7mm, scgments 93,
Colour in ethanol dorsally dark steel grey; pale
ventrally; clitellum buff. Prostomium cpilobous
1/2. Peristomium not ventrally bisected. First
dorsal pore 4/5. Setae in XII, 34;

591

aa:ab:be:cdiyz:zz = 1,8:1.0:0,8:0.8:1.6:2.8; dor-
sal break small but appreciable in the forebexly,
caudally not appreciable and ventral break
scarcely greater than adjacent setal intervals;
about 12 segments from the posterior extremity
60 setae per segment; a few present between the
male pores. Clitellum annular, XIV-XVI. Male
pores narrow elongate slits on two large, trans-
versely oval porophores in XVIII, with centres in
cd, in the setal arc, each porophore bordered
anteriorly and posteriorly by a crescentic glandu-
lar depression. Accessory genital markings: eye-
like markings consisting of a central small
transversely oval papilla and a dark glandular
field with a white rim, not noticeably raised, a pair
presetally in cd and one median in X; in XVH a
presetal, medianly conjoined pair with centres
just lateral of b, and a postsetal pair in d lines; in
XVII, minute markings anterolateral and
posterolateral to the male porophores, approxi-
mately in setal lines 6-7 relative to segment ХІХ;
in XIX a glandular area presetal froma to e lines,
each with two small eyelike markings of which
the lateral is the larger; in XX a presetal pair with
centres in b lines. Female pore unpaired, mid-
ventral and presetal in XIV. Spermathecal pores
2 pairs, in intersegmental furrows 7/8 and 8/9, in
setal lines 5-6, on small papillae.

Dorsal blood vessel single, Last hearts in XII.
A large, moderately compressible gizzard in V.
Three pairs of well defined subspheroidal calcif-
erous glands in X, XI and XII, each sessile but
well constricted off from the oesophagus; the
lumen of each gland with many radial vas-
cularized laminae almost meeting centrally. In-
testine commencing with abrupt expansion at the
anterior septum of XVI. Loose aggregates of
meronephridia jn the pharyngeal region send a
large duct forwards to the yicinity of the peri-
stomium but no enteronephry detected. Caudally
with 10 ог more meronephndia on each side of
which some at least have a long-stalked, in-
trasegmental, posteriorly directed minute funnel.
A single, larger preseptal funnel present on each
side medianly; no enteronephry demonstrated,
Free testes and iridescent funnels in X and XT;
very large racemose, much subdivided seminal
vesicles with large loculi and clusters of many
minute spheroidal bodies, paired in TX and ХП,
Prostates tubuloracemose, restricted to. XVIII;
superficially racemose lobes bul resolvable inta
a flattened S-shaped structure with three contin-
uous adpressed sections; each gland with a stout
looped muscular duct. Ovaries a pairin XIII, each
alarge mass of many oocytes with multiple stalks

592

prostate
duct

prostate gland

1mm

spermathecal
diverticulum

spermathecal
ampulla

FIG. 17. Propheretima eungella sp. nov., Holotype,
QMG?211434. A, right prostate gland. B, right poste-
rior spermathecca,

to the anterior septum; à pair of large funnels on
the posterior septum. Spermathecae 2 pairs in
VIII and IX, each discharging at the anterior
intersegment; each with a large transversely ovoi-
dal ampulla, a bulbous inseminated diverticulum
of approximately equal length, and a short wide
duct.

ETYMOLOGY
From Eungella National Park.

REMARKS

Propheretima eungella resembles only Р, fus-
cus, in the genus, in having a crescentic groove
bordering each male poraphore anteriorly and
posteriorly but differs from the latter species, and
all other members of the genus, in having genital
markings on X, XVII-XX and three pairs of well
developed sessile calciferous glands in X-XIII.

MEMOIRS OF THE QUEENSLAND MUSEUM

Propheretima hugalli sp. nov.
(Figs 18, 19)

MATERIAL EXAMINED

HOLOTYPE: QMG21 1452. Fromc. 28*49'8, 153^ 16 E,
Lismore, Boat Harbour, NSW, Collector A. Hugall. 12
Aug. 1994,

DESCRIPTION

Length unknown, width (midclitellar) 2.4mm,
number of segments unknown. Colour in ethanol
pinkish purple, paler ventrally in the postclitellar
region. Prostomium closed epilobous 1/3. Peri-
stomium very short, bisected ventrally. Body not
canaliculate. First dorsal pore 6/7. Setae peri-
chaetine, 30 in XII; in the forebody, dorsal (zz)
and ventral (aa) breaks small but just appreciable
relative to adjacent intervals; ventral break wider
behind the clitellum; caudally zz and aa remain
narrow and z line is irregular. Setae a and b
present between the male porophores. Clitellum
annular, XIV-X VI; strongly demarcated and pro-
tuberant; setae, dorsal pores and intersegments
obscured. Male pores in c lines towards the me-
dian aspect of a pair of large oval porophores
which are well demarcated medianly but not lat-
erally and which almost fill XVIII an-
teroposteriorly. Each porophore preceded and
succeeded, slightly more medianly, by a roughly
oval depression, in posterior XVII and anterior
XIX, with puckered anterior or posterior margins
respectively; these depressed genital markings
continuous with a relatively depressed area be-
tween the porophores. Other genital markings
consist of a pair of poorly defined transverse pads
equatorially in X which fill the segment longitu-
dinally; each with a small pore-like depression in
the setal arc, in c lines. A small transverse eleva-
tion in XIV appears to bear a pair of female pores
very close together but pairing requires confirma-
tion. Spermathecal pores 2 pairs, in 7/8 and 8/9,
only the posterior left pore visible, in d line,

Dorsal blood vessel single. Last hearts in XII.
Gizzard small, soft, virtually vestigial, in V with
slight muscular sheen which is also seen on the
slightly narrower oesophagus in 1V and anterior
УІ. Ocsophagus also whitish in VII; vascularized
and intersegmentally constricted in VIII to ХШ;
in ХІ, XIE and ХШ with a dorsal longitudinal
groove giving some appearance of extramural
calciferous glands and in these segments with
high internal rugae but not constricted off from
the oesophageal lumen. Intestine commencing in
XVI. Nephridia avesiculate micromeronephridia;
no funnels definitely present in the forebody and
clitellar regions; some loose tufting or aggrega-

NEW EARTHWORMS

peristomium

FIG. 18. Propheretima hugalli sp. nov., Holotype,
QMG211452, Ventral view of forebody and clitellar
region.

tion anteriorly: caudally tending to form lateral
aggregations and with occasional imtrasegmental
funnels observed; presence or absence of median
preseptal funnel not demonstrable; no en-
teronephry observed but absence requires confir-
mation. Free sperm masses and very large,
convoluted iridescent funnels in X and XT; semi-
nal vesicles racemose, in IX and XII. Prostate
restricted to XVIII; each a tongue-like lobe with
a minutely papillate surface indicative of race-
mose structure; the short narrow medianly di
rected duct joined entally by the conjoined vasa
deferentia. Ovaries a pair in XIII, each a relatively
large folded lamina with many chains of large
oocytes with multiple stalks to the anterior sep-

593

tum; a pair of funnels on the posterior septum
with ducts entering the body wall independently
but close together. Spermathecae 2 pairs in VIIT
and IX, discharging at the anterior intersegments;
each with a narrow bulbous ampulla, and a simi-
larly shaped smaller diverticulum which is about
two thirds the length of the spermatheca.

ETYMOLOGY
For Andrew Hugall, the collector.

REMARKS

Prapheretima hugalli differs from the other
species of the genus in having an intersegmental
puckered pit anterior to and behind each male
porophore and a pair of transverse pads equatori-
ally in X with a small presetal, pore-like depres-
sion.

Spenceriella conondalei sp nov.
(Figs 20, 21)

MATERIAL EXAMINED

HOLOTYPE: QMG211422 (ex QMG1 158).
PARATYPES: QMG211423-33, 10 clitellate specimens.
all from 26°45'S, 152°37'E, Conondale Range, Qld.,
in leaf litter. Collected R. Raven, 13-18 May 1976.

DESCRIPTION

Length 26-40mm, mean 33mm. Width 2.2mm.
Segments 81-109, mean 92 (n=4). Pigmentless
buff in ethanol. Prostomium epilobous 1/2. First
dorsal pore 5/6. Setae 18 in Ш; 30 in XII; 32 in
XX; about 26 in caudal segments (Н); аагаЬ;Ье:

prostate gland

vasa deferentia

spermathecal
diverticulum—

1mm

FIG. 19. Propheretima hugalli sp. nov., Holotype,
QMG211452. A, right prostate. B, right posterior
spermatheca,

594

genital markings

male pore and everted
copulatóry pouch

FIG. 20. Spenceriella conandalei sp. nov., Holotype,
QMG21 1422, Ventral view of forebody and clitellar
region.

cdzyz:zz- 1.6:1.0:0.5:0.5:0.5:1.0; dorsal and ven-

tral breaks not apparent; possible vestiges of two

setae, represented only by pore-like dimples, be-
tween the male pores; setae a sometimes absent
in the vicinity of the genital markings, in seg-
ments VI-X, Clitellum annular, best developed

MEMOIRS OF THE QUEENSLAND MUSEUM

on XIV-XVI but some development without tu-
mescence in 1/2XIII, 1/2ХШ and XVII. Male
pores large orifices centred in Б lines of XVIII,
each occupied by a strongly protuberant, whitish
everted copulatory pouch, Genital markings
paired, each consisting of a low whitish subcircu-
lar to elliptical mound, with presetal pore-like
centre; in VIII with the centre in bc; in IX im-
mediately median of c; in X in or very slightly
lateral of b; in XI in the immediate vicinity of a
line; in XVII, in ab and in contact with its partner
medianly; in XIX, in or immediately lateral of b;
in XX, in or immediately lateral of a; the mark-
ings on XIX and XX, but not their pore-like
regions, fused across the midline in each segment
to form a single midventral pad; on XVIII a tumid
area, extends laterally to about setae 5, and bears
a pair of small pore-like markings anterolateral
and another posterolateral to the male pores, lat-
etal of c, sometimes with an additional pore an-
teromedian to the male pore. Some markings,
particularly in УШ and/or ХУШ, may be absent
or unilateral in clitellate specimens. Female pore
single, midventral, shortly anterior to the setal arc
in XIV surrounded by a small elliptical field.
Spermathecal pores transverse slits with thin
white elliptical lips, in 7/8 and 8/9, their median
borders in or immediately lateral of 5 lines
Dorsal blood vessel single. Last hearts in XIII.
Gizzard moderately large, cylindrical and firmly
muscular, in V bnt deflecting septum 5/6 posteri-
orly so as to end posteriorly in VIII relative to
external segmentation. Calciferous glands absent
but oesophagus vascularized and with an encir-
cling blood vessel on each side in each of seg-
ments VIII-XIII. Intestine commencing in XV;
with a low lobulated and not uniformly continu-
ous dorsal ridge but no definite typhlosole; caeca
absent; intestinal contents include quartz grains
and pieces of wood, Several large meronephridial
tufts anterior to the spermathecae, some with
ducts to the pharynx, others exonephric. Several
apparently astomate meronephridia on each side
in anterior intestinal segments. Caudally with a
median preseptal funnel and apparently some
intrasegmental funnels though the latter were not
demonstrable in some specimens. Testes and
large iridescent funnels free in X and XI. Large
racemose seminal vesicles in IX and XII. Ovaries
conspicuous bunches of oocytes in XIII. Prostates
racemose, flattened, subrectangular with a small
accessory lobe behind the junction with the
curved muscular duct. Spermathecae 2 pairs, in
VIII and LX, each with a subspheroidal ampulla

NEW EARTHWORMS

prostate
du
vasa

X deferentia

A
spermathecal
spermaihecal diverticulum
duct
spermathecal
ampulla
B 1 mm

FIG. 21. Spenceriella conondalei sp. nov., Holotype,
QMG211422. A, left prostate. B, right posterior sper-
ca.

and a small, iridescent and therefore inseminated,
clavate diverticulum.

ETYMOLOGY
For the type locality.

REMARKS

S. conondalet is exceptional m Spenceriella in
having a single, not paired, female pore. The
presence of questionable vestiges, as minute pits,
of two setae between the male pores is therefore
noteworthy in view of diagnosis of Prapheretima
and Pheretima sensu lato by the presence of well
developed setae between the male pores. 5. con-
ondalei is excluded from both of these genera
because of the absence of recognizable setae at
these loci and because of its general similarity 10
Spenceriella. The. well developed gizzard places
it in the subgenus Spenceriella although the ab-
sence of calciferous glands is not typical of that
subgenus.

DISCUSSION

The present study has added to the endemic
earthworm fauna of Australia (Megascolecidae),
ane species of Acanthodrilinae (Rhododrilus
glandifera) and nine species of Megascolecinae,

595

in the tribes Perionychini (Heteroporodrilus
montiserratae, Terrisswalkerius windsorl,
Cryptodrilus bunyaensis); Dichogastrini (Digas-
ter lingi, D. moretonensis); and Megascolecini
(Oreoscolex retrocystis, Propheretima eungella
gen et sp. nov., Propheretima hugalli, and
Spenceriella conondalei). Rhododrilus gland-
ifera is the second species of this primarily New
Zealand genus to be described from Australia. It
is possible that it has acquired the microscolecin
arrangement of male pores (prostates one pair
discharging with the vasa deferentia in XVII)
from a precursor with the acanthodrilin arrange-
ment (prostate pores 2 pairs, in XVII and XIX;
male pores in XVIII) independently of the New
Zealand species. If this were so, retention in
Rhododrilus could not be supported and separate,
new generic status would be required.

The type localities of Heteroporodrilus
montiserratae, Terrisswalkerius windsori,
Digaster lingi, D. moretonensis, Oreoscolex
retrocystis and Spenceriella conondalei are well
within the ranges of their respective genera. How-
ever, the insular locality of D. moretonensis, on
Moreton Island, is of zoogeographic interest and
Cryptodrilus bunyaernsis represents a northem
extension for its genus. The new genus Pro-
pheretima, has been erected for species which
resemble the Oriental and Australo-Papuan
Pheretima sensu lato assemblage of genera in
having setae between the male pores but which
are plesiomophic in retaining the gizzard in seg-
ment V, not in the apomorphic location of VIII
seen in all pheretimas. It endorses the author's
view (Jamieson, 1977) that the ancestry of the
Pheretima assemblage may have lain іп Aus-
tralia.

ACKNOWLEDGEMENTS

The author is gzateful to those who collected or
aided in collection of material described in this
paper: E. Bradbury, M. Cunningham, G. Dyne,
A. Hugall, J. Ledger, D. Ledger, E. Ling, K
McDonald, A. Postle, and R. Raven. This study
was made possible by ABRS and ARC grants 10
the author. All illustrations are by the author.

LITERATURE CITED

DYNE, С.К. 1980, Rhododrilus sexpapillanes np.
(Oligochacta: Megaseolecidae) from New Zea-
pone ev Zealand Journal of Zoology 1980 (4):

596

GATES, G.E. 1965. On an Australian species of the
earthworm genus Megascolex Templeton, 1844.
Australian Zoologist 13: 213-215.

JAMIESON, B.G.M. 1974. The indigenous earth-
worms (Oligochaeta: Megascolecidae) of Tasma-
nia. Bulletin of the British Museum (Natural
History) Zoology 26: 201-328.

1972. The Australian earthworm genus Spenceriella
and description of two new genera
(Megascolecidae: Oligochaeta). Memoirs of the
National Museum of Victoria 33: 73-88.

1977. The indigenous earthworms (Megasco-
lecidae: Oligochaeta) of Lord Howe Island. Re-
cords of the Australian Museum 30: 272-308.

1994. Some earthworms from the wet tropics and
from Bunya mountains, Queensland
(Megascolecidae: Oligochaeta). Memoirs of the
Queensland Museum 37: 157- 181.

MEMOIRS OF THE QUEENSLAND MUSEUM

JAMESON, B. G. M. & WAMPLER, J. E. 1979. Bio-
luminescent Australian earthworms. II. Taxon-
omy and preliminary report of the
bioluminescence in the genera Spenceriella,
Fletcherodrilus and Pontodrilus (Megasco-
lecidae: Oligochaeta). Australian Journal of Zool-
ogy 27: 637-669.

LEE, K.E. 1952. Studies on the earthworm fauna of
New Zealand. III. Transactions of the Royal Soci-
ety of New Zealand 89: 23-45.

1959. The earthworm fauna of New Zealand. New
Zealand Department of Scientific and Industrial
Research, Wellington, Bulletin 130: 1-486,

MICHAELSEN, W. 1916 Results of Dr. E. Mjóberg's
Swedish Scientific Expedition to Australia 1910-
1913. Oligochaeten. Kungliga Svenska
Vetenskapakademiens Handlinger, 52 (13): 3-74.

A GROUND-TRAPPING SURVEY FOR SMALL MAMMALS IN CONTINUOUS
FOREST AND TWO ISOLATED TROPICAL RAINFOREST RESERVES

SUSAN G. W, LAURANCE & WILLIAM F. LAURANCE

Laurance, S. G. W. & Laurance, W. F. 1995 12 01: A ground-trapping survey for small
mammals in continuous forest and two isolated tropical rainforest reserves, Memoirs of the
Queensland Museum 38(2), 597-602. Brisbane. ISSN 0079-8335.

We used mark-recapture methods to compare small mammal assemblages between contin-
uous rainforest and two small (c. 500 ha), isolated rainforest reserves in north Queensland,
Lake Eacham and Lake Barrine National Parks. Both parks had abundant populations of
native rodents. Uromys caudimaculatus and Melomys cervinipes were significantly more.
abundant in the parks than continuous forest, while Rattus fuscipes was significantly less
abundant in the parks. A few non-rainforest rodents (Mus musculus and К, lutreolus) and the
edge-Favouring Antechinus flavipes were captured in the remnants, but not in continuous
forest. Hypsiprimnodon moschatus was present in both parks and continuous forest, while
several other rare, forest-dependent mammals (e.g. Antechinus godmani, A, stuartii,
Dasyurus maculatus) were not detected. Several of these trends have been observed in other
rainforest remnants jn the region, suggesting small mammal assemblages in these isolated
parks have been influenced by forest fragmentation. [ ] rainforest mammals, trapping, forest
fragmentation,

Susan С, W. Laurance, Queensland Department of Environment and Heritage, РО Box 834,
Atherton, Queensland 4883, Australia; William F. Laurance, CSIRO Tropical Forest
Research Centre, PO Box 780, Atherton, Queensland 4883, Australia; ! September 1995,

Forest fragmentation can cause a wide range of
ecological effects, including biotic and microcli-
matic changes associated with abrupt forest edges
(Andren & Angelstam, 1988; Kapos, 1989; Wil-
liams-Linera, 1990; Laurance, 19911), the de-
cline or loss of specialised, forest-dependent
fauna (Lovejoy et al., 1986; Laurance, 1991a),
and dramatic changes in abundances of some
species (Terborgh, 1992). Genetic changes, such
as a progressive erosion of allelic diversity and
heterozygosity, also may affect fragmented pop-
ulations (Leung et al., 1993),

Large-scale clearing and fragmentation of trop-
ical rainforest on the Atherton and Evelyn Table-
lands in north Queensland began in the early
J900's and proceeded rapidly for the next several
decades. By 1980, over 76,000ha af forest had
been removed (Winter et al., 1987), leaving only
scattered forest fragments on flatter areas of the
Tablelands, ranging from about 1 to 600ha in area
(Laurance, 19913).

Two of the largest forest remnants on the Ta-
blelands are Lake Eacham and Lake Barrine Na-
tional Parks. The parks were initially designated
as Scenic Reserves in 1888, then gazetted as
National Parks in 1934 (Matthews, 1993), Each
park is about 500ha in area with relic volcanic
lakes occupying 103ha at Вагппе and 52ha at
Hacham. An Eacham Parish map published in
1931 identifies property subdivisions completely

surrounding both parks, suggesting they have
been isolated from primary forest for at least 64
years.

The mammal faunas of the two parks have
received only limited attention. Non-systematic
trapping was conducted in both parks In the
1930's and the 1970's, mainly for mammal taxo-
nomic studies (Taylor & Horner, 1973). J. W-
Winter (pers. comm.) conducted spotlighting and
trapping surveys al Lake Eacham in the mid-
1970's, but Lake Barrinc has not been surveyed
systematically, nor have the mammal fannas in
the parks been contrasted with those in nearby
continuous forest.

The aim of this study was to contrast small
mammal assemblages at Lakes Eacham and Barr-
ine National Parks with those found in nearby
continuous rainforest. These findings are relevant
to the development of park management strate-
gies, such as the creation of faunal corridors,
which are currently being implemented to help
ameliorate the effects of forest fragmentation.

METHODS

Lakes Eacham and Barnne (740-800m eleva-
tion) are located near the NE margin of the
Atherton Tableland, The parks are separated from
Gadgarra State Forest (680-900m elevation) hv а
clearing 1.5-2.5km in width comprised by cattle
pastures and second-growth forest (Fig. 1). At

598

Lake
^j Barrine

NL 0;

`. Eaċham —T* y
"qc ENF o’

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Map of study area on the Atherton Tableland in north Queensland, showing locations of 17 trapping
transects. Lightly stippled areas are rainforest, darkly stippled areas are second-growth forest, and unstippled

areas are mostly cattle pastures.

Gadgarra, upland (>600m elevation) forests
occur in a relatively narrow (4-6km-wide) band
but are contiguous with the extensive Atherton
Uplands (100,000ha). Much of Gadgarra was
selectively logged in the 1950’s and 1960's
(Queensland Forest Service records). Forests at
all three sites are mostly complex mesophyll
vine-forest on basaltic soils, with patches of sim-
ple notophyll vine-forest on poorer metamorphic
soils (Tracey, 1982).

Eacham and Barrine were censused in late July
and early August 1994, while Gadgarra was cen-
sused in early October. Traps were positioned
along a series of 210m-long transects. Each tran-
sect had 8 cage traps (30 x 30 x 60cm) and 14
Elliot box-traps (10 x 11 x 30cm) spaced in a
regular sequence at 10 m intervals. Elliots traps
were baited with a mixture of rolled oats and
vanilla essence, while cages were baited alterna-
tively with the oats-vanilla mixture or dog food

(Chicken Chupp). Traps were operated for 4 con-
secutive nights.

Eacham and Gadgarra were each sampled with
6 transects (528 trap-nights each), while Barrine
had 5 transects (440 trap-nights). At each site, 2-3
transects were used to sample forest interiors,
while the remaining transects began at the forest
edge and proceeded toward the interior (Fig. 1).
Captured mammals were ear-tagged, weighed
and examined to assess sex and reproductive
condition, then released.

RESULTS

ABUNDANT RODENTS

We recorded 603 mammal captures (400 indi-
viduals) during the study. Four native rodents, the
fawn-footed melomys Melomys cervinipes, Cape
York rat Rattus leucopus, bush rat R. fuscipes,
and white-tailed rat Uromys caudimaculatus,

SMALL MAMMALS IN RAINFOREST REMNANTS

were numencally dominant, comprising 94-98%
of individuals captured at each site. The relative
proportions of the four species did not differ
significantly between the two forest fragments
(x226.63, р>0.05). while proportions in each
fragment differed strongly from those in contin-
uous forest (Eacham vs Сайрага: xy2244.41,
p«0.001; Barrine vs Gadgarra: 2237.18,
p<0.001; Chi-square tests for independence,
d.f=3 in all cases). Because rodent communities
in the two fragments similar in com
sition, we pooled data from the fragment sites for
the subsequent analysis,

We compared the mean abundapce of each
species in the two fragments to those in continu-
ous forest (Fig. 2). Melomys (p-0.019) and
Uromys (p=0.012) were both significantly more
abundant in the fragments, while Rattus fuscipes
was significantly less abundant (p=0.001). Rattus
leucopus was quite abundant at Barrine but was
patchily distributed and did not differ signifi-
cantly between fra; ted and continuous forest
(p=0.684), Total rodent abundance (i.e. the com-
bined abundances of the four common species)
did not differ significantly between fragmented
and continuous forest (p-0.546: all Mann-Whit-
ney U-tests), although rodent populations in the
fragments exceeded those in continuous forest by
an average of 14-1656,

RARE MAMMALS

Five species were captured too infrequently to
permit statistical analysis (1-6 individuals). Two
yellow-footed antechinuses Antechinus flavipes,
a swamp rat Rattus lutreolus, and a house mouse
Mus musculus were captured at Barnne. Musky
rat-kangaroos Hypsiprimnodon moschatus were
captured at Eacham (1 animal) and Gadgarra (2
animals), and were occasionally observed during
the day at all three sites. Finally, two long-nosed
bandicoots Perameles nasuta were captured at
cach of the three sites.

DISCUSSION

Our short-term survey was unlikely to provide
a complete census of small mammals in the study
area.

Repeated sampling during the major seasons of
the year, and the use of ground- and arboreal-
traps with both meat and herbivore baits, are
needed to provide a comprehensive sample of
rainforest small mammal communities (Laur-
ance, 1992). There also was a two-month interval
between censuses of the parks and the controls.

599

Results, therefore, should be interpreted with
some caution,

ABUNDANT RODENTS

Despite limited sampling our findings appear
similar in several respects to those observed in
larger (20ha) forest fragments on the southern
Atherton Tableland (Laurance, 1994b), suggest-
ing the small mammal faunas at Lakes Eacham
and Barrine exhibit some effects of forest frag-
mentation.

On average, assemblages of abundant native
rodents were 14-16% more abundant in Eacham
and Barrine than in nearby unfragmented forest,
although this difference was not significant. On
the southern Atherton Tableland (30-40km south
of Eacham and Barrine), the same rodent species
were on average 20-27% more abundant in frag-
ments than continuous forest (Laurance, 19941).
These increases may result from ecological
changes in fragmented rainforest, such as dense
ground and understorey cover associated with
forest edges and disturbed forest, that are fa-
voured hy some rodents. A recent comparison of
forest structural variables at Eacham and Barrine
with nearby continuous forest, suggested the
parks exhibit relatively heavy disturbance, which
may result from increased wind damage near
forest edges compounded by the relatively high
topographical position of the parks (Laurance,
19942). Native rodents may also increase in frag-
ments because of novel foraging opportunities 11)
surrounding pastures or secondary habitats. or
because larger, forest-dependent predators de-
cline in most fragments (Laurance, 1990; 1994p),

The fawn-footed melomys Melamys cervinipes
was significantly more abundant at Eacham and
Barrine than in unfragmented forest. On the
southern Tableland, this species also was more
abundant in both small (1.4-12.7ha) and large
(21-590ha) fragments than in unfragmented for-
est. Melomys appears to favour recent treefall
gaps and other disturbed forest, which probably
accounts for its success in remnants (Laurance,
1994b).

The bush rat Rattus fuscipes was much less
abundant in the parks than continuous forest,
while the Cape York rat R. leucopus was unusu-
ally abundant at Barrine but less abundant at
Eacham and the controls. Cape Yurk rats in-
creased on the southern Tableland in many
smaller («20ha) forest remnants while bush rats
generally declined. Like the fawn-footed
melomys. the Cape York rat appears to favour
forest edges and areas wilh many treefalls and

20
L] Melomys
ШЕШ Aattus fuscipes

15

10

ы М Ме. МЫ

AVAVA

ххх

XX

Individuals per 100 trap-nights

Barrine

Eacham

MEMOIRS OF THE QUEENSLAND MUSEUM

БУ А. leucopus
LAA Uromys

controls

FIG, 2. Standardised abundance estimates (mean + с) of four abundant rodent species from trapping transects in
Lake Eacham and Lake Barrine National Parks, and in nearby continuous forest (Gadgarra State Forest).

dense rattan growth, while the bush rat generally
avoids areas with high densities of Cape York rats
(Laurance, 1994b).

The white-tailed rat Uromys caudimaculatus, a
large (500-900g), aggressive omnivore, was un-
usually abundant in both parks. In a trapping
survey in the 1970's, J. W. Winter (pers. comm.)
also found the white-tailed rat to be the most
frequently-captured small mammal at Lake
Eacham. These results differ from observations
on the southern Tableland, where white-tailed rat
abundance did not differ significantly between
larger (>20ha) fragments and continuous forest
(Laurance, 1994b). Elevated abundances of
white-tailed rats in fragments could result in in-
tensified predation on large rainforest seeds (Os-
unkoya, 1993), nesting birds and small
vertebrates (Laurance et al., 1993; Laurance &
Grant, 1994). In future studies, it would be useful
to contrast the intensity of predation in the parks
with that in continuous forest, using seeds or
experimental bird nests (e.g. Laurance et al.,
1993; Harrington et al., in press),

RARE MAMMALS

Two non-rainforest rodents were captured at
Lake Barrine, the house mouse Mus musculus,
commonly associated with human habitations,
and the swamp rat Rattus lutreolus, which usually
favours open, grassy habitats. J. W. Winter (pers.
comm.) captured Mus musculus and Rattus rattus
at Lake Eacham in the 1970's, although we did
not encounter these species at Eacham in 1994,
Non-rainforest rodents also were encountered,
albeit infrequently, in many forest fragments on
the southern Atherton Tableland (Laurance,
1994b). Thus, forest fragments appear increas-
ingly prone to invasions by non-native rodents
although native rodents, which are abundant and
highly territorial, probably prevent large-scale
invasions of non-forest species.

Interestingly, Lake Barrine is the type locality
for Rattus lutreolus lacus, a disjunct north
Queensland subspecies initially encountered in
1937 (Taylor & Horner, 1973). In the 1970's J.
W. Winter (pers. comm.) attempted to confirm
the presence of R. J. lacus in the parks and inter-
vening paddocks, but did not encounter it. Our
study confirms the presence of swamp rats in the

SMALL MAMMALS IN RAINFOREST REMNANTS

area. The single animal we encountered was cap-
tured inside Lake Barrine Park, 160m from the
nearest forest edge.

The есюру of trapping of Antechinus varies
considerably during the year, peaking from about
March to June (Laurance, 1992), and the present
study would have been unlikely to sample these
species adequately. We captured only two yel-
low-footed antechinuses Antechinus flavipes at
Barrine, and none at Eacham or in continuous
forest. On the southem Tableland, А, flavipes
strongly favoured forest fragments over continu-
ous Forest and usually was captured within 30 m
of forest edges (Laurance 1994b). The Atherton
(A. godmani; Laurance, 1993) and brown an-
techinuses (А. stuartii adustus), both of which are
strongly rainforest-dependent (Laurance,
1994b), also could potentially occur at Eacham
and Barrine. Ideally, both remnants should be
thoroughly sampled during the optimal trapping
period for antechinuses.

The rare spotted-tailed quoll Dasyurus
maculatus was not detected during the study and
is unlikely to persist in the parks, However, both
parks currently maintain good populations of the
musky rat-kangaroo Hypsiprimnadon
moschatus, a rainforest endemic. This ies
appears to have disappeared from even the largest
(up to 590ha) fragments on the southern Atherton
Tableland (Laurance, 1994b), although it was
detected in a few small fragments on the north-
central Tableland that nearly abutted continuous
forest (Grey, 1994). The musky rat-kangaroo is
strongly rainforest-dependent, thus populations
in the two parks may be genetically and demo-
graphically isolated from other such populations.

A recent initiative by the Queensland Depart-
ment of Environment and Heritage to establish an
SOm-wide faunal corridor between Lake Barrinc
and Gadgarra State Forest, may facilitate dis-
persal of musky rat-kangaroos and other forest-
dependent species, reducing the isolation of
faunal populations at Barrine, The effectiveness
of such corridors has not yet been rigorously
assessed (cf, Noss, 1987; Simberloff & Cox,
1987; Bennett, 1990), however, and future re-
search is clearly needed to test the efficacy and
design of faunal corridors, especially in the trop-
Ics.

ACKNOWLEDGEMENTS

We thank John Winter, Les Moore, Luke
Leung and Peter Green for commenting on earlier
drafts of the manuscript: Eleanor Collins, Arthur

601

Sellers and others from the Lake Eacham Na-
tional Park Volunteers Group assisted with field
work. This project was partially supported by a
senior research fellowship awarded to WFL bv
the Wet Tropics Management Authority.

LITERATURE CITED

ANDREN, Н. & ANGELSTAM, P. 1988. Elevated
predation rates as an edge effect in habitat islands:
experimental evidence, Ecology 69:544-547.

BENNETT, A. F. 1990. Habitat corridors: their role in
wildlife management and conservation. (Arthur
Rylah Institute: Heidelberg, Victoria).

GREY, А. 1994. Fragmentation: effects on the musky
rat kangaroo (H'ypsiprimnodon moschatus) on the
Atherton Tableland. Unpublished report. (School
for International Training: Caims, Queensland),

HARRINGTON. G.N., IRVINE, A.K., CROME,
F.HJ., MOORE, L.A. & SANDERSON, K.D. (in
press), Problems faced by large-seeded trees in
rainforest fragments. ‘Tropical forest remnants;
ecology, management and conservation of frag-
memed conununities’. (University of Chicago
Press: Chicago, Illinois),

KAPOS, V, 1989. Effects of isolation on the water status
at forest patches in the Brazilian Amazon, Journal
of Tropical Ecology 5;173-185,

LAURANCE, W. F. 1990. Comparative responses of
five arboreal marsupials to tropical forest frag-
mentation. Journal of Mammalogy 71:641-653.

199]a, Ecological correlates of extinction proneness
in Australian tropical rain forest mammals. Con-
servation Biology 5:79-89,

1991b. Edge effects in tropical forest fragments:
application of a model for the design of nature
reserves, Biological Conservation 57:205-219,

1992. Abundance estimates of small mammals in
Austrilian tropical rainforest: a comparison of
four trapping methods. Wildlife Research
19;651-655,

1993. The pre-European and present distributions of
Amechinas godmani (Marsupialia: Dasyuridac),
a restricted rainforest endemic. Australian Mam-
malngy 16:23-27.

19943. Hyper-disturbed parks: ecology and man-
agement of isolated rainforest reserves in tropical
Queensland. (Published abstract, Annual Mect-
ing of the Seciety for Conservation Biology,
Gaudalajara, Mexico).

1994b. Rainforest fragmentation and the structure
of small mammal communities in tropical
Queensland. Biological Conservation 69-23-32,

LAURANCE, W. F., GARESHE_J. & PAYNE, C. W.
1993. Avian nest predation in modified and natu-
ral habitats in y Queensland: an experimen-
lal study. Wildlife Research 20:711-723.

LAURANCE, W, F. & GRANT, J, D, 1994, Phato-
graphic identification of grownd-nest predators in

602

Australian tropical rainforest. Wildlife Research
21:241-248.

LEUNG, L. K. P., DICKMAN, C. R. & MOORE,L. A.
1993, Genetic variation in fragmented populations
of an Australian rainforest rodent Melomys
cervinipes. Pacific Conservation Biology 1:58-65.

LOVEJOY, T. E., BIERREGAARD, R. O., RY-
LANDS, A. B., et al. 1986. Edge and other effects
of isolation on Amazon forest fragments. Pp. 257-
285 in M. A. Soule (ed). ‘Conservation biology:
the science of scarcity and diversity'. (Sinauer:
Sunderland, Massachusetts).

MATTHEWS, C. 1993. Draft management plan for
Lake Eacham and Lake Barrlne National Parks.
(Department of Environment and Heritage:
Cairns, Queensland).

NOSS, R. F. 1987. Corridors in real landscapes: a reply
to Simberloff and Cox. Conservation Biology
1:159-164.

OSUNKOYA, O. O. 1994. Post-dispersal survivorship
of north Queensland rainforest seeds and fruits:

MEMOIRS OF THE QUEENSLAND MUSEUM

effects of forest habitat and species. Australian
Journal of Ecology 19:52-64.
SIMBERLOFF, D. S., & COX, J. 1983. Consequences
© and costs of conservation corridors. Conservation
Biology 1:63-71.

TAYLOR, J. M. & HORNER, B. E. 1973. Results of
the Archibald expedition 98: Systematics of native
Australian Rattus (Rodentia, Muridae). Bulletin of
the American Museum of Natural History 150:1-
130.

TERBORGH, J. 1992. Maintenance of diversity in trop-
ical forests. Biotropica 24:283-292.

TRACEY, J. G. 1982. ‘The vegetation of the humid
tropical region of north Queensland’, (CSIRO:
Melbourne).

WILLIAMS-LINERA, G. 1990. Vegetation structure
and environmental conditions of forest edges in
Panama. Journal of Ecology 78:356-373.

WINTER, J. W., BELL, F. C., PAHL, L. I. &
ATHERTON, R. G. 1987. Rainforest clearing in
northeastern Australia. Proceedings of the Royal
Society of Queensland 98:41-57.

PALORCHESTES SELESTIAE, A NEW SPECIES OF PALORCHESTID MARSUPIAL
FROM THE EARLY PLIOCENE BLUFF DOWNS LOCAL FAUNA, NORTHEASTERN

QUEENSLAND
BRIAN MACKNESS

Mackness, B.S, 1995 12 01, Palorchestes selestiae, a new species of palorchestid marsupial
from the early Pliocene Bluff Downs Local Fauna, northeastem Queensland. Memoirs of the
Queensland Museum 38(2): 603-609. ISSN 0079-8835

Palorchestes selestiae sp. nov. from the early Pliocene Bluff Downs Local Faunais described
on the basis of an isolated LM!, It is larger than P. painei and P. parvus but smaller than P.
агае in both length and width of lophs, It is also differentiable from all other palorchestids
on the basis of the combination of links between, or stemming from lophs. It is the second
undoubted Pliocene species and shares features of both P. parvus and the Pleistocene P.
azael, With its relatively derived anterior morphology and plesiomorphic posterior morphol-
ogy, Р, selestiae challenges current notions about the phylogenetic relationships of
diprotodontid marsupials which suggest that small size and simple structure are probably
pleisomorphic features. [ ] Palorchestes, palorchestids, Bluff Downs Local Fauna, Pliocene,
Diprotodontia, Marsupialia.

Brian Mackness, School of Biological Sciences, University of New South Wales, Р.О. Box
І, Kensington NSW 2033, Australia; і August 1995.

Palorchestes was described by Owen (1873) on
the basis of an anterior portion of a cranium
including the rostrum. The holotype was obtained
by Dr Ludwig Becker and described by Owen
(1874) as P. azael from an unspecified deposit in
Victoria interpreted by Mahoney & Ride (1975)
as the River Tambo in Gippsland. Owen
(1873:387) considered Palorchestes to be”... the
largest form of kangaroo hitherto found".

Owen regarded Palorchestes to be a
macropodid, a view followed by Simpson (1945)
who placed the genus within the subfamily

inae. Raven & Gregory (1946), how-
ever, placed jt in the subfamily Sthenurinae while
Tate (1948) placed it in a new subfamily of kan-
garoos, the Palorchestinae. Woods (1958) was
the first to recognise the diprotodontid rather than
macropodid affinities of Palorchestes, Archer &
Bartholomai (1978) followed this assignment but
elevated Tate's Palorchestinae to family level.

Palorchestes selestiae sp. nov. is described
here on the basis of an isolated LM! from the
fluviatile and lacustrine deposits of the Al-
lingham Formation, northwest of Charters Tow-
ers, northeastem Queensland. This formation
contains a diverse assemblage known as the Bluff
Downs Local Fauna, and has been the subject of
many studies, most recently including those of
Vickers-Rich (1991), Mackness et al. (1993),
Boles & Mackness (1994) and Mackness (in
press). Archer & Wade (1976) assigned an early
Pliocene age to the assemblage on the basis of the
intepreted age of the overlying Allensleigh Ba-

salt. Archer, in Archer & Wade (1976) noted, bul
did not name three other diprotodontoids from the
Bluff Downs Local Fauna, a species of Eu-
ryzygoma, a species of Zygomarurus and an un-
identified *notothere',

TERMINOLOGY

Dental homology of cheekteeth follows
Luckett (1993) such that the first adult molari-
form tooth is M! and the deciduous molariform
tooth is dP5. Terminology of crown morphology
(Fig. 1)followsthat used by Archer (1976) except
that the metaconule is used instead of hypocone
following Tedford & Woodburne (1987). The-
gotic terminology used follows Every (1972).
Abbreviations for specimen numbers: QMF,
Queensland Museum fossil collection; UCMP,
University of California, Berkeley; P, South Aus-
tralian Museum.

SYSTEMATICS

Order DIPROTODONTIA Owen, 1866
Suborder VOMBATIFORMES
Woodburne, 1984
Infraorder VOMBATOMORPHIA
Aplin & Archer, 1987
Family PALORCHESTIDAE (Tate, 1948)
Archer & Bartholomai (1978)
Palorchestes Owen. 1873

TYPE SPECIES
Palorchestes azael Owen 1873.

1, mel.

ті.

FIG. 1. Diagram showing terminology of an M! of P.
selestiae. Regions used for measurements in Table 1
are also presented using the holotype (QMF12455).
Arrow indicates anterior of molar. Abbreviations: ab.f
anterobuccal fossa; a.c. anterior cingulum; l.f. lingual
forelink; m: metaloph; mc. metacone; mel.
metaconule; ml. midlink; pa. paracone; p.c. posterior
cingulum; pl.f. posterolingual fassa; pr. pratocone,

Palorchestes selestiae п, sp.
(Fig. 2)

MATERIAL EXAMINED

HoLorYPE ОМЕ12455, an isolated LM', Main
Quarry (19° 43'8, 145° 36 E), Allingham Formation,
Bluff Downs Station, northeastem Queensland,

ETYMOLOGY
For Selesti Smith of Bluff Downs Station.

DIAGNOSIS

Small LM! with lingual forelink terminated in
the cingular valley and not in contact with the
cingulum; second incipient buttress on the back
of the metaloph; posterolingual fossa well devel-
oped; posterior cingulum rises to an apex mid-
length; incipient buttressing on the posterior wall
of metaloph; one midlink and two incipient mid-
links; double forelinks.

DESCRIPTION

The holotype consists of a complete, relatively
unwom LM!, lacking any remnant of the roots.
The metaconule is the tallest cusp with the pro-
tocone, paracone and metacone subequal in
height. Protocone linked to the paracone by a
well-defined sinuous protoloph. Metaconule
linked to the metacone by a well-defined crescen-
tic metaloph whose ends are posteriorly de-
flected,

MEMOIRS OF THE QUEENSLAND MUSEUM

The anterior cingulum fails to extend around
the base of the protocone. A slight anterobuccal
cingulum is continuous with the anterior cingu-
lum. Otherwise there are no lingual or buccal
cingula. Anterior cingulum high resulting in deep
fossae where the forelinks join the cingulum.
Posterior cingulum closer to the metaloph than
the anterior cingulum is to the protoloph. As a
result, the posterior cingular valleys are not as
deep as the anterior ones. There is a conspicuous
interproximal wear facet on the anterobuccal cor-
ner of the tooth, presumably caused by abrasion
against P3,

There are a nümber of blades on the crown.
Where the enamel has been breached, both sets
of leading and trailing blades (sensu Every, 1972)
can be seen on the protoloph and metaloph. A
number of secondary blades are also present. One
blade (the apical margin of the lingual forelink)
leads from the protocone io the anterior cingu-
lum. Another blade (the apical margin of the
buccal forelink) connects the protoloph, approx-
imately one third of the way between the
paracone and the protocone, to the anterior cin-
gulüm. A well-developed primary midlink joins
the protoloph to the metaloph. The posterior end
of this midlink contacts the trailing edge of the

TABLE 1. Comparative measurements pf Palorchestes
M! in mm.

|P.selestiae [Holotype — — 225 [166 169 |
|P. parvus

ОМЕ12476
| omma әз fns
| ü [OME |i93 fiso [12 |

QMF2967 jiga Jiss [iss |

| | o |gMEss X || Jus |
| мето ||
| —— paora ë ise jna (136 |
|. [gMrsx [258 [207 [97 |
|. [eso ——— [зз [sis fais |
Г am |жз [2258 mo |
| | [Ben — (261 |9 |219 |
| ОСМР70553(1) [168 | 13.7

| ducMP7som) he7 hisa [зт |
[— [ocres [v7.8 aor [132° |

* = approximation

NEW SPECIES OF PALORCHESTID MARSUPIAL

TABLE 2: Summary of links in Palorchestes M!
Q Incipient link; - Link/s worn, * cast.

Мо. — |Forelinks |Midlink | Hindlinks |

He

pem ы bt J
dign 7 CHONGNU Rand ШШЕ!

[| — owe [xy b p |
омет p ë h b |
[ — low |» — 2 | |
Totes eB ШЕШИП
[gama [owe 2 [im w |
[emer һ e
painei CMP70553

емее — ПИШЕ ПИШЕ

postenor metaloph blade. Another less developed
midlink occurs on the lingual side of the tooth:
This smaller link is steeply v-shaped rather than
shallowly v-shaped as is the primary midlink.
There is a very deep fossa between these two
midlinks

There is also a swollen buttress on the posterior
flank of the protoloph, buccal to the primary
midlink. Another steeply inclined buttress de-
scends posteriorly to the posterior cingulum from
the tip of the metaconule and supports an incipi-
ent blade that connects the apex of the
metaconule to the posterior cingulum. A smaller
blade-like structure is present on the posterior
wall of the metaloph, buccal to the primary pos-
terior blade, There is some slight crenulation of
the enamel at the base of the lophs.

n

N
ғә

кә
[5]

M

REMARKS

P. selestiae 1s approximately 3/4 the size of P.
azael Owen and is 1/5 larger than P. parvus De
Vis. Whilst closer in size to P. parvus, P. selestiae
is morphologically closer to P. azael. P. azael
differs in that the lingual forelink links the pro-
tocone to the anterior cingulum, and lacks the
second incipient buttress on the back of the
metaloph. P. selestiae has a better developed
posterolingual fossa and the apex midlength on
the cingulum is absent in P. azael. In P. parvus
the anterobuccal fossa is less developed, both
forelinks join the anterior cingulum, there are two
well-defined hindlinks which join directly with
the posterior cingulum, and two midlinks are
present. P. panei Woodburne has a definite mid-
link, absent in P. selestiae, and it lacks the double
forelink and the two incipient buttresses on the
posterior wall of the metaloph.

605

PHYLOGENETIC AFFINITES

The early Miocene Nagapkaldia tedfordi and
Pitikantia dailyi (Stirton, 1967) from the Tirari
Desert of South Australia have previously been
regarded as the pleisomorphic sister group (sub-
familiae Incertae) of the apparently more derived
Palorchestinae based on synapomorphies in the
auditory region and neurocranium (Stirton et al.,
1967). It has been further suggested by Stirton et
al. (1967:154) that Pitikantia" ., is much nearer,
if not in, an ancestral position to Palorchestes".
Archer (1984), however, observed that there was
little evidence to regard the two groups as mono-
phyletic, with species of Ngapakaldia and
Pitikantia lacking the specialized molars and re-
tracted nasal bones of palorchestines, even
though their basicranial morphology was similar.
Murray (1986) described the middle Miocene
Propalorchestes novaculacephalus from the Bul-
lock Creek Local Fauna and, based on its cranial
base, glenoid fossa morphology and auditory re-
gion, placed it within the subfamily Pal-
orchestinae, Subsequently, Murray (1990)
described the dentition of P. novaculacephalus as
well as a new taxon, P. ponticulus from D-site
(System A), Riversleigh, Queensland. The latter
was also placed within the Palorchestinae.

The M!, the most diagnostic tooth for pal-
orchestids (Woods, 1958; Woodburne, 1967), is
characterised by elaborated forc-, hind- and mid-
links. These features are lacking in species of
Ngapakaldia and Pitikantia but present or incip-
ient in Propalorchestes novaculacephalus, con-
sidered to be the sister group of Palorchestes
Murray (1986, 1990). Palorchestes painei has the
simplest system of links with one fore-, one hind,
and one midlink which 1s the condition also scen
in P. novaculacephalus. All other species of Pal-
orchestes have two forelinks as a synapomorphy.

FIG. 2. Palorchestes seiestiae holotype QMF12455.
Actual size. Occlusal view stereo pair. Anterior at lop,

E
B
=
E
$

22 24
TOOTH LENGTH

P. paínel P. selestiae
ANTERIOR WIDTH o o

POSTERIOR WIDTH * .

FIG. 3. Bivariate plots of tooth length against anterior width (") and tooth length

against posterior width M! of Palorchestes species.

Palorchestes selestiae and P. azael share acces-
sory midlinks and the reduction of hindlinks as
synapomorphies while P. parvus has two
hindlinks and two midlinks as autapomorphies.
The development of accessory hindlinks and the
lingual forelink not extending to the anterior cin-
gulum in P. selestiae are considered au-
tapomorphies for this taxon.

Palorchestes selestiae indicates need for cau-
tion in determining phylogenetic relationships
within diprotodontoid lineages based on size
alone. Stirton ef al. (1967) suggested that molar
size and complexity in diprotodontoids were in-

MEMOIRS OF THE QUEENSLAND MUSEUM

verse correlates of age. By
those measures, P. painei,
as the smallest species,
ought to be the oldest fol-
lowed, in decreasing age
and increasing size by P.
parvus, P. selestiae and P.
azael. However, P. selestiae
comes from a demonstrably
early Pliocene site, whereas
the smaller P. parvus comes
from early to middle
Pliocene sites which does
not support the
biostratigraphic argument
of Stirton er al. (1967).

Several authors (Archer,
1976b; Murray, 1990) have
proposed that the bilophod-
ont upper molars of species
of Palorchestes are deriv-
able from subselenodont an-
imals such as wynyardiids.
Murray (1990:49) suggests
"|. .The possibility of
paraphyletic bilophodonty
could not be ruled out, par-
ticularly with respect to the
diprotodontids". In this
view those animals such as
the late Miocene P. painei
which had the least develop-
ment of selenodont features
would be regarded as the
most derived. It would then
follow that, barring conver-
gence, the more elaborate
the cresting, the more
plesiomorphic the animal.
The seledont groundplan is
still evident in P. selestiae,
even though it has been ob-
scured to some degree by
the development of lophs.

PALAEOECOLOGY

Palorchestids are uncommon elements in most
Australian marsupial faunas although they have
a long Tertiary and Quaternary history (Murray,
1991). The oldest known representatives are spe-
cies of Ngapakaldia and Pitikantidia from the
Oligo-Miocene deposits of central Australia and
Riversleigh in northwestern Queensland. At the
younger end of their time range, some may be

NEW SPECIES OF PALORCHESTID MARSUPIAL

REDUCTION GF
HINDLINKS

DEVELOPMENT
OF MIDLINKS

FIG. 4. Cladogram of taxa based on M! morphology.

only 30 000 yBP (Murray, 1991) and it has been
suggested that Palerchestes was contemporane-
ous with humans. Murray & Chaloupka (1984)
have interpreted an Aboriginal rock painting ir
the Arnhem Land escarpment as a possible ren-
dition of a species of Palorchestes although this
has been questioned (Lewis, 1986; Chaloupka &
Murray, 1986; Mackness, 1992).

The reduction and retraction of the nasals.
along with a conspicuous fossae for large
nasomaxillolabialis muscles and a narrow, pro-
tracted rostrum have been used as a basis for
reconstructing palorchestids with a trunk and the
basis of their popular description as ‘marsupial
tapirs’ (Bartholomai, 1978; Flannery, 1983;
Flannery & Archer, 1985; Murray, 1991). The
long, narrow deeply grooved symphysis suggests
that palorchestids had long protrusible giraffe-
like tongues (Murray, 1991). The spatulate lower
incisors are almost parallel to strongly developed
diastemal crests and interpreted by Murray
(1991) as probably being used to strip leaves or
crop clumped vegetation. Little has been pub-
lished about the postcranials of palorchestids.
Woods (1958) cast doubt on the association of
postcranials with skull remains suggested by sev-
eral authors (Owen, 1876; Gregory, 1902;
Fletcher, 1945). Archer (1984), however, has
noted that Palorchestes has. powerful forelimbs

607

equipped with large laterally compressed claws.
Flannery & Archer (1985) have used postcranial
material and teeth to attempt a reconstruction of
two species. Their analyses suggested the possi-
bility that species of Palorchestes were medium-
sized folivores that may have used their powerful
claws and arms.to rip bark off trees for food or to
uproot shrubs with tuberous roots.

ACKNOWLEDGEMENTS

I thank Michael Archer and Suzanne Hand for
helpful comments on the manuscript. Sue
Creagh, Peter Murray, Neville Pledge and Ralph
Molnar for access to other palorchestid speci-
mens. Jack, Rhonda, Bram, Troy and Selesti
Smith of Bluff Downs Station continue to provide
tremendous support for this ongoing research. |
thank Henk Godthelp, Darren Sprott and Frank
Coffa for their assistance. Collection of the Bluff
Downs material was supported in part by an ARC
Program Grant to M. Archer; a grant from the
Department of Arts, Sport, the Environment,
Tourism and Territories to M. Archer, S. Hand
and Н. Godthelp: a grant from the National Estate
Progràm Grants Scheme to M. Archer and A.

Pleistocene

o
c
o
o
2
a

Miocene

FIG. 5. Distribution of species of Palorchestes.

Bartholomai; and grants in aid to the Riversleigh
Research Project from Wang Australia, ICT Aus-
tralia and the Australian Geographic Society,

LITERATURE CITED

APLIN, K, & ARCHER, А. 1987, Recent advances in
marsupial systematics with а new syneretic clas-
sification. Pp. xv-Ixxii in Archer, M. (еф) 'Pos-
sums and Opposums: Studies in Evolution,
(Surrey Beatty & Sons and the Royal Zoolagical
Society of New South Wales: Sydney)

ARCHER, M. 1976. Phascolarctid origins and the po-
lential of the selenodont molar in the evalution af
diprotodont marsupials. Memoirs of the Queens-
land Museum 17; 367-371.

1984. The Australian marsupial radiation, Pp, 633-
BOS, In Archer, M & Clayton, О, (eds.) “Verte-
brale Zoogeography and Evolution in
Australasia. (Hesperian Press: Carlisle).

ARCHER, M. & BARTHOLOMAL А. 1978, Tertiary
mammals of Australia: a synoptic review. А}-
cheringa 2: 1-19,

ARCHER, M., GODTHELP, H., HAND, SJ. &
MEGIRIAN, D. 1989. Fossil mammals of
Riversleigh, northwesiem Queensland: prelimi-
nary overview of hiostratigraphy, correlation and
environmental change. Australian Zoologist 25:
29-65.

ARCHER, М, & WADE, М, 1976. Resulls of the Ray
E. Lemley Expeditions, Part I. The Allingham
Formation and a new Pliocene vertebrae fauna
from narthem Australia. Memoirs of the Queens-
land Museum 17: 379-97,

BARTHOLOMAI, A, 1978. The rostram in Pal-
orchestes Owen (Marsuptalia: Diprotodontidac).
Results of the Ray E. Lemley expeditions, Part 3.
Memoirs of the Queensland Museum 18: 145-
149,

BOLES, W.E. & MACKNESS, B.S. 1994. Birds from
the Bluff Downs Local Fauna, Allingham Forma-
tion, Queensland. Records of the South Australian
Museum 27: 139-149.

CHALOUPKA, G. & MURRAY, P, 1986. Dreamtime
or reality? Reply to Lewis. Archaeology in Ocea-
nia 21: 145-147.

EVERY, R.G, 1972, А new terminology for mamma-
lian teeth: founded on the phenomenon of thego-
sis. (Pegasus Press Christcharch).

FLANNERY, T.F. 1983. A unique trunked giant. Pal-
orchestes azael. Pp. 54-55. In Quirk, S & Archer,
M. (eds. ) ‘Prehistoric Animals of Australia (Aus-
tralian Museum: Sydney).

FLANNERY, TF, & ARCHER, М. 1985. Palerchestes
Owen, 1874. Large and small palorchestids. Рр,
234-239, In Rich. РУ. &G. V. Tets "Kadimakara.
Extinct Vertebrates of Australia. (Pioneer Design
Studio: Lilydale),

FLETCHER, Н.О. 1945. Palorchestes - Australia's
aoe oa t A Australian Museum Мар-

MEMOIRS OF THE QUEENSLAND MUSEUM

GREGORY, J.W. 1902. Some remains of an extinct
kangaroo in the dune-rock of the Sorrento Penin-
sula, Victoria. Proceedings of the Royal Society
of Victoria 14: 139-144.

LEWIS, D. 1985. Comment ‘The Dreamtime Animals’;
a reply. Archaeology in Oceania 21: 140-14,
LUCKETT, P. 1993, An ontological assessment of

dental homologies in therian mammals, Pp. 182-
204. In Szlay, F,S., Novacek, М.Ј, & McKenna,
М.С. (eds.) “Mammal Phylogeny: Mesozoic dif-
ferentiation, multituberculates, monotremes, early
therians and marsupials (Springer-Verlag: New

York).

MACKNESS, B.S. 1992, Aboriginal animal motifs in
the fourth dimension. Australian dame PP
Society Conference, Canberra, September. (Ab-
stract)

IN PRESS, Anhinga malagurala, a new species of
darter from the early Pliocene Bluff Downs Local
Fauna, northeastern Queensland. Emu.

MACKNESS, B.S., MCNAMARA, G., MICHNA, P,
COLEMAN. 5. & GODTHELP, H, 1993, The
Sprang Park Local Fauna, a new late Tertiary fossil
assemblage from northern Australia. Conference
on Australasian Vertebrate Evolution, Palaeonta-
logy and Systematics. Adelaide, 19-21 April
1993, Programme and Abstracts,

MAHONEY, J.A. & RIDE, W.D.L. 1975. ‘Index to the
qus and species of fossil Mammalia described

Australia and New Guinea between 1838
and 1968'. Western Australian Museum Special
Publication 6; 1-250,

MURRAY, P. 1986, Propalorchestes
novaculacephalus gen, et sp. nov, a mew pal-
orchestid (Diprotodontoidca: Marsupiatiay from
ihe Middle Miocene Camfield Beds, Northern
"Territory, Australia. The Beagle, Occasional Pa-
pers of the Northern Territory Museum of Arts and
Sciences 3: 195-211.

1990. Primitive marsupial tapirs (Propalorchestes
novaculacephalus Murray and P. ponticulus sp.
nov.) from the mid-Miocene of North Australia
(Marsupialia: Palorchestidae). The Beagle, Occa-
sional Papers of the Northern Territory Museum
of Arts and Sciences 7: 39-51.

1991. The Pleistocene megafauna of Australia. Pp.
1071-1164. In Vickers-Rich, P., Baird, R.F.,
Monaghan, J. & Rich. T.H. (eds.) ‘Vertebrate
Palaeontology of Australasia (Pioneer Design
Studio and Monash University Publications
Committee: Melbourne).

MURRAY, P. & CHALOUPKA, G, 1984, The Dream-
time animals: extinct megafauna in Arnhem Land
rock art, Archaeology in Oceania 19: 105-116.

OWEN, В. 1873. On the fossil mammals of Australia.
Family Macropodidae. Genera Macropus,
Pachysiagon, Leposiagon, Procoptodon and Pal-
orchestes. Part IX. Proceedings of the Royal So-
ciety 21: 128

1874. On the fossil mammals of Australia, Part TX.
Family Macropodidae. Genera Macropus.

NEW SPECIES OF PALORCHESTID MARSUPIAL

Pachysiagon, Leptosiagon, Procoptodon and
Palorchestes.Philosophical Transactions of the
Royal Society of London 164: 783-803.

1876. On the fossil mammals of Australia. Part X.
Family Macropodidae: mandibular dentition and
parts of the skeleton of Palorchestes; additional
evidences of Macropus titan, Sthenurus, and
Procoptodon. Philosophical Transactions of the
Royal Society of London 166: 197-226.

1880. Description of a portion of mandible and teeth
of a large extinct kangaroo (Palorchestes crassus
Ow.) from ancient fluviatile drift, Queensland.
Transactions of the Zoological Society of London
11: 7-10.

RAVEN, H.C. & GREGORY, W.K. 1946. Adaptive
branching of the kangaroo family in relation to
habitat. American Museum Novitates 1309: 1-14.

SIMPSON, G.G. 1945. The principles of classification
and a classification of mammals. Bulletin of the
on Museum of Natural History History 85:
1-350.

STIRTON, R.A. 1967. The Diprotodontidae from the
Ngapakaldi Fauna, South Australia. Bureau of
Mineral Resources, Geology and Geophysics Bul-
letin 85: 1-44.

STIRTON, R.A., WOODBURNE, M.O. & PLANE,
M.D. 1967. A phylogeny of the Tertiary
Diprotodontidae and its significance in correla-
tion. Bureau of Mineral Resources, Geology and
Geophysics Bulletin 85: 149-160.

TATE, G.H.H. 1948. Results of the Archbold Expedi-
tions. No.59. On the anatomy and phylogeny of
the Macropodidae (Marsupialia). Bulletin of the
American Museum of Natural History 91: 233-
352.

TEDFORD, R.H. & WOODBURNE, M.O. 1987. The
Tlariidae, a new family of vombatiform marsupials
from Miocene strata of South Australia and an
evaluation of the homology of molar cusps in the
Diprotodontia. Pp. 401-418. In Archer, M. (ed.)
‘Possums and Opossums: Studies in Evolution.
Volume 1 (Surrey Beatty & Sons Pty Ltd and the
Royal Zoological Society of New South Wales:
Sydney).

VICKERS-RICH, P. 1991. The Mesozoic and Tertiary
history of birds on the Australian plate. Pp.722-
808. In Vickers-Rich, P., Baird, R.F., Monaghan,
J, & Rich, T.H. (eds.) ‘Vertebrate Palaeontology
of Australasia (Pioneer Design Studio and
Monash University Publications Committee:
Melbourne).

WOODBURNE, M.O. 1967, The Alcoota Fauna, cen-
tral Australia: an integrated palaeontological and
geological study. Bureau of Mineral Resources,
Geology and Geophysics Bulletin 87: 1-187.

WOODS, J.T. 1958. The extinct genus Palorchestes
Owen, Memoirs of the Queensland Museum
13(4): 177-193.

610

GEHYRA DUBIA (MACLEAY, 1877) CONFIRMED AS
SENIOR SYNONYM OF PEROCHIRUS MESTONI DE
VIS, 1890. Memoirs of the Queensland Museum 38(2):610.
1995:- The gecko Perochirus mestoni De Vis, 1890 was
described from a specimen collected from Bellenden Ker in
north-east Queensland. Following examination of the holo-
type (Queensland Museum QMJ236), the name was placed in
the synonymy of Gekyra variegata (Dumeril & Bibron, 1836)
by Kluge (1963), This synonymy was followed by Cov-
acevich (1971), Cogger et al. (1983) and Ingram (1990) ,

Recently, Bauer & Henle (1994) noted that the type local-
ity and re size of the hol: were more consistent with
Gehyra dubia (Macleay, 1877) than Gehyra variegata, and
tentatively synonymised P. mestoni with the former, though
recommending that the holotype be re-examined.

The holotype of P, mestoniis faded, dessiccated, contorted
and damaged, The left mandible and associated tissues are
missing, all toes and fingers other than the fourth toe on the
right pes are missing, and the body is broken almost into two,
probably by previous attempts at measurement of snout-vent
length.

Nonetheless, diagnostic features (King, 1985) of Gehyra
dubia are present on this specimen, a female with greatly
distended endolymphatic sacs. The rostra] scale is about 1,5
times wider than tall, gabled, and has at least three internasals
bordering its dorsal margin. The right postmental contacts
only the first infralabial, and is short, approximately twice as
long as broad. The fourth toe on the right pes has nine
expanded lamellae, the apicalmost divided, the remainder
deeply creased. The snout-vent length 3s not able to be accu-
rately measured, butis certainly greater than 50mm, Although
Bauer & Henle (1994) interpret De Vis’ (1890) statement
"length 106mm" as snout-vent length, comparison of
measurments of head and limb with the holotype indicates that
this is à total length measurement,

The pattern, though faded, consists of at least broken fine
dark vermiculations over the head and neck, consistent with
photographs of live G. dubia (Wilson & Knowles, 1988).

The type locality lies outside the known distribution of G.
variegata, but well within the known distribution of G, dubia
(Ingram & Raven, 1991).

Consequently, Perochirus mestoni is formally syn-
onymised with Gehyra dubia. This action requires no changes
to current nomenclature.

Kluge's erroneous synonymy is understandable given the
state of knowledge of Australian Gehyra systematics. At the
lime of his paper, G. variegata was differentinted from a
composite G, australis largely on the basis of division of the
distal subdigital lamellae. Additional characters distinguish-

MEMOIRS OF THE QUEENSLAND MUSEUM

ing variegata from the G. australis complex and recognition
of additional species in the latter group awaited the work of
Mitchell (1965) and King (1985).

Acknowledgements
1 thank Patrick Couper and Jeanette Covacevich for assis-
tance during my work at the Queensland Museum.

Literature Cited

Bauer, А.М. & Henle, К. 1994. Familia Gekkonidae (Reptilia,
Saura) Part I. Australia and Oceania. Das Tierreich
(109): 1-306.

Cogger, H.G., Cameron, E.E. & Cogger, H.M. 1983. 'Zoolog-
ical Catalogue of Australia. Vol. 1. Amphibia and
Reptilia’. (Australian Government Publishing Service:
Canberra). 313pp.

Covacevich, J. 1971. Amphibian and reptile type-specimens
in the Queensland Museum. Memoirs of the Queens-
land Museum 16: 49-67,

De Vis, C.W. 1890. Description of two lizards of genera new
to Australian herpetology. Proceedings of the Linnean
Society of New South Wales (2)4: 1034-1036.

Dumenl, A.M.C. & Bibron, G. 1836, Repetologie Generale
ou Histoire Naturelle Complate des Reptiles Vol. 3.
(Librairie Encyclopedique de Roret: Paris)

Ingram, GJ. 1990, The works of Charles Walter De Vis,
alias'Devis',alias "Thickthorn'. Memoirs of the
Queensland Museum 28: 1-34.

Ingram, G.J. & Raven, R.J. 1991. “An Atlas of Queensland's
Frogs, Reptiles, Birds & Mammals’. (Queensland Mu-
seum; Brisbane). 391рр.

King, M. 1985. The Gehyra australis species complex (Sau-
ria: Gekkonidae). Amphibia-Reptilia 4; 147-169,
Kluge, А.С. 1963, The systematic status of certain Australian
and New Guinean gekkonid lizards. Memoirs of the

Queensland Museum 14; 77-86.

Macleay, W. 1877. The lizards of the "Chevert" Expedition,
second paper, Proceedings of the Linnean Society of
New South Wales 2; 97-104,

Mitchell, F.J. 1965, Australian geckos assigned to the genus
Gehyra Gray (Reptilia, Gekkonidae). Senckenbergiana
Biologica 46; 287-319,

Wilson, S.K. & Knowles, D.G. 1988. "Australia's Reptiles. A
photographic reference to the terrestrial reptiles of
Australia’ (Collins; Sydney), 447pp,

Glenn М, Shea, Department of Veterinary Anatomy, Univer»
sity of Sydney, NSW 2006, Australia; 1 August 1995,

STATUS OF THE LATE PLEISTOCENE FOSSIL DARTER ANHINGA LATICEPS
(DE VIS, 1906)

BRIAN S, MACKNESS & GERRY F. VAN TETS

Mackness, B.S, & van Tets, G,F. 1995 12 01. Status of the late Pleistocene fossil darter
Anhinga laticeps (De Vis, 1906). Memoirs of the Queensland Museum 38(2): 611-614,
Brisbane, ISSN 0079-8835.

Plotus (=Anhinga) laticeps (De Vis. 1906), described from the late Pleistocene sediments of
Shine: mel neyaneCoopers Creek, was compared with 31 specimens efie ESCAR Ario pa melanie gave! чүт леру nf de тучи!
qal otn? fugenovaehallandiae. While A. laticeps has a large interórtital: Avidthand Wasefrontal ingas * Ene imt! nda
(0 17у з se pirHhonrelative to extant darters, these do not constitute sufficient diffésemies to justify issepratieme iisdem neret
wr уу! of 4 tas a species. Anhinga laticeps is, therefore, recogRised Sed Gunter eynonig irc of Ac de clon, venons s
^ лаада) Ave novaehollandiae. C) Anhinga laticeps, darter, taxoroWV/ PleisreéWe, Алта Ave, ces irony, emm

Seu) BAO, AD aer Brian Mackness, School of Biological Sciences, Universiry Df Neve Seul Valle; PO Bod Soo o UMa nn New
Mugan чу УИНН wreKensington, New South Wales 2033; Gerry Е. van ТЇ CSIKO Division bi УРЦ ала етту o m n, CH n
p, Ecology, РО Box 84, Lyneham, A.C.T. 2602 (deceased) ES Julti? Bus SA Coole C ONS due tenes

Mociviess CsCharles ЖААП Vis described numerous fossil Chevaehillakdives Misckness"(im^press] Has тезү coo ood
мү, ‘eel Бїт erwaen- 4885 and 1911 (van Tels & Rich? centIy described ио алге ге ерат got; Cen ood а
suns 6031999): 8o0me ofthese were based on a collection ^ Plióverie (Blufts Сенту Есау Füunaj thironly ices is Iu E
ber Sow Ümade'BysPrefes&or Gregory of over 200 smalh=other Extihetiomettberfyont thesAnhingidad redi 15 =а ны! sc
alh wiih pones fram the deposits around Lake Eyre (Detticoritedfrim usiitumie sire ус (Oe corded frum Ан!
Vis, 1906). A fossil cranium and partial реіуі =. 79067. A ose] cem une руа Sohu
ILS AND Мт Codpers Creek were described as a darter ш ocoWEATBRUAES-ANE-METHODS © ys FRAG
Platus (-Anhinga) laticeps (De Vis, 1906). Atos te ngo) latiemes 1 За We 1906) А
taal va museum label, in the handwriting of De Vis, als6/2^7 pe GAY) кые ass eld! НЫ the De Vie ope n
mo Pusea lista eeracoid-(INo. 70) on the same card as thes" EMI йзейп O Rëss Yens oF other teens Л
- \nares!'eatworformenspdcimens but this bone or bone frag-* o fers ver ime Atal Natal is = were isih
D Le ySoumentwas'ti¢t megtioned by De Vis in his origina = нас CSIRO Division df Wildlife anf olson CSU
Son ops eseniplien. порах it been seen or noted hy any ^ Epdlogy, On foai from ела А Com-co у, Олї funr
Se wits 21 subsequent nhie and so must, therefore, be preu оуу s Weve "nde With 31 specimens of. ihgansons wens чи

niiTanddios sused Insti! fossi Suife&tant A. пт, movaehollandige as well as all fossibt 270.4 Wye
> (hs 12x07. BecMis (1906-18), in his description of P. айай т Ito this taxén: T8xXenofic posia tes] ferred |

orld din crdditeepysstdted tat ". . . this cranium is in all боон ofthe Ога World darters’ folthws*Mayr dor ef the (Yd У
3 Mamban dini@asions, Srfiewhat larger than that of PivmCettée)r (19795 und Marchand Higgins (990p eu) (1979) un
27 y пао wevaeltallándiae;it prohibits us referring it to the dn^recegnisin рой Гу sone ре ое s Inga тосол ce
iminelogy zIsealtsspocios previously described P. рагу" айат ауы n eninelegy'ot bone sig from yarn parley Ta
o ASADA pap De Vis (1906) has since been lodyn Drieschit976). ABhréevibtians for specimeri-? ass | 5 71
Ure: MWewn.io OE" a Little Pied Cormorant cmimbers >AM, (Амека Miseumc^ АМУУ Ое АМ, /
= Waits ( ePbedravthelanoleucos (Miller, 1966). — ^-sAustraliauNatianal Wildlife Collectimn, CSIRO srs fen Nes iin
їс and Entong iett feviewed the cranium and pelvic ^IDivisipwen Waldhfearid-Ezalogy; MN Museum «ion if Wich
Quens а mie bed by De Vis and nominated thé esol Vidora Му Queensland) Museum (ROMY! V toi ОМ,
scm; ЗАМ The lectotype of Anhinga laticeps (ag cacRoyal Ontatio Museunt, SAM, South Australian ys Onamo. Wa
Ve үези, ЖБ hd not formally specify a type) апд“ Museum; ICMP, Unive sityrefuCalifomia Mus иН " 2 Mi

sa) Collse ылгый the pelvic fragment to the extant AeleseumPalcamelogicalCollectiane cxtpr А. 80Р сотик

melanogaster. novaehollandiae. While Miller !@inyexter novaenallasdiae. Worle Molle:
(1966) considered A. lariceps to be a valid spe- MIERSDWEMENTS. Шер to o5 д valid аре мїзлңшчнм КТ

їс» cies, Brodkerbs&)Mourer-Chauviré (1982) havéic*. punc d ipio "arr ЧЕП Реве Mcuroremem

3-0 Ve" singe oquestiened-this view. Miller (1966:31 EJ "«(1906j were made using vermercallpers абга! 9 15 were vel
тта sed aisorteviewediarnumber of Pleistocene darter sac) Уей "and Sürfimerised below: Siilishcal^ s and :
пазува отта! Australian deposits and сон analysis GP lhesémeaturenieants' is provided тагу я of locke
se modorelsdeditham they differ in no aspects of size ot Fables]. ARTAS ай motd Specimens used le T. oc foe
Wort саннар ст sesigaing all to the extant А, mi зб comparisons, were considered: Lo represent)! So s

TABLE 1. Measurement (mm) of cranium. Measure-
ments as defined below: range; mean I standard devi-
ation; standard error.coffiecient of variation (96); 9556
confidence interval of the mean.

s WT 8
49.6

| 47 431.24
сс. 001, 2.6

3713414
20.8-24.7
22,810.78
0.14, 34

22. 6-23. 1

‚20, 4.
19.3-20.1
i 5, I

fully grown or mature individuals, based on the

absence of the juvenile condition of a”. . pitted
surface of the bone and incomplete ossification
of the articular facets" (Campbell, 1979:17). The
measurements taken were as follows:

Cranium greatest length (CGL). Measured as
the greatest distance from the protuberantia oc-
cipitalis externa to the incisivium.

Cranium greatest width (CGW), Measured
across the linea nuchales superior.

Cranium orbital width (COW). Measured as the
smallest breadth between of the pars nasalis of
the frontale.

Cranium greatest postfrontal width (CPW).
Measured as the greatest breadth across proces-
sus postfrontales,

Cranium greatest depth (CGD). Measured from
the basitemporale in the median plane to the
highest and median point of the braincase.

Cranium incisivium width(CIW). Measured as
the greatest width at the base of the incisivium.

COMPARATIVE MATERIAL

Skeletons (catalogue number, sex, locality) of
Anhinga m. novaehollandiae are as follows:

Anhinga m. novaehollandiae AMO.62367 ?
Australia; AMO.65078 9 Magela Floodplain,

MEMDIRS OF THE QUEENSLAND MUSEUM

Northern Territory, AMO.65077 ? Magela
Floodplain, Northern Territory; AMO,65076 3
Magela Floodplain, Northern Territory;
АМО.65075 d Magela Floodplain, Northern
Territory; ANWC (PELS 38) d Papua New
Guinea; ANWC (PELS 316) ? Burrinjuck Dam,
New South Wales; ANWC (PELS 318) 9 New
South Wales; ANWC (PELS 319) ? Burrinjuck
Dam, New South Wales; ANWC (PELS 320) 2
Burrinjuck Dam, New South Wales; ANWC
(PELS 356) 2 Woolgarlo, Piney Ridge, New
South Wales; ANWC (PELS 37) & New South
Wales; MV W4754 ? Victoria; MV WS082 ?
Victoria; MV W5913 ? Victoria: MV W8972 ?
Victoria, MV W12746 ? Victoria; MV W13183
? Healesville Sanctuary, Victoria; MV B8674 $
Melbourne Zoo, Victoria; MV B8675 ? Reedy
Lakes, Kerang, Victoria, MV B11664 & Reedy
Lakes, Kerang, Victoria; MY B16242 2 Top
Marsh, Kerang, Victoria; MV B17254 ? Lake
Mokoan, Victoria; MV B17255 F Lake Mokoan,
Victoria; MV B17595 ? Lake Mokoan, Victoria:
MV B18970 ? Lake Mokoan, Victoria; QM
21032 ? Queensland; QM 21031 ? Queensland;
QM 20798 ? Queensland; ROM 157468 d Aus-
tralia; SAM 31636 ? Lashmars Lagoon, Kanga-
roo Island, South Australia.

RESULTS

All cranial measurements of A. laticeps, except
the cranium orbital width (COW) and the cra-
nium incisivium width (CIW), fall within the
observed range of those for extant darters (Table
1). There is a depression in the region of the pars
nasalis of the frontal bone which is clearly
illustrated in Plate V1 of De Vis’ original descrip-
tion and marked with а small "a" (De Vis, 1906).
It runs from the frontal region commencing be-
hind the processus postfrontalis and continues
through to the processus frontalis of the premax-
illa. This appears to have been a post-mortem
fracture as there is no sign of any bone regrowth.
The depression probably contributes to an artifi-
cial widening of the interorbital area through the
flexure of the orbital bones on either side to
accommodate the depressed bone piece. The
large cranium orbital width is, therefore, consid-
ered to be artefactual.

The cranium incisivium width of A, laticeps is
not significant at two degrees of freedom and,
therefore, not considered to be of taxonomic im-

riance. The cranium of A. laticeps lacks most
of the features on the dorsal side and retains just
a remnant of the basisphenoid rostrum. De Vis

STATUS OF ANHINGA LATICEPS

(1906:18) remarked that the "presphenoid ros-
trum is higher and much stronger than it is in the
recent bird." Comparison with the 3] specimens
of the living A. m. novaehollandiae has shown
this feature to be extremely variable and that A.
laticeps fits well within that variation. In all other
cranial features, А. laticeps compares very well
with the modem A, m, novaehollandiae,

COMPARISON WITH FOSSIL MATERIAL

Several specimens of fossil darters were col-
lected from the vicinity of the type locality and
identifed as A, laticeps by R.H. Tedford and his
team in the late 1950s. Some of these were re-
viewed by Miller (1966) and assigned to A. т.
novaehollandiae. A single vertebra and bone
fragment (UCMP 56351), a proximal end of an
ulna (UCMP 56319) and a proximal end of a
humerus (UCMP 94681) collected by R.H.
Tedford from Coopers Creek, all compare well
with extant A. m. novehollandiae in both size and
features (humerus greatest width: Aij.
novaehollandiae 22.8mm - 19.4mm (Mackness,
unpublished data); A. ?laticeps 20.2mm).

Other Anhinga specimens have been collected
from the Kalipiri Formation of Lake Kununka,
eastern Lake Eyre Basin. These were questioned
by Vickers-Rich (1991) as being of Pliocene age
but are clearly labelled by their collector К.Н,
Tedford as being from the Katipiri Sands, à
Pleistocene deposit. A vertebra (UCMP 112825)
is too wom for diagnosis but compares well with
the extant A. m. novehollandiae in size as does a
cervical vertebra (UCMP 56852). Two proximal
ends of humen (UCMP 60545, greatest width:
18.4mm; UCMP 56885 20.9mm) compare well
with extant А, m, novehollandiae in both size and
features, A distal end of an ulna (UCMP 60863)
from the Pleistocene beds of Warburton River in
South Australia is also regarded as inseparable
from the extant darter,

DISCUSSION

In his original description of A. Jaticeps, De Vis
(1906) compared the fossil with only one speci-
men of the extant darter Anhinga m.
novaehollandiae. Miller (1966), in his subse-
quent revision of Australian darters, used seven
specimens of А. mt novaehollundiae and two
specimens of A, anhinga. This study utilised an
examination by Mackness (unpublished data) of
56 darter skeletons (12 specimens of A. anliingu;
13 of A. melanogaster rufa and 31 of the extant

613

A. m. novaehollandiae) as well as fossil material
referred to this taxon. Because of the obvious
large size of A. laticeps, only A. m.
novaehollandiae, the largest of the extant darters,
was used in the statistical analysis.

Miller (1966:317) supported the retention of A,
laticeps as a valid species on the basis that "the
measurement of laticeps exceed the mean of the
modern material by more than three times lhe
standard deviations". With a much wider data sct,
only one of these mcasurments now falls within
this category and that measurement is suspect
owing to post-mortem fracturing. Several speci-
mens of extant darter fall outside two degrees of
freedom, particularly ANWC (PELS) 316, a large
female from Burrinjuck Dam in New South
Wales. It is clear that there is significant variation
within certain darter measurements and caulión
should be exercised in making taxonomic deci-
sions before first comparing any specimen wilh a
suitable data set encompassing such variation,

There is not more than one darter species oc-
curing in any one location anywhere in the world
within extant populations (Dorst & Mougin,
1979), nor is there any evidence of this in the
fossil record (Mackness, in press), With demon-
strable specimens of A. m. novaehollandiae from
the Pleistocene of Australia and а new species of
darter from the Pliocene (Mackness, in press), it
is unlikely that a second form of darter lived
during the Pleistocene, Even if the large interorbi-
tal width of A. laticeps proves to be not
artefactual, the continued recognition of this pal-
aeospecics cannot be justified, and its synonymy
with A, m. novaehollandiae is the best solution.

ACKNOWLEDGEMENTS

We thank Waller Boles, Australian Museum,
and Rory O'Brien, Museum of Victoria for their
gencrous support and assistance dunng Ihis
study. Michael Archer and Suzanne Hand pro-
vided helpful comments on the manuscript. Les
Christidis, Museum of Victoria; Matthew Shaw,
Queensland Museum; Richard Schodde and John
Wombey, CSIRO Division of Wildlife and Ecol-
ogy; AJ. Baker, Royal Ontario Museum: Phillipa
Horton, South Australian Museum, made com-
parative material available, This study was sup-
ported in part by an ARC Program Grant to М,
Archer; a grant from the Department of Ans,
Sport, the Environment, Tourism and Territories
to M. Archer, S. Hand and Н, Godthelp; a grani
from the National Estate Program Grants Scheme
to M. Archer and A, Bartholomai; and grants in

614

aid to the Riversleigh Research Project from
Wang Australia, ICI Australia and the Australian
Geographic Society.

LITERATURE CITED

BRODKORB, P. & MOURER-CHAUVIRÉ, C. 1982.
Fossil anhingas (Aves: Anhingidae) from Early
Man sites of Hadar and Omo (Ethiopia) and
Olduvai Gorge (Tanzania). Geobios 15: 505-515.

CAMPBELL, K.E., JR. 1979. The non-passerine
Pleistocene avifauna of the Talara Tar Seeps,
northwestern Peru. Royal Ontario Museum, Life
Sciences Contribution 118: 1-203.

DE VIS, C.W. 1906. A contribution to the knowledge
of the extinct avifauna of Australia. Annals of the
Queensland Museum. 6: 33-36.

DORST, J. & MOUGIN, J.L. 1979. Order
Pelecaniformes. Pp. 155-193. In Mayr, E. &
Cottrell, G.W. (eds.) *Check-list of birds of the
world. Vol 1. 2nd ed. (Museum of Comparative
Zoology: London).

MACKNESS, B.S. in press. Anhinga malagurala, a
new pygmy darter from the early Pliocene Bluff

MEMOIRS OF THE QUEENSLAND MUSEUM

Downs Local Fauna, northeastern Queensland,
with a review of the world anhingid fossil record.
Emu.

MARCHANT, S. & HIGGINS, P.J. (Coordinators).
1990. ‘Handbook of the Birds of Australia, New
Zealand and Antarctica. Vol. 1. (Oxford Univer-
sity Press: Melbourne)

MILLER, A.H. 1966. An evaluation of the fossil an-
hingas of Australia. The Condor 68: 315-320.

VAN TETS, G.F. & RICH, P.V. 1990. An evaluation
of De Vis’ fossil birds. Memoirs of the Queens-
land Museum 28: 165-168.

VICKERS-RICH, P. 1991. The Mesozoic and Tertiary
history of birds on the Australian plate. Pp. 722-
808. In Vickers-Rich, P., Baird, R.F., Monaghan,
J. & Rich, T.H. (eds.) *Vertebrate Palaeontology
of Australasia (Pioneer Design Studio and
Monash University Publications Committee:
Melbourne).

VAN DEN DREISCH, Р. 1976. A Guide to the Mea-
surement of Animal Bones from Archaeological
Sites. Peabody Museum of Archaeology and Eth-
nology, Harvard University, Peabody Museum
Bulletin 1: 1-137.

A BLUE WHALE BALAENOPTERA MUSCULUS (LINNAEUS, 1758) FROM ST
LAWRENCE, QUEENSLAND

ROBERT A. PATERSON & STEVE VAN DYCK

Paterson, R.A & Van Dyck, S.M. 1995 12 01: A blue whale Balaenoptera musculus
(Linnaeus, 1758) from St Lawrence, Queensland. Memoirs of the Queensland Museum

38(2):615-621. Brisbane. ISSN 0079-8835.

The decomposing carcass of a juvenile blue whale Balaenoptera musculus musculus
(Linnaeus, 1758) washed ashore at St Lawrence, Queensland in February 1994, Its skeleton
was retrieved together with some baleen. Comparison was made with its vertebral measure-
ments and those of a pygmy blue whale B. m. brevicauda. Pathological changes, possibly
post-traumatic, in the caudal vertebrae are described and the sparse records of blue whales
in Queensland waters are documented, [ ] Blue whales, skeletal and baleen descriptions,

Queensland.

R.A. Paterson & S.M. Van Dyck, Queensland Museum, PO Box 3300, South Brisbane,
Queensland, Australia 4101; 16 October 1995,

Inlate January 1994 the carcass of alarge whale
was found drifting in Broad Sound by Chris Mc-
Namara à local fisherman, It subsequently
washed ashore between Waverley and St Law-
rence Creeks 11km east of St Lawrence (22^19' S,
149*38'E). On 17 February Grahame Byron of

Queensland Department of Environment and
Heritage (Rockhampton) identified it as a large
rorqual measuring between 16.5-17.5m in length.
The uncertainty in measurement resulted from
advanced decomposition and the disposition of
the carcass (Fig. 1). Assisted by Peter Cross and

FIG. 1. Carcass of QMJM10415 photographed at low tide on 17 February 1994.

616

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 2. Carcass of QMJM10415 photographed at high tide on 23 February 1994,

Kel Fowler of Queensland Boating and Fisheries
Patrol (Mackay), we retrieved the available skel-
eta] material on 23-24 February and 8 March
1994. (Details of skeletal elements, considered to
have been lost prior to collection, will be given
later.) The discovery of some baleen 80m from
the carcass enabled its identification as a blue
whale. The location of the carcass presented con-
siderable logistic difficulties. They included a
tidal range of approximately 5.5m on an exposed
shore with a steep mud bank lined with living and
dead mangroves. Rapid tidal flow posed dangers
due to movement of the skull and mandible with
their attached soft tissues (Fig. 2), The journey to
the nearest boat ramp was 7km. During one jour-
ney, with the mandible lashed to the sides of the
5.2m boat and the skull being towed, the nasals
and premaxillae became detached and were lost.
Despite these and other difficulties the skeleton
was transported initially to St Lawrence before its
removal to Brisbane. Itis registered QMJM10415
in the mammal collection of the Queensland Mu-
seum and ts the first confirmed record of a blue
whale in that collection.

SKELETAL APPRAISAL

Yochem & Leatherwood (1985) consider that
blue whales are 6-7m long at birth whereas Slijper
(1962) considered that they are approximately
7.5m long at birth which, in the Southern Hemi-
sphere, occurs during May-June. He also noted
that weaning occurs at seven months when length
averages 16m. Accordingly, QMJM10415 may
have died about the time of weaning. The length
at physical maturity is 24.5-26.5m (Slijper, 1962)
whereas the pygmy form (B.m. brevicauda) de-
scribed by Ichihara (1966) reaches physical ma-
turity at 20-22m.

The skull of QMJM10415 measured 4.32m in
length and is shown, from its ventral aspect, in
Fig. 3. The left side of the mandible measured
4.34m in length and the right 4.36m. There were
fourteen pairs of ribs. The epiphyses of only the
first two cervical and last twelve caudal vertebrae
were attached to their centra. Fifty-nine vertebrae
were collected but there were four additional
pairs of central epiphyses making a vertebral
count of sixty-three. The appearance and size of
the additional epiphyses suggest that the four
missing vertebrae are D9, 10, 11 and L2, The

BLUE WHALE

FiG, 3. Skull of QMJM10415 from ventral aspect.

vertebral formula C7; D14; L15; Ca27=63 is
consistent with others reported for the species
although variations of 63-66 have been noted
(Tomilin, 1957;Ichihara, 1966; Omura et al.,
1970).

Pathological changes were seen in the spinous

processes and neural arches of Cal0-11 (Fig. 4).

Considerable hyperostosis was noted but there
was no involvement of the centra as seen in
spondylitis deformans, considered to be associ-
ated with advanced age in cetaceans (Slijper,
1936, Van Bree & Duguy, 1970; Lagier, 1977;
Omura, 1972 & 1975; Paterson, 1984; ; ), The

617

degree of hyperostosis, localisation to two verte-
brae and lack of associated bony destruction tend
to exclude metabolic bone disease, osteomyelitis
and recent trauma as causes of these changes.
However, trauma in the perinatal period is possi-
ble as the advanced hyperostosis is consistent
with a process present for months rather than
weeks. One of us (RAP) has observed hyperos-
totic changes localised to two vertebrae (Ca7-8)
in a juvenile (central epiphyses unfused) blue
whale skeleton in the British Museum of Natural
History. The registration number of that speci-
men is BMNH1865.8.23.1. Slijper (1936) noted
examples of similar pathology to that in JM10415
in juvenile cetaceans and considered that onset
prior to birth was possible, given the extent of the
changes.

Omura et al.(1970) demonstrated in tabular and
graphic form vertebral measurements of an
18.6m long pygmy blue whale captured at
42°08'5, 44'09'E in December 1966. Its skull
measured 4.86m in length. The epiphyses of all
vertebrae were fused to the centra. In Fig. 5 the
vertebral measurements of that specimen are
compared with those of QMJM10415 whose
maximum length was 17.5m. The vertebrae of the
pygmy blue whale are larger in all respects than
those of QMJM10415 with the exception of the
lengths of the centra, particularly in the caudal
region, This latter finding, admittedly in the con-
text of only two specimens of disparate physical
maturity, may be significant in that Ichihara
(1966) named the pygmy blue whale B. т.
brevicauda on the basis that its tail, on external
measurement, was relatively shorter than that of
B. т, musculus.

In addition to the missing vertebrae, discussed
above, some other bones were not retrieved and
may have washed away prior to our visits to the
site. They were the sternum, a stylo-hyal, an ulna
and some carpals and metacarpals.

BALEEN

The number of baleen plates reported for blue
whales 15 324+6 (Mackintosh & Wheeler, 1929).
A portion totalling 88 plates from the right baleen
row of QMJMIO0415 was recovered and one of
the larger plates 15 shown in Fig. 6. The lengths
of all plates are shown in Fig. 7 and the steep
decline in measurements suggests that the plates
were from the posterior aspect of the row.
Illustrations of complete baleen rows from a large
balaenopterid in the British Museum of Natural
History (Beddard, 1900) and humpback whales
(Megaptera novaeangliae) in the Queensland

618 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 4. Caudal vertebrae of QMJM 10415 in (A) lateral and (B) dorso-ventral projection. Hyperostoses are seen
on the spinous processes and neural arches of Ca 10 and 11, In the fresh specimen the pathological areas were
"bridged" by fibrous tissue.

Museum (Paterson & Van Dyck, 1991) The equivalent ratio in the largest recovered plate

demonstrate a much steeper decline in the lengths ,

of the posterior plates than the anterior plates. ЗОМ Epa ois 2:001, Аав the: plates АН
Ichihara (1966) measured length/breadth ratios ОМЈМ10415 may not include the largest, the

of the largest baleen plates in 77 specimens of B.

m. musculus and 118 B. m. brevicauda. The ratios

in the latter did not exceed 1.93 in any specimen. В. m. musculus.

ratio of 2.05 suggests that the plate is typical of

BLUE WHALE

HEIGHT, GREATEST

HEIGHT, CENTRUM

MEASUREMENTS IN MM

BREADTH. GREATEST

BREADTH, CENTRUM

619

e

IGHT, GREATEST

- е
HEIGHT, CENTRUM

BREADTH, CENTRUM

LENGTH, CENTRUM

SS a

FIG. 5. Comparative vertebral measurements of QMJM10415 (right) and pygmy blue whale (left), The latter

measured 18,6m in length and the former 17.5.

QUEENSLAND BLUE WHALE RECORDS

There is an earlier record of B. musculus in the
Queensland Museum (Paterson, 1986). The spec-
imen (J4807) is a small portion of baleen from a
whale which stranded in 1928 at Couti Uti
(22°20°S, 150"08'E) not far from the locality of
QMJ10415. However, re-examination of the
specimen indicates that it is from a fin whale, B.
physalus (Linnaeus, 1758). The Queensland Mu-
seum also has a photographic record of a large
rorqual, possibly a blue whale, which stranded at
O'Regan's Creek, Dundowran (25'18'S,
152'46'E) in the late 1930s. Eye witness accounts
indicated that the whale exceeded 22m in length
(M. Campbell, pers. comm.) and they are sup-
ported by the photographic evidence (Fig. 8).

Inthe period (1952-1962) of whale exploitation
from the shore-station at Tangalooma (27°11'S,
153'23 E) 6277 humpback whales were captured
and the commercial viability of the station de-
pended exclusively on that species
(Chittleborough, 1965). One blue whale measur-
ing 20.5m in length was captured during the 1954
season. In the station's terminal phase a fruitless

search for humpback whales was made at dis-
tances exceeding 60km from the coast. During
that off-shore search two blue whales were seen
(L. Nash, pers. comm.). Paterson et al. (1994)
assessed humpback whale population recovery
from Point Lookout (27°26’S, 153°33’E) during
June-October from 1978-1992 and saw only one
blue whale. Identification was made on the basis
of the extremely tall blow, which exceeded 10m
in height and is considered to be characteristic of
the species (Horwood, 1986). These isolated cap-
ture, sighting and stranding records suggest that
blue whales are infrequent visitors to Queensland
coastal waters whereas they are more frequently
sighted in the cooler waters of southern Australia
(Paterson, 1982) and stranding records are more
numerous from those regions (Dixon & Frigo,
1994: Kemper & Ling, 1991).

ACKNOWLEDGEMENTS
We owe particular thanks to Peter Cross and

Kel Fowler of QBFP. Without their generous
assistance and local knowledge the retrieval

620

FIG. 6. One ofthe largest baleen plates of QMJM10415.

would not have been possible. Grahame Byron of
QDEH performed the initial inspection and pro-

MEMOIRS OF THE QUEENSLAND MUSEUM

50

40

30

20

MEASUREMENTS IN CM

10

20 40 60 80
NUMBER OF BALEEN PLATES

100

FIG. 7. Graph demonstrating the length of the baleen
plates, considered to be from the right posterior row
of QMJM10415.

vided Fig. 1. Ben Ingram and Richard McTaggart
of the St Lawrence Police Station provided sup-
port and staff of the Broadsound Shire Council
and Queensland Railways assisted with transport
and loading. The hospitality of Ken Johnson and
family of the Sportsman's Arms Hotel at St Law-
rence was much appreciated, Staff at the Luggage
Point Wastewater Treatment Plant at Pinkenba
kindly allowed us to store the skeleton there
during cleaning and ensured its safety. A gener-
ous donation from the Australian Whale Conser-
vation Society defrayed steam-cleaning costs.
Bruce Cowell of the Queensland Museum pre-
pared the photographs and Sue Gray prepared the
figures.

LITERATURE CITED

BEDDARD, F.E. 1900. ‘А book of whales’ (John Mur-
ray: London).

CHITTLEBOROUGH, К.С. 1965, Dynamics of two
populations of the humpback whale, Megaptera
novaeangliae (Borowski). Australian Journal of
Marine and Freshwater Research 16:33-128.

DIXON, J.M. & FRIGO, L. 1994. "The cetacean collec-
tion of the Museum of Victoria. An illustrated
catalogue’. (Australian Deer Research Founda-
tion: Croydon, Victoria).

HORWOOD, J.W. 1986. The distribtuion of the south-
em blue whale in relation to recent estimates of
abundance. Scientific Reports of the Whales Re-
search Institute, Tokyo 37:155-165.

ICHIHARA, T. 1966. The pygmy blue whale,
Balaenoptera musculus brevicauda, a new sub-

BLUE WHALE

621

FIG, 8. Skeleton of large rorqual, possibly a blue whale, at Dundowran in the 1930s. The skull and mandible are
situated in an inverted position, to the right of the photograph.

species from the Antarctic. Pp.79-113. In Norris,
K.S. (ed), “Whales, dolphins, and porpoises’ (Uni-
versity of California Press: Berkeley & Los Ange-
les).

KEMPER, C.M. & LING, J.K. 1991. Whale strandings
in South Australia (1881-1989). Transactions of
the Royal Society of South Australia 115(1):37-
52.

LAGIER, К, 1977, Spondylosis in the whale - an an-
atomicroradiological study of an osleophyte.
Scandanavian Journal of Rheumatology 6:245-
249.

MACKINTOSH, М.А, & WHEELER, J.F.G. 1929,
Southem blue and fin whales. Discovery Reports
1:257-540.

OMURA, H. 1972. An osteological study of the
Cuvier's beaked whale, Ziphius cavirostris, in the
northwest Pacific. Scientific Reports of the
Whales Research Institute, Tokyo 24:1-34.

1975. Osteological study of the minke whale from
the Antarctic, Scientific Reports of the Whales
Research Institute, Tokyo 27:1-36.

OMURA, H., ICHIHARA, T. & KASUYA, T. 1970,
Osteology of pygmy blue whale with additional
information on external and other characteristics.
Scientific Reports of the Whales Research Insti-
tute, Tokyo 22:1-27.

PATERSON, R. 1982. ‘Whale watch’ sighungs in-
crease again. Australian Fisheries 41(5):25-29.

PATERSON, R.A. 1984. Spondylitis deformans in a
Bryde's whale (Balaenoptera edeni Anderson)
stranded on the southern coast of Queensland.
Journal of Wildlife Diseases 20(3):250-252.

1986. A list of specimens of the Order Cetacea in
the Queensland Museum. Memoirs of thc
Queensland Museum 22(2):309-311.

PATERSON, R., PATERSON, P. & CATO, D.H. 1994.
The status of humpback whales Megaptera
novaeangliae in east Australia thirty years after
whaling. Biological Conservation 70:135-142.

PATERSON, R.A. & VAN DYCK, S, 1991. Studies of
two humpback whales, Megaptera novaeangliae,
stranded at Fraser Island, Queensland. Memoirs of
the Queensland Museum 30(2):343-350.

SLUPER, E.J. 1936. Die cetacean vergleichend - an-
atomisch und systematisch. Capita Zoologica
7(2):305-590.

1962. ‘Whales’ (Hutchinson: London).

TOMILIN, А.С. 1957. Mammals of the U.S,S.R. and
adjacent countries: Cetacea, Israel Program for
Scientific Translations, Jerusalem. 7 17рр.

VAN BREE, P.J.H. & DUGUY, R. 1970. Sur quelques
aberrations pathologiques chez les petits Cétacés.
Zoologische Garten 39(1/6):11-15.

YOCHEM, P.K. & LEATHERWOOD, S. 1985. Blue
Whale Balaenoptera musculus (Linnaeus, 1758),
Pp.193-240. In Ridgway, S.H. & Harrison,
R.(eds), ‘Handbook of Marine Mammals’ Vol.3,
(Academic Press: London & New York.)

622 MEMOIRS OF THE QUEENSLAND MUSEUM

FIRST RECORD OF PALAEMON CONCINNUS DANA,
1852 (CRUSTACEA: DECAPODA: PALAEMONIDAE)
FROM AUSTRALIA, Memoirs of the Queensland Museum
38(2):622. 1995:- Palaemon concinnus 3s widely distributed
in the Indo-West Pacific region but has not been previously
recorded from Australia, despite being known from
neighbouring areas such as New Guinea (Holthuis, 1982) and
New Caledonia (Holthuis, 1970). Four specimens collected
from Palm Island, northeast Queensland by Mr Hamar Mid-
gley in 1977, were found in the unidentified collections of the
Queensland Museum. Further material was discovered at the
Northern Fisheries Centre, Cairns, after an invitation by Mr
Warren Lee Long (Queensland Department of Primary Indus-
tries) to examine and identify Crustacea collected during their
Johnstone River Catchment Survey. Abbreviations: QM,
Queensland Museum; QDPI, Queensland Department of Pri-
mary Industries; NEQ, northeast Queensland. All measure-
ments are carapace length from the orbit to the posterior
margin.

Palaemon concinnus Dana, 1852
Restricted synonymy
Palaemon concinnus Dana, 1852; 26.
Paluemon exilimanus Dana, 1852: 26.
Leander longicarpus Stimpson, 1860: 40.
Palaemon lagdoensis Blanco, 1939: 167, pl.l.
Palaemon (Palaemon) concinnus; Holthuis, 1950: 61, fig.12.
Palaemon concinnus: Chace & Bruce, 1993; 40,

Material Examined

QMW 14793, 29 9 (12.6, 12.8mm), Palm Is., airport
swamp, МЕО, July 1977, S.H. Midgley; QMW16427, 15
(9.2mm), 12 (13.6mm), Ibid.; QMW18730, 18 (19.9mm),
19 (6.4mm), Bamboo Ck, Innisfail, NEQ, freshwater tidal
reach, silt, fringing rainforest, water clarity high, Vallisneria,
Blyxa, Aponogeton, fallen timber, leaf litter, DO? 60% satu-
ration, 1m depth, electrofished, 1992, QDPI Northern Fisher-
ies Centre, Johnstone R, Catchment Survey; QMW 19615, 1 9
(8.1mm), unnamed tributary of Johnstone R., МЕО,
17°29.0°S, 146°01.9'E, freshwater, lotic, water temperature
28.2°C, DO2 87.095, turbidity 2.0 NTU, electrofished,
10/4/1992, QDPI Northern Fisheries Centre, Johnstone Ё.
Catchment Survey,

Remarks

Male specimens examined have a rudimentary appendix
interna on the endopod of the first male pleopod, a feature that
distinguishes P. concinnus from other species of the genus.
Other diagnostic features such as the rostrum and antennular
flagellae also agree well with published descriptions and

figures. Bruce (1987) provided a key to the six species of
Palaemon previously recorded from Australia - P. debilis
Dana, 1852; P. litoreus (McCulloch, 1909); P. macradactylus
Rathbun, 1902; P, semmelinkii (de Man, 1881); P. serenus
(Heller, 1862); and Р. serrifer (Stimpson, 1860).

This species is recorded in the literature from salt, brackish
and freshwater, The Australian records are from coastal low-
land. fresh waters in the wet tropical region of northeast
Queensland.

Literature Cited

Blanco, G.J. 1939. Two new decapods from the Philippines.
The Philippine Joumal of Science 69(2): 167-171,
plates 1,2.

Bruce, A.J. 1987. Records of three Palaemonid shrimps new
to the Australian fauna, The Beagle, Records of the
Ur АШ Territory Museum of Arts and Sciences 4(1):

-60.

Chace, F.A, & Bruce, A.J. 1993. The Caridean shrimps
(Crustacea: Decapoda) of the Albatross Philippine Ex-
pedition, 1907-1910, Part 6; Superfamily Pal-
aemonoidea, Smithsonian Contributions to Zoology
543: 1-152.

Dana, J.D. 1852. Conspectus Crustaceorum etc., Conspectus
of the Crustacea of the Exploring Expedition under
Captain C. Wilkes, U.S.N. Proceedings of the Academy
of Natural Sciences of Philadelphia 6: 10-28.

Holthuis, L.B. 1950. The Decapoda of the Siboga Expedition.
Рап X. the Palaemonidae. Siboga-Expeditie. Leiden
39(а9): 1-268.

1970. Etudes hydrobiologiques сп Nouvelle-Calédonie
(Mission 1965 du Premier Instit de Zoologie de
l'Université de Vienne), (Suite) EX. The freshwater
shrimps (Crustacea. Decapoda. Natantia) of New Cal-
edonia. Cahiers ORSTOM 3(2): 87-105, figs1-4.

1982. Freshwater Crustacea Decapoda of New Guinea, in,
Gressitt, J.L., “Biogeography and ecology of New
Guinea’, Monographiae Biologicae. Vol.42, (Dr W.
Junk: The Hague).

Stimpson, W. 1860. Crustacea Macrura. Pars VIII of Pro-
dromus descriptionis animalium evertebratorum, quae
їп Expeditione ad Oceanum Pacificum
Septentrionalem, a Republica Fedorata missa,
Cadwaladaro Ringgold et Johanne Rodgers Ducibus,
observavit et descripsit. Proceedings of the Academy
of Natural Sciences of Philadelphia 1860; 22-47,
printed 1861,

John W. Short, Queensland Museum, P.O. Box 3300, Sourh
Brisbane, Queensland 4101, Australia; 7 November 1995

THREE NEW MYGALOMORPH SPIDER GENERA FROM THAILAND AND CHINA

(ARAENAE)
ROBERT J. RAVEN AND PETER J. SCHWENDINGER

Raven, R.J. & Schwendinger, P.J. 1995 12 01: Three new mygalomorph spider genera from
Thailand and China (Aranac), Memoirs of the Queensland Museum 38(2). 623-641. Bris-
bane, ISSN 0079-8835.

Three new mygalomorph genera and four new species are described: Angka gen. nov, Angka
hexops sp. NOV., à cyrtaucheniid from Thailand; nemesiids, Sinopesa gen. nov., Sinupesa
maculata sp, nov., from Thailand and Sinopesa guangxi sp. nov. from China; and a diplurid,
Lepiothele gen. nov., L. bencha sp. nov., trom Thailand. Generic limits, cladistic assessments.
and systematic placements within the 3 families are discussed, notes on natural history are
given. A character initially considered diagnostic in Phyxioschema suthepium Raven &
Schwendinger is shown to be variable, [ ] Mygalomorphae, Angka, Sinopesa, Leptothele ,
China, Thailand.

Robert John Raven, Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101,
Australia; Peter J. Schwendinger, Institute of Zoology & Limnology, University of Innsbruck,

Technikerstr, 25, A-6020 Innsbruck, Austria;

In biodiversity studies, it is customary to com-
pare faunal diversities of regions and countries
with one another. Judged by an early catalogue of
spiders (e.g. Roewer, 1942), the mygalomorph
fauna of Thailand has been poorly represented (2
theraphosids and 1 atypid). The reasons are his-
torical. Early arachnologists focused their col-
lecting efforts on British and other European
colonies. The protracted absence of harsh western
techniques for land use no doubt ensured that the
biodiversity of Thailand was not so heavily af-
fected until World War II but equally it was little
known. The prolonged absence of a country so
rich and diverse as Thailand from phylogenetic
and biogeographic studies of spiders has led to
significant weaknesses in such analyses. Wide
discontinuities in both form and space invite ex-
planations of rapid evolution in the known fauna,
and the intrusion of major barriers and/or cata-
clysmic extinction. The cffect, however, is prob-
ably no more detrimental than inadequately
funded taxonomy in key regions like India, and
was succinctly demonstrated in the New Zealand
area by Sphenodon (Daugherty et al, 1990)
where subspecies became extinct because of the
limited taxonomic study on the genus.

This paper is one of several (by PJS with
others) on the spiders of the rich forests of Thai-
land and highlights these points. The
Aganippinae were thought to be highly
specialised endemic Australian taxa (e.g. Main,
1981) with the closest relatives being the Indian
genysine genus Scalidognathus Karsch, 1878
(Raven, 1985), The description of Prothemenops
Schwendinger, 1991 from Thailand removed the

28 August 1995.

mystery of that uniqueness and quickly bridged
the gap. Here we address further exciting new
species of initially uncertain taxonomic status
discovered in Thailand. Relationships became
clear by searching for sister taxa, For each species
а new genus is established; in one, a new species
from China is also included. The new taxa show
new relationships among southeast Asian and
Australian mygalomorphs,

MATERIALS AND METHODS

Abbreviations are standard for the Araneae and
follow Raven (1994). Number of structures on the
opposite body side are given in parentheses. Mea-
surements, if not otherwise indicated, are in mm;
total length includes chelicerae. Colour is de-
scribed from alcohol preserved specimens unless
noted. Museum Acronyms: MCZ, Museum of
Comparative Zoology, Harvard University, Cam-
bridge, Massachussets; MHNG, Muséum
d'Histojre naturelle, Genève; NHMW,
Naturhistorisches Museum, Wien; QM, Queens-
land Museum, Brisbane, Collectors: AP, A.
Payne; PIS, P.J. Schwendinger.

SYSTEMATICS
Family CYRTAUCHENIIDAE Simon

RELATIONSHIPS.

This group was given family status first by
Raven (1985) who radically restructured the
Mygalomorphae, Only Goloboff (1993) has since
addressed the weaknesses of the family tenuously

624

constituted by Raven (1985). Goloboff's fine
analysis used a rigorous method and exhaustive
tested alternatives. Goloboff found only some of
Raven's groups could be supported but that
"Raven (1985) was impressively close to an op-
timal solution". However, Goloboff (1993) con-
cluded that "the present results were too
preliminary to warrant nomenclatural changes".
Hence, the classification of Raven is followed
here.

Angka gen. nov.

TYPE SPECIES
Angka hexops sp. nov.

DIAGNOSIS

Closely related to Kiama Main & Mascord,
1969 but only 6 eyes (Fig. 6I), a narrow fovea
(Fig. 6A) and much smaller PMS (Fig. 6K).

Angka hexops .
Sinopesa maculata

Leptothele bencha

MEMOIRS OF THE QUEENSLAND MUSEUM

Maxillae narrow, ventral surface not distended.
Labium short; posterior sternal sigilla small, mar-
ginal. Legs armed with spines, also on dorsal
tibiae and metatarsi of posterior legs. Legs of 9
scopulate. Transverse ridges on trichobothrial
bases confined to upper ledge (Fig. 4A). Tarsal
organ low. Female with 2 rows of teeth on all
paired claws (Fig. 3A, D). PMS very small. Meta-
tarsi of d leg I modified (Fig. 6H). Male palp with
dorsal spines on tibia and tarsus; bulb with cork-
screw-shaped embolus (Fig. 6C, D).

DESCRIPTION

Carapace wide, glabrous; cephalic part wide in
front and moderately arched (Fig. 6A, F, J).
Group of 6 eyes, not on tubercle, occupies 1/3-1/4
of head width at that point, no PME. Fovea
slightly procurved, deep, transverse, narrow.
Chelicera robust, with teeth only on promargin of
fang groove; rastellum absent. Maxillae moder-
ately wide, with flat ventral surface and few spin-

CHINA

3
* Sinopesa guangxi

Localities

1 Doi Inthanon

2 Khao Phanom Bench National Park
3 Guangxi Province

FIG. 1. Occurrence of Angka hexops sp. nov., Leptorhele bencha sp. nov., Sinopesa maculata sp. nov. in Thailand

and 5. guangxi sp. nov. in China.

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

FIG. 2, A-C. Habitus. A, B, Angka hexops sp. nov. A, juvenile 3. B, ?. C, Leprothele bencha sp. nov., ©.

dle-shaped cuspules on probasal corner. Labium
wide, short, no cuspules. Sternum almost as wide
as long; labiosternal suture a broad groove; all
sternal sigilla small, marginal. Legs long, slender;
cuticle of distal leg segments with fine reticulated
texture (Fig. 5B). Strong spines ventrally on all
tibiae and metatarsi and on dorsal side of legs III,
IV and on palpal tibia and tarsus. Leg tarsi as-
pinose. Scopula light on leg tarsi and distal meta-
tarsi of I, II. No preening combs. Trichobothrial
collar with transverse ridges only along upper
ledge (Fig. 5B). Tarsal organ low, with detached
low mounds nearby (Fig. 4A). Paired tarsal claws
with 2 rows of several long juxtaposed cylindrical
teeth, less numerous and long on III, IV (Fig. 3A,
D). Unpaired claw bare, short and hooked. Abdo-
men pale, no pattern. Booklung apertures with
many setae on tissue just inside entrances. PMS
very small; PLS long, apical joint digitiform.
Pumkiniform spigots absent.

Male: strong spines but no coupling spur on
tibia I (Fig. 6G); metatarsus I sigmoid, aspinose
and narrowed in its basal half (Fig. 6B, G); bulb
pyriform, with long slender corkscrew-shaped
embolus (Fig. 6C, D). No intercheliceral tumes-
cence,

Vulva with 2 spermathecae, each divided into
two anterior lobes, the outer ones branching from
the distal portion of the spermatheca and bent
towards the anterior (Fig. 6L).

ETYMOLOGY

From the type locality, Doi Angka Luang, the
old name of Doi Inthanon; ang ka Thai, meaning
crow’s bowl. (Doi = mountain, luang = large,
great).

DISTRIBUTION
As for species.

INCLUDED SPECIES
Angka hexops sp. nov.

RELATIONSHIPS

The search for the sister group of Angka begins
by eliminating alternatives. Several
mygalomorph families have strong au-
tapomorphies not present in Angka. Two possible
families with close affinities are Nemesiidae and
Cyrtaucheniidae. The arched cephalic area and
reduced eye tubercle clearly associate Angka with
the Cyrtaucheniidae. No nemesiids with such
deep and rounded maxillae are known. The sim-
ilarly elevated tarsal organs of Kiama (see Raven,
1981b and Fig. 5C) and Angka (Fig. 4A), the
elevated caput and sessile eyes, the wide shape of
the maxillae, and leg spination indicate they are
sister groups. Differences between the two are not
phylogenetically contradictory. The strongly pro-
curved fovea of Kiama is plesiomorphic for the
Rastelloidina (Raven, 1985) and its large sternal
sigilla are simply autapomorphic. In both genera.
the spermathecae are bilobed “1 + 1" (see Raven.
1985), d lack a tibial spur, and the spiders are
light coloured, not dark as for many burrowing
mygalomorphs.

BIOGEOGRAPHY

Although Angka and the Australian Kiama are
considered sister groups, the suggestion that their
occurrence in Australia arose by invasions
through New Guinea (e.g. according to Main,
1981, 1982, Queensland was invaded by idiopids,

626 MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 3. A-F, Tarsal claws, lateral views. A, D, Angka hexops sp. nov. A, G, leg LD, 9, leg IV. B, C, Sinopesa.
B, S. maculata sp. nov., d, leg IV. C, S. guangxi sp. nov., d, leg IV. E, F, Leptothele bencha sp. nov., 9, leg
I. E, all claws. Е, unpaired claw.

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

D s xi qv

627

FIG. 4. A-E, tarsal organ, and trichobothrial base, dorsal views. A, Angka hexops sp. nov., d , leg I. B-D. Sinopesa.
B, C, S. guangxi sp. nov., d, leg IV, B, trichobothrium and base, C, tarsal organ. D, S. maculata sp. nov., d,

leg IV. E, Leptothele bencha sp. nov., 9, leg I.

theraphosids, and some barychelids) lacks merit.
The taxonomy, distributions, and family place-
ments of mygalomorphs in Main (1982) were
tentative and some were confused. Hexathelidae
(notably the Atracinae) are not known from New
Guinea. Main (1982) noted doubts in locality
records of the Chevert Expedition (Rainbow,
1920). In fact, the combination of mygalomorph
genera listed from Bradley's Chevert material is
exactly that of his home suburb in Sydney, Aus-
tralia.

Mygalomorph genera known from New
Guinea are fromBarychelidae (Nihoa,
Monodontium; Raven, 1994), Ctenizidae (Con-
othele; Main, 1982), Dipluridae (Masteria Koch,
1873; Raven, 1979) or Theraphosidae
(Selenocosmia, Phlogiellus, Coremiocnemis;
Queensland Museum records). A similarly botan-
ically diverse region as New Guinea is the Wet

Tropics World Heritage Area (WHA), in north-
eastern Australia. Queensland Museum database
records (unpublished) list 9 mygalomorph fami-
lies and 24 genera from the Wet Tropics WHA:
Actinopodidae (Missulena), Barychelidae (Jdi-
octis, Idiommata, Mandjelia, Moruga, Ozicrypta,
Trittame, Tungari, Zophorame), Ctenizidae
(Conothele), Dipluridae (Cethegus, Masteria,
Namirea), Hexathelidae (Hadronyche), Idi-
opidae (Cataxia, Homogona, Misgolas), Migidae
(Migas), Nemesiidae (Aname, Chenistonia,
Namea) and Theraphosidae (Phlogiellus,
Selenocosmia, Selenotypus). Hence, the known
mygalomorph fauna of New Guinea can be re-
garded as depauperate. Also, the terrestrial com-
ponent of New Guinea is geologically young,
having been pushed up in the Oligocene by the
collision of the Australian and Asian plates
(Raven & Axelrod, 1972). The three most diverse

628

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 5. A-G, trichobothrial base and cuticle, oblique dorsal view. A, C, Kiama lachrymoides Main & Mascord,
д. A, trichobothrium & base . C, w ith tarsal organ & cuticle . B , Angka hexops sp. nov., д , leg I. D, E, Acontius
species, d, D, leg I, trichobothrial base, dorsal view. E, claws, lateral view. F, С, Leptothele bencha sp. nov.,
F, serrula, axial view. G, maxilla, ventral view.

mygalomorph families in Australia (Idiopidae,
Nemesiidae, Barychelidae) are either absent (Idi-
opidae, Nemesiidae) or far less diverse
(Barychelidae: 3 genera vs 10 in Eastern Aus-
tralia). Significantly, Idiopidae and Nemesiidae
occur west of Wallace's Line but not east until

Australia. Equally, few Australian
mygalomorphs show affinities with Oriental taxa
indicative of dispersal. Large Australian the-
raphosids of the genera Selenocosmia and
Phlogiellus are widespread through northern
Australia and are found also in New Guinea, parts

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

of the Pacific and southeast Asia. The enigmatic
minute litter mygalomorph Masteria is known
from the Philippines, through the Pacific (north-
em Australia, New Guinea, Fiji, and New Cale-
donia) to South and Central America (Raven,
1979, 1991). The group probably predates the
breakup of Gondwanaland. The ctenizid Con-
othele occurs through the western Pacific and
much of northern Australia. In the Australian
Barychelidae (Raven, 1994), only the Indo-Pa-
cific genera Sason and Jdioctis (Raven, 1986,
1988) occur on both sides of Wallace's line,

One alternative for this widely split distribution
of the Cyrtaucheniidae lies in India's northward
raft carrying part of the fauna including the àn-
cestral sister group of both Kiama and Angka.
After the separation, each genus diverged.

Angka hexops sp. йоу,
(Figs 1, 2A, В, ЗА, D, 4A, 5B, 6)

MATERIAL EXAMINED
НоготуғЕ: QMS4167, d, Doi Inthanon, 18733'N,
98728'E, 2530m altitude, Chiang Mai Province, Thai-
land, 18 Apr-23 May 1987, PJS.
PARATYPES: QMS29274, same dala as holotype: 9
dd, 1 9. 1 penultimate d, 13 Jan 1993, MHNG,
NHMW,
ETYMOLOGY

Greek, hexa (six), and ops (eye).

DiAGNOSIS
As for genus.

DESCRIPTION

Holotype male QMS4167: Total length, 12.5.
Carapace 5.2 long, 4.9 wide. Abdomen 5.6 long,
3,7 wide.

Carapace orange brown, chelicerae, palps and
leg I red brown, other legs orange brown. Abdo-
men entirely pallid yellow brown, posterior lung
covers with tiny dark patches at bases of setae.

Carapace broad with many long brown bristles
on lateral and posterior margins, posterior inter-
strial ridges, in foveal region, along anteromedial
ridge, on clypeus, between PME, and on caput
behind eye group. Pars cephalica broad, arched.
Fovea deep, slightly procurved, 0.94 wide, occu-
pies ca. 1/5 of carapace width at that point. Clyp-
eus very narrow, 0.25 wide.

Eyes six, PME absent, lateral eyes evanescent,
unpigmented; tubercle absent, AME on common
mound. Group 0.42 long, front width 1.07, hack
width 1.10, occupies ea. 1/3 of headwidth. Sizes
and interdistances; AME 0.20, ALE 0.2), PME

629

absent, PLE 0.14; AME-AME 0.15, AME-ALE
0.08, PLE-PLE 0.77, ALE-PLE 0.08.

Chelicerae porrect, broad, with broad band of
stiff black bristles dorsally and narrower band of
shorter bristles laterally. Intercheliceral tumes-
cence and rastellum absent; rastellar region with
fine bristies only. Furrow promargin with 12(13)
teeth, hasomesally with 3 tiny denticles. Maxillae
broad, 1,64 long in front, 2.40 behind, 1.20 wide,
with 21-25 cuspules in small prolateral-proximal
corner; heel rounded; antenor lobe indistinct.
Serrula region with small low scales, Labium
0.60 long, 1.00 wide: no cuspules.

Sternum 2.60 long. 2.60 wide; sigilla distinct,
oval, marginal. Labiosternal suture, two tear-
shaped sigilla just touching medially.

Leg formula 4123. Eighi thorn spines on ventral
and prolateral tibia 1. Leg 1 much thicker than
ПТУ, Metatarsus I slightly sigmoid. Preening
combs absent. Anterior leg tarsi slightly spindle-
shaped, all aspinose, Scopula light on tarsi and
distal metatarsi of legs 1, if Spines; T, fe p1, d1w,
ра 0, ti p2, v4 + 2 megaspines, me 0; П, fe p3.
dlw, pa 0, ti p3, v3, me pl, v7; III, fe all weak,
p2, dl, rI, pa pl, ti p2, di, 12, v7, me p4, r4, v6;
IV, fedl, rl, parl, ti p3, 12, v7. me p4 13 v7; palp,
fe pl, pa 0, ti p3 v6, cymbium 6. Trichobothria
on legs I-IV and palp: tarsi 8-10 in zig-zag row,
metatarsi [0-11 in one straight row, tibiae 6-9 in
Two proximally divergent rows. Paired claws with
two rows of 8-11 teeth; unpaired claws bare,

Leg and palp measurements:

I п П IV Palp
Femur 475 425 375 468 306
Patella 28] 231 212 212 1.75
Tibia 3.94 331 243 3.50 2.18
Metatarsus 3.31 2.94 318 450 -
Tarsus 187 168 168 L87 0.96
Total 1668 1449 13.16 16,67 6.03

Palpal tarsus spinose, wilh two rounded lobes.
Bulb pyriform, embolus long and slender, with
corkscrew-shaped tip.

PMS small, 0.27 long, 0.13 wide, PLS 2.1 long:
apical segment digitiform, 0.87 and 0.45 mid-
width, median 0.50, basal 0.77 long.

Pararvpe female QMS29274: Total length, in-
cluding chelicera 15, Carapace 4,9 long, 4.0 wide.
Abdomen 7.4 long, 4.7 wide. Asin б except:

Body generally pale yellow brown, chelicera
and sternum slightly darker.

Carapace glabrous with distinct black bristles
along margins and on caput. Caput more dis-
tinctly raised than in d , highest well behind eyes.
Fovea slightly procurved, 0.9 wide, occupies 1/4
of carapace width at that point.

630

MEMOIRS OF THE QUEENSLAND MUSEUM

Fic. б. A-L. Angka hexops sp. nov. A-I, д. J-L, 9. A, body (no legs), dorsal view. B, tibia and metatarsus, I,
ventral right, ventral view. C, D, palpal tibia, tarsus and bulb. C, ventral view. D, prolateral view. E, chelicerae,
sternum, maxillae and labium, ventral view. F, carapace, lateral view. G, leg I, ventral right, prolateral view. H,
metatarsus [, dorsal view. I, eyes dorsal view. J, carapace, lateral view. K, spinnerets, ventral view. L,
spermathecae, dorsal view. Scale lines = 1.0mm (А-К), 0.1 mm (L).

Eye group 0.26 long, front width 0.85, back
width 0.84, occupies ca. 1/4 of headwidth at that
point. Only AME distinct; others small, translu-
cent. Sizes and interdistances: AME 0.14, ALE
0.14, PME 0.04, very indistinct and present only

on left side, PLE 0.06; AME-AME 0.09, AME-
ALE 0.08, PLE-PLE 0.69, ALE-PLE 0.05.
Chelicerae strong, 2.3 long, no rastellum; fur-
row with 13 teeth on promargin and 2(3)
basomesal denticles. Maxillae 1.2 wide, front
length 1.6, back length 2.1, with ca. 30 cuspules

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

on distinct mound on prolateral-proximal corner.
Labium 0.5 long, 1.0 wide; no cuspules. Sternum
2.2 long, 2.2 wide. Posterior sigilla distinct, oval,
0.25 long; others indistinct, submarginal.

Leg and palp measurements:
I Ii Ш ІУ Palp

Femur 3.5 3.1 2.8 3.7 3.0
Patella 2.] 1.9 17 19 L6
Tibia 23 2.2 1,9 2.6 1.8
Metatarsus 1.9 19 23 3.2 -

Tarsus 14 14 14 1.7 17
Total 116 105 101 131 8.1

Leg formula 4123. All tarsi apinose, Scopula
light on tarsi and distal metatarsi of leg I, II.
Spines: 1, ti v3(4), me v6; II, ti v4, p1(2), me v7,
pl; Ш, ti v4(5), r1(2), p2(3), dl, me v7, 13, p3,
dl; IV, ti v6 (weak), rl, pl, dl, me v7, r3, p2(3),
d]; palp, ti v6, ta v3. Trichobothria on legs I-IV
(palp): tarsi 10-12 (7-8) in a zig-zag row, meta-
tarsi 9-11 in one straight row, tibiae 7-11 (7) in
two proximally divergent rows. Claws with two
rows of 6-8 cylindrical teeth, confined to basal
portion of paired claws; unpaired claws bare.
Palpal claw with prolateral row of 6 teeth.

PMS 0.3 long, PLS 1.9 long: apical segment
digitiform, 0,6, median 0.5, basal 0.8 long.

Spermathecae two, each divided into short
inner lobe and longer, bent outer lobe.

VARIATION

Carapace lengths of 5. 4.8-6.1 (n=10). Some
d have one PLE reduced. The juvenile 2 shows
small pigmentation where rhe left PME would be
situated, In the mature $ , a rediiced cornea of the
PME occurs on the left side.

DISTRIBUTION, HABITAT & BURROW

Known only from cloud forest near the summit
of Doi Inthanon (2565m), Chiang Mai Province,
Thailand, Fernale and juveniles were found under
rotten logs on the forest floor; d were trapped in
pitfalls. In captivity, the mature 2 built a бст
long burrow without any silk lining. The only silk
observed was a thin web spun onto the substratum
before the burrow was made. No other specimens
were observed to produce silk or construct a
burrow. Two possibilities emerge: the spiders
may inhabit temporary retreats and possibly for-
age on the ground. Vagrant females, however.
were not caught in any of the 5 pitfalls at the
locality during the 14 month study. Or, as in the
case of Xamiatus rubrifrons Raven, 1981, only
smal] amounts of web may be used to line the
burrow (Raven, 1981b),

631

Family NEMESIIDAE Simon
Subfamily ANAMINAE Simón

Sinopesa gen. nov,

ТҮРЕ SPECIES
Sinopesa maculata sp. nov.

DIAGNOSIS

Sinopesa is close to Entypesa Simon, 1902, but
differs in lacking posterior median spinnerets
(Figs 7J, 8B) and a serrula; in d, the palpal tarsus
1s short (Fig. 7D-F), metatarsus I modified (Figs
IK; 8E), and the intercheliceral tumescence is
absent.

DESCRIPTION

Carapace glabrous, narrow in front, caput a
little higher than thoracic region in d. (Fig. 7B),
clearly arched and higher in 9 (Fig. 7C). Eight
eyes on low but distinct tubercle, Fovea deep,
very narrow, transverse and slightly recurved.
Chelicera weak, with teeth only on promargin of
fang furrow. Intercheliceral tamescence and
rastellum absent. Maxillae long, narrow, with
distinct posterior heel and few cuspules on pro-
basal comer, not on mound; по sertula. Labioster-
nal suture a broad groove, Stemum wide, with
small, marginal sigilla (Figs 7M, 8D). Legs long,
slender. strongly armed with spines on patellae to
metatarsi, except on dorsal side of legs I, П. Leg
tarsi aspinose. Scopula weak on tarsi L Ш and
distal metatarsi. No preening combs. Paired
claws with 2 rows of several long, cylindrical
teeth; unpaired claw bare, quite Jong and little
bent. Tarsal organ low (Fig. 4B-D). Trichobothr-
ial base with transverse ridges confined to upper
ledge (Fig. 4B), Leg cuticle with fine scaly sculp-
ture (Fig. 4C, D). Abdomen with or without pat-
tern. PMS absent; PLS long, apical article
digitiform (Figs 7J, 8A, B). Males with large
spines but no spur on tarsus J; metatarsus I sig-
moid, narrowed in its basal half, with one dis-
toventral spine (Figs TH, K, L, SE). 3 palp with
spines on tibia and dorsal tarsus. Embolus pyri-
form, with long, slender, more or less corkscrew-
shaped embolus tip (Figs 7D-F, 8C).

ETYMOLOGY
The generic name is an arbitrary combination
of letters. The gender is feminine.

INCLUDED SPECIES
Sinopesa maculata sp. nov.
Sinopesa guangxi Sp. nov.

632

MEMOIRS OF THE QUEENSLAND MUSEUM

Fic. 7. A-M. Sinopesa maculata sp. nov., C, G, І, J, 9; rest, б. A, body no legs, dorsal view. B, C, carapace,
lateral view, B, d. C, 9. D, palpal tibia, tarsus and bulb. D, E, retrolateral view, F, prolateral view. G,
spermathecae, dorsal view. Н, leg I, retrolateral view. 1, 1, abdomen, 9. I, dorsal, J, ventral. К, metatarsus 1, д,
dorsal view. L, tibia and metatarsus I, retroventral view. M, chelicerae, stemum, maxillae and labium, ventral

view. Seale lines = 1.0mm, except G, 0.1mm.

Sinopesa maculata sp. nov.
(Figs 1, 3B, 4D, 7)

MATERIAL EXAMINED

HoLorvPE QMS6207, d, Doi Inthanon, 18°33'N,
98°28'Е, 2300m altitude, Chiang Mai Province, Thai-
land, 14 Jul-20 Aug 1987; PJS.

PARATYPES: QMS6208, 1 9,23 May 1987; 3 d d, 14
Jul-20 Aug 1987 & 20 Aug -8 Oct 1987, PJS, MHNG,
NHMW.

DIAGNOSIS

Males of S. maculata are much smaller than
those of 5, guangxi sp. nov., have a dark pattern
dorsally and ventrally and the embolus tip is
distinctly corkscrew-shaped.

ETYMOLOGY
Latin, maculatus, spotted.

DESCRIPTION

Holotype male QMS6207: Total length, 7.7.
Carapace 3.4 long, 3.0 wide. Abdomen 3.4 long,
2.3 wide.

Prosoma, legs and palps brown; abdomen light
brown, with dark dorsal and ventral (less pro-
nounced) pattern and dark brown posterior lung
covers (Fig. 7J).

Caput low, narrow. Fovea short, slightly re-
curved, 0.25 wide, occupies c. 1/12 of carapace
width at that point (Fig. 7A).

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

Eye group on distinct tubercle, 0.31 long, front
width 0.61, back width 0.63, occupies slightly
less than 1/2 of head width. Sizes and interdistan-
ces: AME 0.12, ALE 0.18, PME 0.12, PLE 0.14;
AME-AME 0.08, AME-ALE 0.02, PME-PME
0.21, PME-PLE 0.02, ALE-PLE 0.04. MOQ 0.27
long, front width 0.26, back width 0.41.

Chelicerae 0.7 long, no rastellum; furrow with
9(10) teeth on promargin and 5 tiny basomesal
denticles. Maxillae 0.6 wide, front length 1.0,
back length 1.3, with 14 cuspules at prolateral-
proximal comer. Labium 0.2 long, 0,6 wide, no
cuspules. Sternum 1.7 long, 1.6 wide. Labioster-
nal suture a broad groove.

Leg and palp measurements:

I H ш IV

Femur 2.7 23 2.1 2.8 1.7
Patella 1.7 14 13 m 1,0
Tibia 2.1 17 1.4 2.2 1.3
Metatarsus 1.8 17 2.1 3.0 -

"Tarsus 1.1 1.2 13 4 0.6
Total 94 8.3 $2 108 46

ventral tibia I (Fig. 7H, L). Metatarsus I sigmoid,
with distinct prolateral flattening distally (Fig.
7K). Preening combs absent, Tarsi aspinose.
Scopula very light on metatarsi and tarsi 1, IT.
Spines: І, fe pl, d3, rl, pa 0, ti p3, v5, me p2; II,
fe p3, 44, pa pl. ti p3, v6, me pl, v6; III, fe p3,
d3, 12, pa p4 r2, ti p3, d2, 13, v8, me p4, r4, v8;
ТУ, fe p2, d3, rl, pa p4, rl, ti p3, 42, r3, v7, me
p3, r3, v9; palp, fe pl, 04, pa 0, ti p3, ta б apical.
Trichobothria on legs I-IV (palp): tarsi 7-8 (6) in
a zig-zag row, metatarsi 7-10 in onc straight row,
tibiae 5-7 (6) in two proximally divergent rows.

Paired claws with teeth in two rows of 4-7;
unpaired bare.

Palpal cymbium spinose, with two rounded
lobes. Bulb pyriform; embolus long, thin, with
spiral tip (Fig. 7D-F).

PMS absent; PLS 1.6 long: apical segment dig-
itiform, 0.5, median 0.4, basal 0.5 long.

Paratype female QMS6208: Total length, in-
cluding chelicera 6.6. Carapace 2.3 long, 1.9
wide. Abdomen 3.1 long. 1.9 wide, As for б
except:

Body generally pale brown, chelicera slighily
darker. Abdominal pattern less evident.

Caput more distinctly raised than in d (Fig.
7С). Fovea straight, 0.3 wide, occupies cu. 1/5 of
carapace width at that point.

Eye group 0.27 long, front width 0.48, back
width 0,48, occupies almost half of head width at
that point. Sizes and inlerdistances: AME 0.08,
ALE 0.14, PME 0.10, PLE 0.14; AME-AME

633

0.08, AME-ALE 0.02, PME-PME 0.16, PME-
PLE 0.02. ALE-PLE 0.03.

Chelicerae weak, 0.6 long, no rastellum; furrow
with 8 teeth on promargin and 7 tiny basomesal
denticles. Maxillae quite narrow, 0.5 wide, front
length 0.7, back length 1.0, with 8(9) cuspules at
prolateral-proximal comer, Labium 0.2 long, 0.5
wide, no cuspules. Sternum 1.1 long, 1.1 wide.

Leg and palp measurements:

I

1! Hi IV Palp
Femur 1.6 1.3 1.2 1.6 1.3
Patella 11 0,9 0.8 10 Q8
Tibia 1.1 0,9 0,8 1.2 0.8
Metatarsus 0,8 0.3 1.0 1.6 -
Tarsus 0.6 0.7 07 1.0 0.8
Total 5.2 4.6 4.5 6.4 33

Leg formula 4123. Tarsi aspinose. Scopula
very light on tarsi of palp and leg I. Spines: 1, fe
d5 (long stout bristles); II, fe d3 (long stout bris-
tles), me v6; III, fe d3 (long stout bristles), pa p2,
T3, ti v7, p2, d4, me v7, p2, d5; IV, fe d3 (long
stout bristles), pa p2, r3, ti v7, p2, r2, dl, me v7,
p2, 95; palp, fe d2 (long stout bristles), ti v2, ta
v2 + several short stout bristles. Trichobothria on
legs I-IV (palp): tarsi 5-8 (6) in a zigzag row,
metatarsi 6-8 in one straight row, tibiae 5-6 (S)in
two proximally divergent rows. Paired claws with
2 rows of 4-6 cylindrical teeth, confined to hasal
half of claw on posterior legs; unpaired claw long,
only slightly bent. Palpal claw with prolateral row
of 4 teeth.

PMS absent. PLS 1.3 long: apical segment dig-
itiform, 0.4, median 0.4, basal 0.5 long.

Spermathecae two, triangular, each divided
into short pointed inner lobe and longer knoh-
shaped outer lobe bent forward.

VARIATION
Carapace lengths in @3.2-3.6, width 2.7-3.1
(n24).

DISTRIBUTION & HABITAT

Known only from Doi Inthanon, Chiang Mai
Province, Thailand at 2000-2300m altitude.
Males were trapped in pitfalls and juveniles were
found in leaf litter and under rotten wood on thc
forest floor. The young 9 kept in captivity con-
structed no web or burrow,

Sinopesa guangxi sp. nov.
(Figs 1, 3C, 4B, D, 8А-Е)

MATERIAL EXAMINED
HoLotyre:; MCZ, 3 , Dragon Lake, Liuzhou Guangxi,
China, Jan 1982, AP.

wH MEMOIRS OF THE QUEENSLAND MUSEUM

Fic. 8. A-E. Sinopese guangxi sp
Abdomen, ventral view, C, Palpa
ventral view, E, libia & metatarsus 1, proventral view.

. NOV., holotype g, A, Carapace, chelicerae & abdomen, dorsal view. B,
tibia, tarsus & bulb, ventral view, D, Sternum, maxillae, labium & chelicerae,

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

ETYMOLOGY
For the type locality.

DIAGNOSIS

Larger than S. maculata; lacking abdominal
pattern, posterior sigilla larger, and embolus tip
only slightly corkscrew-shaped. 2 unknown.

DESCRIPTION
Holotype male: Total length, 12. Carapace 5.31
long, 4.81 wide. Abdomen 5.13 long. 3.00 wide.
Carapace orange brown, legs orange to red-
brown. Abdomen dorsally and ventrally pallid
yellow-brown, no pattern (Fig. 8A).

Cephalic bristles short ой caput and interstrial
ridges; long on lateral margins; 4 long on clypeal
edge; many anteromedially. Fovea short, shal-
low, slightly recurved (Fig. 8A). Clypeus absent.
Caput elevation low, thorax sloping. Striae dis-
linet,

Eye tubercle low, with almost sessile eyes save
AME. Group occupies 2/5 of head-width. Two
rows: front row 34, back row 85. Eye group front
width, back width, length, 32:34:15. MOQ front
width, back width, length, 14:22:13.
AME:ALE:PME:PLE, 5:11:6:7. AME-AME,
0.6; AME-ALE, 0.3; PME-PLE, 0.2; PME- PME,
2; ALE-PLE, 0.2.

Chelicerae narrow, porrect, with long bristles.
Rastellum absent. Fang smooth. Furrow pro-
margin with 9 teeth, plus 1 small tooth, intermedi-
ally with 6-7 fine basally; several deep parallel
ndges near intercheliceral tumescence area but
not pallid; no teeth on outer margin of furrow.

Maxillae 1.30 long in front, 1.92 long behind,
0.84 wide; with c. 3-4 cuspules along edge. Heel
produced over labium; anterior lobes just evident.
Serrula absent. Labium 0,82 wide, 0.42 long.
Labiosternal suture with 2 double, pallid, lateral
areas, not grooved. Labium lacks cuspules; ante-
rior edge indented. Sternum 2.28 long, 2.13 wide;
distinct bristles along posterior edge. Sigilla all
marginal, oval. Sigilla length, distance to margin:
posterior 0.4, 0,16; middle 0.22, 0.16; anterior
0.19, 0.16, No prolateral coxal bristles; inner
coxae not produced or angular.

Leg and palp measurements:

1

M Ш IV. Palp
Femur 375 344 313 400 25
Parella 231 206 1.69 200 1.5
Tibia 294 262 238 3.50 1.78
Metatarsus 2.69 2,50 3,00 431 -
Tarsus 1.56 1.63 1.56 1.81 072
Total 13.25 12.25 11.75 15.63 6.50

Leg formula 4123. Setation light brown, ad-
pressed. Tibia 1 darker and thicker than other

635

tibiae with ventral spines thicker and 3 on distal
portion very raised, bases all movable. Metatar-
sus I curved for two-thirds of proximal length
(Fig. 8E). No preening combs. No scopula on
metatarsi; very thin and divided on tarsi I, absent
on I-IV. Spines: J, fe pl, d3, rl, pa 0, ti p3, v5,
me v1; ТЇ, fe pl, d2, pa pl. ti p3, v6, me v6: HI,
fe p3, d3, r2, pa p2, r1, ti p3, d2, r3, v7, me рб, r3,
v7; IV, fe pl, d4, r1, pa p2, r1, ti p3, d3, r3, уб,
me pó, r3, v7; palp, fe pl, d3, r1, pa pl, ti p3. ri,
v6, ta 10 distally. Paired claws with 2 rows of
teeth, 6-7 per row juxtaposed (Т), longer on IV
than TIT; on TV, teeth basal on outer face and distal
on the inner face; unpaired claw on I hooked,
bare, on IV longer, bare. Trichobothria in two
rows, each of 11 on tibiae; full curving row of
10-15 on metatarsi: 15 in 2 bands for three quar-
ters of length on tarsi.

Palpal bulb rugose, pyriform; embolus small
with distally hooked tip. Tarsus short, with large
spines similarly and deeply incised; no scopula.
Tibia not incrassate; distal ventrally excavated
(Fig. 8C).

PMS absent, without vestige. Length of basal,
middle, apical, and total articles of PLS 0,75,
0.50, 0.63, 1.88, respectively: 0.55 apart. Apical
segment digitiform. Australotheline crescent ab-
sent but sclerotised disc on ventral elastic cuticle
between hirsute basal article and abdomen
Booklung apertures broad, ovoid (Fig. 8B),

DISTRIBUTION

Known only from Dragon Lake, Liuzhou,
Guangxi, China.

Family DIPLURIDAE Simon, 1892

RELATIONSHIPS

As with the Cyrtaucheniidae, Goloboff (1993)
found that the Dipluridae of Raven (1985) were
paraphyletic, However, a possible new genus
secn in the MNHN, Paris, is intermediate be-
tween Ischnothelinae and Euagrinae and may
reduce that paraphyly significantly (Goloboff.
pers. comm, ).

Main (1993) also attempted to test the syn-
apomorphies of the Dipluridae in establishing the
relationships of the cave-dwelling genus
Troglodiplura Main, 1969 and arrived at the same
conclusion as Raven (1985),

636

MEMOIRS OF THE QUEENSLAND MUSEUM

Fic. 9, А-К, Leptothele bencha sp. nov. A-H, К, d; 1, J, 9, A, body (without legs), д, dorsal view. B, eyes,
dorsal view. C, maxillae, labium and sternum, ventral view. D, abdomen, ventral view. E, F. Tibia and metatarsus
of leg I], retroventral view (E) and leg I, prolateral view (F). G, Н. Palp, prolateral (G) and ventral (Н) view. I,
Leg П, prolateral view. J, К, Spermathecae of two 9 ©, dorsal view. Scale lines = І тт, except B, E-I 0.5mm,

J, K 20.1 mm.

Subfamily EUAGRINAE Raven, 1985
Leptothele gen. nov.

TYPE SPECIES
Leptothele bencha sp. nov.

DIAGNOSIS

Leptothele is closely related to the genus group
Euagrus Ausserer, 1875 plus Phyxioschema
Simon, 1889 (Coyle, 1988), which share inter-
locking spinule patches on d femora I and II, a
median d tibia II mating apophysis with
megaspines and pseudosegmented apical articles
of PLS. Leptothele resembles Euagrus Ausserer
but differs in a single ventral process on

metatasus II and a small transverse distal ridge on
ventral tibia П of d (Figs. 9E, I); the sper-
mathecae lack sclerotised stalks, scopula light on
tarsi I, Il of б and leg tarsi in both sexes aspinose.
It also resembles Masteria in the small size and
squat maxillae with glabrous tumid anterior edge
but differs in dark pigmentation of the carapace,
abdomen, and legs, in a narrow but band-like
serrula (Fig. 5F) and in lacking an australotheline
crescent.

DESCRIPTION

Tiny euagrine spiders with hirsute carapace and
pit-like fovea (Fig. 9A). Eight eyes on tubercle.
Chelicera geniculate, teeth only on promargin.
Serrula a narrow band 3-5 denticles wide (Fig.

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

SF). Leg tarsi of both sexes aspinose; б cymbium
apically with cluster of thick dark, spine-like
setae. No preening combs. Trichobothrial bases
deeply corrugiform, as in Masteria, tarsal organ
a low dome with concentric ridges (Fig. 4E); leg
cuticle smooth. Paired claws with one row of
teeth in slight S-shape; unpaired claw with sev-
eral fine teeth, basally on asingle process, distally
sessile (Fig. 3E, F). Four spinnerets, PMS widely
separated; PLS very long, with pseudosegmented
apical joint; no australotheline crescent (Fig. 9D).

Males: tibia I unmodified but armed with thorn
spines; tibia П with 2 megaspines on low spur
medially and a small transversal ridge distally;
metatarsus П bearing pointed ventromedian thom
(Fig. 9E, I); palpal bulb pyriform, with elongated
embolus; cymbium dissimilarly lobed and api-
cally with distinctly thick setae (Fig. 9G, H),
Interlocking spinules on femora I. П.

Females with two spermathecae, densely cov-
ered by small protuberances. Each spermatheca
divided in two lobes, the inner one with a narrow
but not sclerotised stalk.

ETYMOLOGY

Greek, lepros (small, thin, weak) and shele,
(wart, nipple = ѕріппсгеі, in figurative sense).
The gender is feminine.

RELATIONSHIPS
Absence of cuspules on maxillac and
seudosegmentation of 3 tarsi indicate that affin-
ities of Leptothele are clearly within the euagr-
ine+masteriine clade (Raven, 1985, fig. 6).
Presence of a spur and megaspines оп tibia П and
the associated absence of an australotheline cres-
cent unequivocally places Lepiothele in a trichot-
omy with Evagrus and Phyxioschema. However,
dentition of the third claw severely tests the no-
tion of the common process on the Masleriinue.
At its basal end, the group of teeth resembles that
of the Masteriinae—long teeth set on a broad base
off the claw, whereas the base of the distal tooth
is clearly on the claw (Fig. 3F), No doubt similar
problems arise in determining the cheliceral den-
tition in plurident and fissident salticids,

Since Raven (1985), two new diplurid genera
have been described. Of those, only Chilehexops
Coyle, 1986 need be considered further. The
trichobothrial bases of Chilehexops are unlike
any other diplurids in that they lack the multiple
longitudinal corrugations so well developed in
Masteria (Raven, 1979; fig. 21), The segments of
the posterior lateral spinnerets are more nemesoid
in relative size—the apical segment is barely as

637

long if not shorter than the median segment. As
Coyle (1986) noted, Chile hexops genus shows
no close relationships with any known euagrine
genus and may represent a separate clade within
the Dipluridae where its inclusion requires an-
other character change (spinneret relative length
reversal) and a weakening homoplasy in the syn-
apomorphies proposed by Raven (1985). In any
case, it does not affect the position of Leptothele
in the Dipluridae.

Leptothele bencha sp. nov.
(Figs 1, 2C, ЗЕ, F. АЕ, 5F, G, 9A-K)

MATERIAL EXAMINED

HOLOTYPE: QMS29275, à, Khao Phanom Bencha
National Park, 8°15°N, 98755" E, 260 m altitude, Krab!
Province, Thailand, 21 Sep 1992, PJS. PARATYPES:
QMS29276, MHNG, NHMW, 1 3 (matured 22 Oct
19925, 4 9 2. data as for holotype.

ETYMOLOGY
Far the type locality,

DIAGNOSIS
As for genus.

DESCRIPTION

Holotype male: QMS29275. Total length 3,7,
Carapace 1,49 long, 1.33 wide, Abdomen 1.74
long, 1.07 wide.

Carapace brown, mottled with dark brown on
cephalic and coxal areas. Abdomen dorsally dark
brown, apart from light anterior patch, densely
covered with small grey hairs and few inter-
spersed long setae in the anterior part. Legs and
chelicerse light brown dorsally, Ventral side of
body yellow brown. Spinnerets dark brown, with
light ventral spots. In life, the spmnerets have a
conspicuous white ventral side, like an icing,
which is lost in alcohol.

Carapace oval, almost flat, highest near fovea,
covered with fime brown hairs; setae absent.
Fovea a deep pit, occupies са, 1/10 of carapace
width at that point (Fig. 9).

Eyes on dark raised tubercle; two rows, front
row slightly procurved, back row indistinctly re-
curved. Group 0.14 long, front row 0.31 wide,
back row 0,34 wide, occupies slightly more ihan
1/3 of carapace width at that point. Sizes and
interdistances: AME 0.06, ALE 0.10, PME 0.07,
PLE 0.08; AME-AME 0.04, AME-ALE 0.01,
PME-PME 0.11, PME-PLE 0.02, ALE-PLE
0.02,

Chelicerae small, 0.29 long, geniculate, with
dorsal setac projecting anteriorly. No inter-

638

cheliceral tumescence. Fang groove with 11 teeth
on promargin and with 6-7 tiny median denticles
(granules). Maxillae 0.24 wide, 0.37 long in front,
0.40 behind. No cuspules or anterior lobe, Gla-
brous tumid area along proventral margin.
Serrula a band 3-5 denticles wide (Fig. ЗБ). La-
bium 0.09 long, 0.30 wide. No cuspules. Anterior
edge with setae and transverse glabrous tumid
area. Sternum 0.88 long, 0.79 wide, cordate, cov-
ered with long setae. Sigilla indistinct. marginal.
Labiostemal suture a broad groove.

Leg and palp measurements:

I I Ш IV Palp
Femur 135 119 LU 140 075
Patella 073 0.66 O58 O66 044
Tibia 0.92 084 0.78 113 0.52
Metatarsus 0.90 0.97 1.01 133 -
Tarsus 067 069 0.69 079 034
Total 447 425 417 531 2.05

Leg formula 4123. Legs covered with fine
brown hairs and setae. Preening combs absent.
Patch of spinules on retrodorsal femur T and pro-
dorsal femur II form interlocking mechanisms.
Ventral and prolateral tibia T with 9 long thick
spinge: coupling spur absent (Fig. 9F). Ventral
tibia П with 2 megaspines on low median process
(1ndistinct spur) and low transverse distal ridge.
Ventral metatarsus П with short anteriorly in-
clined process like a cuticular thorn (Fig. 9E, K).
Scopula weak on tarsi I, IL Spines: leg tarsi
aspinose; J, fe dô long, pa v4, ti p2, v7, me v4; 11,
fe d6 long, pa vd (weak), ti p2, v12-2 megaspines,
me p2, v2; Ш, fe dB long, pa d5, r1. v3, ti d5, p2,
r2, v6, me d4, р1(2), v6(7); IV, fe d8 long, ра d4,
rl, v3, ti d5, p2, 13, v6, me 07, p2, r1, v5. Palp: fe
dé long, pa d3, ti d3, p2, v4, ta d2(3). Trichoboth-
ris discemible only with difficulty. About 5 in
two rows on tibiae, 4-7 in one row on metatarsi
and tarsi.

Paired claws with ca, 9 teeth im one sigmoid
row; 3-5 needle-like teeth on unpaired claw and
heel-like process below (Fig. 3E, F).

Palp. Cymbium dissimilarly lobed, retrolateral
lobe more pointed and longest, bearing 2-3 short,
terminal spines, Bulb pyriform, embolus long and
tapering (Fig. 9G, Н).

PMS 0.32 long, 0.08 wide, 0.32 apart; PLS 2.17
long, articles: basal 0,66, median 0.67, apical
pseudosegmented 0.84 long, mid-diameter of
basal article 0,18, of apical 0,09, Australotheline
crescent absent (Fig. 9A, D).

Paratype female: QMS29276, Total length 4.8.
Carapace 1.85 long, 1.46 wide. Abdomen 2.34
long, 1.63 wide. As for d. except;

Posterior portion of ventral abdomen darker
than anterior.

MEMOIRS OF THE QUEENSLAND MUSEUM

Carapace hair cover more dense than in à; a
pair of long bristles in front of fovea, one long
sigmoid bristle in front of PME. Fovea occupies
1/12 of carapace width at that point.

Eyes group 0.20 long, front row 0.34 wide,
back row 0.37 wide, occupies slightly more than
1/3 of carapace width at that point. Sizes and
interdistances: AME 0.07, ALE 0.14, PME 0.08,
PLE 0.10. AME-AME 0.04, AME-ALE 0.02,
PME-PME 0.12, PME-PLE 0.02, ALE-PLE
0.02.

Chelicerae 0.38 long. Fang groove with 12
teeth on promargin and with 9-12 tiny median
denticles (granules). Maxillae 0.38 wide, anterior
length 0.50, posterior length 0,58, Labium 0.12
long, 0.40 wide. Sternum 0.99 long, 0.93 wide.

Leg and palp measurements:

I I nt IV Palp
Femur 127 122 116 142 091
Paiella 0.76 073 067 076 O55
Tibia 0.88 0.78 O76 105 0.63
Metatarsus 0.84 079 O91 LI? -~
‘Tarsus 0.56 056 064 064 0.69
Total 431] 408 414 304 278

Leg formula 4132. No spinules on femora I, M,
Scopula absent. Spines: leg tarsi aspinose; І, fe d6
long, pa v3 weak, ti p2, v4, me v3(4); IL fe d6
long, pa v4(5) weak. ti p3, v4, me p2, v4(5); Ш,
fe d8 long, ра d4, cl. v3(4) weak, ti 03(4), p2, r2,
v5(6), me d4, v5(7); IV, fe d9 long, pa d3, rl, v3
weak, ti d3, p2, r3, v5, me d4, p2, гі, v5, Palp: fe
d5 long, ti v6, ta уб, Trichobothria discernible
only with difficulty. About 6-7 in two rows on
tibiae, 5-6 in one row on metatarsi and tarsi,
Paired claws with 9-14 teeth in one sigmoid row,
decreasing in size posteriorly; 13 (14) on palp,
and ca. 5 needle-like teeth on unpaired claw,

PMS 0.43 long, 0.12 wide, 0.47 apart; PLS 2.22
long, articles: basal 0.69, median 0,69, apical,
pseudosegmented, 0.84 long, mid-diameter of
basal article 0.23, of apical 0.14.

Spermathecae two, each divided in two anterior
lobes; inner lobe, knob-like, its stalk narrow,
smooth and not selerotised; all with dense cover
of small pointed protuberances (Fig, 9T, J),

VARIATION

Body measurements (d, п=2, ?n-4) total
length (including chelicerae) 4.41-4.97, 4.8-6.1,
carapace length 1.83-2.09, 1.9-2.0, carapace
width 1.46-1.65, 1.4-1.6. One 9 lacks one PMS,
in another the fovea is short, straight, and trans-
verse.

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

* THAILAND

Phyxioschema suthepium

639

tion of Phyxioschema
= cox vi

500

^ Kilometres

FiG. 10. Occurrence of Phyxioschema suthepium Raven & Schwendinger in Thailand; and of Phyxioschema

(inset).

DISTRIBUTION & NATURAL HISTORY

Khao Phanom Bencha National Park, covering
an isolated mountain near the west coast of south-
em Thailand. Spiders were found on slopes along
a stream in semi-evergreen rainforest at altitudes
of 250-280m. They live in small funnel-shaped
webs, like those of Phyxioschema suthepium,
under debris ànd in crevices on the forest floor.

One mature ó was taken from the forest in late
September, another matured in captivity one
month later. Females moult twice per year;
moulting data: 25 Sep and 17 Jan (1st), 26 Nov
and 16 Jan (2nd), 9 Oct (3rd), 18 Jan (4th), 20 Mar
(5th $). One 9 built its first lenticular egg case
suspended in the web on 24 Sep (12 spiderlings
hatched 6 Sep), its second in Oct (9 eggs) and its
third on 12 Nov (5 spiderlings hatched 3 Jan); two
other females had 16 and 13 eggs in their first egg
case, Mating in captivity was not observed,

Phyxtoschema Simon

Phyxioschema suthepium Raven & Schwendinger
(Figs 10, 11A-F)

Phyxioschema suthepia Raven & Schwendinger, 1989, p.
55 (emended to suthepium by Platnick, 1993, p. 91.)

REMARKS

The diagnostic character "3 megaspines on
ventral spur of d tibia IT", first given (Raven &
Schwendinger, 1989) to distinguish P. suthepium
from P. raddei Simon, is intraspecifically vari-
able. Among 48 d later re-examined, two had
three megaspines on one side, and only one had
three on both sides (Figs 11B-D); all others have
only two megaspines (Fig. 11A). Three
megaspines in P. suthepium males, thus is the
rarer of two states. Other characters given in the
description, however, sufficiently support P.
suthepium.

MEMOIRS OF THE QUEENSLAND MUSEUM

Fic 11. A-F. Phyxioschema suthepium Raven & Schwendinger. A, normal type of d tibia and metatarsus II,
ventral view. B-D, variation in number of megaspines on d tibia II, ventral view. E, palp, retroventral view. F,

leg II, prolateral view. Scale lines 2 0.5 mm.

DIAGNOSIS (EMEND.)

P. suthepium differs from P. raddei in lacking
foveal setae and having metatarsal preening
combs on legs II-IV. Males with two rounded
instead of three sharp ventral keels on proximal
metatarsus II; 2-3 megaspines on tibia II.

DISTRIBUTION & NATURAL HISTORY

New records from the provinces Mae Hong
Son, Nakhon Sawan, Cholburi and Rayong in-
dicate that P. suthepium occurs widely in low-
lands (altitudes 30-750m) of northern,
southeastern and central Thailand (Fig. 10).

One d, which matured on 19 Dec 1989, at-
tempted a surplus moult on 5 May 1990. The
exuvium was not completey shed, as the ster-

num detached from the coxae, and the spider died
in the process. Supernumerary ecdysis in mature
8 d may be the result of exceptional longevity
due to conditions in captivity.

ACKNOWLEDGEMENTS

PJS thanks Prof. Dr Panee Chiowanich and Dr
Porntip Chantaramonkol (Biology Department,
Chiang Mai University) for facilitating research
in Thailand, which was supported by the Depart-
ment of Technical and Economic Cooperation
and by the Thai Ministry of University Affairs
(Bangkok). The Royal Forest Department pro-
vided permission to collect specimens in pro-
tected areas, the National Research Council of

NEW MYGALOMORPH GENERA FROM THAILAND & CHINA

Thailand approved the research work. The
"Österreichische Forschungsgemeinschaft" sup-
ported a visit by PJS to the Queensland Museum.
Bronywn Mitchell drew Figs 6A-C, E, 7A, D, L,
M, and 8A-E. We are grateful to Dr Pablo
Goloboff, Instituto "Miguel Lillo", Argentina,
Prof. МІ. Platnick (AMNH), and Prof, F.A.
Coyle, Westem Carolina University, for com-
ments on the manuscript.

LITERATURE CITED

COYLE, F. A. 1986. Chilehexops, a new funnelweb
mygalomorph spider genus from Chile ( Araneae,
Diplundae), American Museum Novitates, 2860;
1-10.

1988. A revision of the American funnel-web
mygalomorph spider genus Euagrus (Araneae,
Dipluridae). Bulletin of the American Museum
Natural History 187 (3): 203-292.

DAUGHERTY, C.H., CREE, E., HAY, J.M., &
THOMPSON, M.B, 1990. Neglected taxonomy
and continuing extinction of tuatara (Sphenodon).
Nature 347 (6289): 177-179.

GOLOBOFF, P.A. 1993. A reanalysis of mygalomorph
spider families (Araneae). American Museum
Novitates 3056: 1-32.

MAIN, B.Y. 1981. Eco-evolutionary radiation of
mygalomorph spiders in Australia. In Keast A.
(ed.), Ecological Biogeography of Australia. The
Hague, pp. 853-872.

1982. Some zoogeographic considerations of fami-
lies of spiders occurring in New Guinea. Pp.
583-602. In Gressitt, J. L. (ed.), Biogeography
and Ecology in New Guinea. (Dr W. Junk: The
Hague).

1993, Biogeographic significance of the Nullarbor
cave mygalomorph spider Troglodiplura and its
taxonomic affinities. Journal of the Royal Society
of Western Australia 76; 77-85.

MAIN, В. Y, & MASCORD, R. 1969. A new genus of
diplurid spider (Araneae: Mygalomorphae) from
New South Wales. Journal of the Entomological
Society of Australia 6; 24-30.

PLATNICK, N.I. 1993. ‘Advances in Spider Taxon-
omy 1988-1991, With synonymyies and transfers
1940-1980." (New York Entomological Socicty &
American Museum of Natural History: New
York).

641

RAINBOW, W.J. 1920. Trap-door spiders of the Che-
vert Expedition. Records of the Australian Mu-
seum 12: 77-85.

RAVEN, P.H. & AXELROD, D.L 1972. Plate tectonics
and Australasian Paleobiogeography. Science
176: 1379-1386.

RAVEN, R.J. 1979, Systematics of the mygalomorph
spider genus Masteria (Masteriinae: Dipluridae:
Arachnida). Australian Journal of Zoology,27:
623-636.

1981a. The mygalomorph spider genera Phy-
xioschaema Simon and Stenygrocercus Simon
(Dipluridae: Ischnothelinae), Bulletin of the Brit-
ish Arachnological Society 5 (5): 225-231.

1981b. A review of the Australian genera of the
mygalomorph spider subfamily Diplurinae
(Dipluridae: Chelicerata). Australian Journal of
Zoology, 29: pp. 321-363.

1985, The spider infraorder Mygalomorphae
(Araneac): cladistics and systematics. Bulletin of
the юп Museum of Natural History 182:
1-180.

1986. A revision of the spider genus Sason Simon
(Sasoninae, Barychelidae, Mygalomorphae) and
its historical biogeography. Joumal of Arachno-
logy 14: 47-70.

1988. A revision of the mygalomorph spider genus
Idioctis (Araneae, Barychelidae). American Mu-
seum Novitates 2929: 1-14.

1991. A revision of the mygalomorph family
Diplundae (Araneae) in New Caledonia. In J.
Chazeau and S. Tillier (eds), Zoologica
Neocaledonica 2. Mém, Museum nationalc
Histoire Naturelle, A, 149: 87-117.

1994, Mygalomorph spiders of the Barychelidae in
Australia and the western Pacific. Memoirs of the
Queensland Museum 35(2): 291-706.

RAVEN, R. J. & SCHWENDINGER, P. J. 1989. Ona
new Phyxioschema (Araneae, Mygalomorphae,
Dipluridae) from Thailand and its biology. Bulle-
tin of the British Arachnological Society 8 (2):
55-60.

ROEWER, C.F. 1942. Katalog der Araneae von 1758
bis 1940. (Paul Budy Bd.: Bremen) 1: i-viii +

1-1040,
SCHWENDINGER, Р. J. 1991. Two new trap-door
spiders from Thailand (Araneae,

Mygalomorphae, Idiopidae). Bulletin of the Brit-
ish Arachnological Society 8 (8): 233-240.

642

AN ALIEN FLATWORM IN AUSTRALIAN WATERS,
Memoirs af the Queensland Museum 38 (21642. 1995). In
the Botanical Gardens on Mount Coot-Tha, Brisbane are
several . One displays blue water lilies, Nymphae spp
some of which are Introduced, There, 100, are simal! planarian
Darwarnms (S) Sem long) with à triangular head wilh rwo
white patches each containing a dark суе, and a yellowish
brown body with mimerpus Whitish spots,

Thougli this population exclusivebv asexual, exter-
nal aranceis sufficient lo identify Girardin figrimi, noto-
rious for the exse with which it has recently extemied ils
Histriubonal range chrough accidental introductions С,
figrina is а presumed native of Nori America; sexually
reproducing animals found in Brazil and Uruguay are thought
the result of lusman-aided imnuigratioa. In recent times G.
tigrina has been introduced With aquarium plants imo Europe
(бани & Gourhault, 1978; Ribas et al. 1989) where il now
occurs in natural water bodies. In Japan, this reduced species
was initially fand only in culture ponds and fish tanks, hut a
naturalised lation has been recently reported Irom Naga-
saki (Kn u etal., 1993),

in Australia, unidentified, asexual specimens nf Girardia
р. (possibly G. tigring) have been reported from Унлопа
{ alt 1974: Нау & Ball, 1979), Further indirect evidence for
the occurrence of G. figrina in Awstralia comes Irom à caure
in Japan of Australian crayfish, Cherax fenyirmanns, Piro-
duced from Westem Australia in 1984. In 1985 an asexual
population was present in she culture ponds (Femurs ct al..
1985). 1 is probable that Girandia has already achiewed quite
а large distribution within Australia, most likely through
Jotreductions and unintentional Wansfer within the country,

Colonisation into new territories is facilitated by the
Species’ capacity 10 readily reproduce by transverse fission-
ing; one animal can be the parent of an entire population.
Several other plananan species have enlarged their dis-
tributional range as a result Of human activities, The land
planarian Bipaliam kewense, originally confined to S.E. Asia,
has been reported from hothouses worldwide, as it is easily
transported in pots of soll with exotic plants, Another land
planarian, Artioposthia triangulata, was introduced from
New Zealand [mo Britain, and was reported in 1963 from
pardens near Belfast (Ball & Reypoldson, 1981), Presently,
the species has spread over Northern Ireland and Scotland,
and Ñs predatory activities have caused serious depletion of
local earth woens populations (Ogren, 1995; Putnam, 1994),

Other freshwater planarians have extended their ranges by
Iruman-aided dispersal, For example, Phagocata woodwarthi
was introduced into Loch Ness attached to North American
p meris used during searches for the "Loch Ness Monster?
(Reynaldson et al, [981). Introduction of G tigrina into
Australia сой! potentrally threaten indigenous, МИ poorly
studied, native planarians. G, 1ígrina feeds on small crusta-
ceans, oligochactes and gastropods hy penetrating the body af
its prey with its pharynx and suckang out the contents. In
periods. of lowered food supply they are able to shrink, calling
on their body reserves. There are few known predators, Cou-
pled with Meir powers of regeneration, this makes them
fonnidably resistant. In continental Europe no ill effects of G
tigrina on the planarian fauma have been observed, hat in
Britain Reynoldson ( 1985) reported a rapid and drastic decline
of two native planarian species after introduction of G. rigri&a
into an English coastal lake. According to Keynoldson, re-
инем of native plunarisms by G. tigrina probably results

interspecific competition for food. Australia has the
greatesa diversity and the most primitive representatives of thé
Dugesixdae to which G. riprima belongs. It would be tragsc to
have our indigenous fauna Pall prey To American Invaders

Australia has a long history of planned and unplanned
емеше wpodhuetuóns Anhingtòn & Mitchell (1986) gh-

MEMOIRS OF THE QUEENSLAND MUSEUM

lighted the need For insqeecron and quarantine. walt reg ant no
disease, of introduces aquatic plants and @sh. Small worms
such as С. tigriaa are not pathogens, but merely pers
of plants and will produce nò obvious damage, Hence, they
may be easily overlooked and easily tran either with
planis or with water associated wilh plants and animals. They
are sinall, generalist predators with evidently wide tolerances.
and, because they can reproduce through fission as well as
Sexually, thé worms сап in а manner similar to many
plants, This will be panicularly enhanced through inadvertent
human intervention, All thres components of invasion re-
parted hy Arthington and Mitchell; introduction, establish-
ment and dispersal, are ensy for С, tigrina. Hopefully, the
npportunily may exist to limit rhe spread of ris alien, and
monitor bath its spread and impact on the native fauna as was
advocated in Britain (Ball & Reynoldsan, 1981).

Literature Cited

Arthington, А.Н. & Mitchell, D.S. 1986. Aquatic invuiling
Species, Pp 34-53, In Groves, К.Н, & Burdon, 3), (eds)
Ecology of hiolagical invasions: an Australian c
tive (Cambridge University Press: Cambie)

Ball. L R., 1974. A new genus and species of freshwater
planarian from Australia (Platyhelminthes: Turbella-
па), Journal of Zoology 174: 149-158.

Ball, |, R. & Reynohtson T, B., 198}. ‘British Planarians’.

(Cambridge University Press: Cambridge).
Dahm, А, G, & Gourhault N.. 1978, Tricladida et Tem
nocephalida. Pp. 16-20. in Mies, J. (ed.) “Limnofauna

Europaea’ (Fischer Verlag: Stuttgart)

Hay, D. A, & Ball |. R., 1979. Contributions to the biology of
freshwater planarlans (Turbellaria) from the Vietorian
Alps, Australia, Hydrobiologia 62: 137-164.

Ka'wakatsa, M., Tamura, S. Takai, M., Yamamoto, K., Ueno
R, & Oki 1, 1993. The first record of occurrence of n
naturalised ulation of Dugesia tigrima (Girard,
1850} at Nap: aki, Kyushu, Japan (Turbellaria;
Treladida: Paludicola), Bulletin of the Biogeographi-
eal Society, pn ts 48: 28-34,

Ogren, К. E., IN PRESS. Predation behaviour of land planar-
dans, Hydrobiologia 305,

Putnam, C., 1994. War of worms rages across Scotland. New
Scientist 18 June, no. 1930: 18.

Reynoldsom, T. B., 1965, An article on invasion – Take-over
ofan Anglesey lake hy an American species of triclad
-the potential threat to the native triclad fauna, Bulletin
af the British Ecological Society 16: 80-86.

Reynokison, T. B, Smith B. D. & Maitland P. 8, 1981. A
species of North American triclad (Paludicola;
‘Torbellana) Dew tò Britain found in Loch Ness, Scot-
Jand, Journal of Zoology 193; 531-539.

Kihes, Mu Riutort M, & Варцпа J., 1989, Morphological and
biochemical variation in populations of Dugesia (G.)
Nerina (Turbellaria, Tricladida, Paludicola) from the
westerm Mediterranean) biogeographical) and
ўпісана implications Journal of Zoology 218%;

2
Tamura, 5., Oki, L, Kawakatsu, M.. Ninigawz, М.
Mutsusato, T, & Suzuki H., I985, A note on an intro-
duced species of freshwater planarian, Dugesia tigrina
(Girard, 1850), found in culture ponds of Australian
crayfish in Kagoshima prefecnire, Japan. The Вые
Fuit Women's College 23: 133-137.
Ronald Sluys, Intipure for Systematics & Population Biology,
Zoological Museum, University af Amsterdam, Baris Joffe,
Zoological Insvitwe. Rucsinn Academy of Sciences, St. Peiers-
burg, Russia, & Lester R. С. Cannon, Queensland Museum,
PO Box 3300, South Brisbane, Queensland 4104, Australia;
25 Дикт 1995,

THE RESULTS OF TEST EXCAVATIONS IN THREE ROCKSHELTERS
IN SOUTHWEST QUEENSLAND

RICHARD ROBINS

Robins, R. 1995 12 01: The results of test excavations in three rockshelters in southwest
Quem Memoirs of the Queensland Museum 38(2): 643-666. Brisbane. ISSN 0079-
5.

In order to document the archaeology of southwest Queensland and to establish a chronology
of Aboriginal occupation, test excavations were undertaken in three rockshelters to determine
their suitability for further investigation and, if possible, to date the age of the deposits. All
proved to have shallow deposits and the cultural component of each was sparse. The deposits
from two were dated to the late Holocene. While occupation deposits from these types of
sites can provide chronological evidence, it is likely to be limited to the late Holocene.
Investigation of other types of sites, such às open sites or features associated with relict
sediments, will be required ta provide a chronology of Aboriginal occupation in southwest
Queensland.

Richard Robins, Queensiand Museum, РО Box 3300, Sowrh Brisbane, 4101 Australia; 1

August 1994,

Despite considerable knowledge gains over the
last three decades, much of Australia remains
archaeologically unknown. White and O' Connell
(1982:3) observed that Sahul prehistory *... is
based on appallingly small samples...’ Not only
are the samples small, they are also very biased
їп their distribution. While some areas of Aus-
tralia have been subjected to considerable
amounts of archaeological investigation which
has enabled, amongst other things, the establish-
ment of basic regional chronologies of Aborigi-
nal occupation, other areas have received little or
no attention. Southwest Queensland, where only
one unpublished date has been obtained from a
salvage excavation (P. Hiscock, pers. comm.) and
only one brief report of a test excavation pub-
lished (Hughes & Lampert, 1980), falls into ihe
latter category. This paper builds on this small
data base by presenting the results of test excava-
tions undertaken in 1984 in three rockshelters in
southwest Queensland. The work was undertaken
as part of a larger baseline study designed to
explore a range of aspects, including the dating
of sites, relating to the archaeology of southwest
Queensland (Robins, 1993).

A BRIEF HISTORY OF ARCHAEOLOGICAL WORK
UNDERTAKEN IN SOUTHWEST QUEENSLAND
The early archaeological record for southwest
Queensland is one characterised by infrequent
arlicles, some in popular magazines and the oc-
casional report in professional journals. Reports
in the former category include that of Lewis's
account of artefacts and campsites on the
Cuttaburra and Kulkyne Creeks (Lewis, 1966);

those from the latter include those of Riddell
(1923), who described a stone arrangement on
Durham Downs, and Elkin (1949), who described
the rock engravings on Nappermerrie Station and
recounted their associated mythology,

Тл the late 1960's and early 1970's, Federal and
State government-sponsored site recording pro-
grammes resulted in the recording of a number of
sites throughout the southwest, the aims being to
compile an inventory of sites and site types. A
largely descriptive, nan-theoretical approach was
adopted. From these recordings, information
about the nature and distribution of stone arrange-
ments (Rowlands & Rowlands, 1976, 1978) and
the style and content of rock engraving sites
(Robins, 1977) was expanded. New site types
were recorded, including burials, burial plat-
forms, stone cairns, rockshelters with occupation
deposits, scarred trees (as Ше result of obtaining
wood far boomerangs and bark for canoes), pos-
sum trees, stone quarries, extensive artefact scat-
ters, axe grinding grooves, clay and stone hearths
and huts frames (Heritage Branch files, Queens-
land Department of Environment and Heritage;
Robins, 1981; McKellar, 1984).

Oil and gas exploration in the Eromanga basin
in the lale 1970's, brought with it the need for
Impact Assessments. This work, largely confined
1o the area around the Jackson oilfield between
the Wilson River and Cooper Creek, resulted in
anumber of surveys being undertaken (e.g. Lance
& Hughes, 1983; Lilley, 1983; Richardson, 1983;
Hiscock, 1984а; Alfredson, 1987; Lance, 1990,
1992), In the course of these surveys large num-
bers of stone artefact scatters were recorded, and

MEMOIRS OF THE QUEENSLAND MUSEUM

w

wn
MT dd PA

оп;

(>

Cunnamulla

|

-

FIG. 1. Topographic map of southwest Queensland showing the location of Kilcowera and Kyeenee Stations

(after Dawson, 1974).

in some cases quarries, scarred trees and burials
were also identified, Lance (1990) comments that
most of these surveys were descriptive in nature,
limited in scope and intensity, and were designed
to identify sites to be avoided. The exception was
a predictive base line study of the Naccowlah
Block (Hiscock, 1985), in the vicinity of Cooper
Creek and the Wilson River, where variations in
the frequency and nature of sites were shown to
depend on distances from water sources, raw
materia] sources and proximity to land system
boundaries.

Only two archaeological excavations have
been undertaken in southwest Queensland. The
first was by Hughes & Lampert (1980), who sank
a ] x Im test pit into the clayey core of a dune
near Nappermerrie Station, on Cooper Creek. No
artefacts were recovered from the core, although
numerous artefacts were observed in a lag at the
surface of the dune flanks. These artefacts were
presumed to be Holocene in age, most of them
younger than 5,000 years. Other artefacts were
observed embedded in indurated sand above the

clay cores of dunes. From their geomorphic con-
text, Hughes & Lampert deduced that they were
late Pleistocene or early Holocene in age.

The second was a rescue excavation under-
taken in 1983 where an oil pipeline intersected
the lunette on the eastern side of Lake Toomaroo,
on the Dynevor Lake system just east of
Thargomindah. The excavator, Peter Hiscock,
undertook the collection of part of a surface scat-
ter of artefacts and carried out some test excava-
tions. In one of the squares a date of 1744+50yBP
(Beta 7640) was obtained from charcoal at a
depth of about 20cm (P. Hiscock, pers. comm. ).

This brief summary illustrates the poverty of
archaeological information, particularly that re-
lating to the temporal evidence of Aboriginal
occupation, in southwest Queensland,

The situation in adjacent areas of New South
Wales and South Australia 1s not greatly differ-
ent. In northwestern New South Wales the most
detailed work has been undertaken by Witter
(1992), although some basic site recording work
had been done previously (e.g. Stockton, 1977;

ROCKSHELTER TEST EXCAVATIONS

з 5

535 Murderers Bore повы?

615

290m - 250m

FIG. 2. Map showing the location af Murderers Bore rocksheltet.

Rowlands & Rowlands, 1976. 1978). Witter
(1992) undertook a survey in the Tibooburra area
as part of a larger project designed to characterise
stone artefact industries in three areas of New
South Wales. During the survey, he recorded
large numbers of artefacts differentially spread
across the landscape. Although no excavation or
dating was done, Witler assumed that the evi-
dence is all mid-to-late Holocene in age on the
basis of its general character and content, inclnd-
ing the presence of artefact types that have else-
where been dated to that time period.

Archaeological investigations along the Lower
Cooper Creek in South Australia have deter-
mined that human occupation of that area extend
back al least 1 2000y BP (Veth & Ham, 1989; Yeth
et al, 1990). ^ survey along the riverine corridor
coupled with transects across the adjacent desert
resulted in the recording of 204 archaeological
sites. Distinct differences in site densities and
assemblage structure between the eastem and
western sections of the Lower Cooper Creek were
interpreted às the product of differing Holocene
settlement and suhsistence strategies. The exea-

FIG. 3. Murderers Bore rockshelter.

vation of hearths buried within calcrete dune
cores provided evidence of late Pleistocene (с.
12000yBP) occupation.

Survey work to the north around the Coongie
Lakes by Williams (1988) recorded extensive
mussel shell middens, termite mound heat retain-
ers and stone artefacts. These sites were interpre-
ted as late Holocene in age and reflect habitation
near permanent or semi-permanent water. Exca-
vations to the south of Cooper Creek, in the
Strzelecki Desert, revealed evidence of late
Pleistocene habitation. At the JSN site, charcoal

MEMOIRS OF THE QUEENSLAND MUSEUM

from hearths (Smith et al., 1991) produced dates
ranging from about 14000y BP to about 2500yBP.

]n summary, archaeological investigations in
southwest Queensland over the last seventy years
have resulted in a number of types of sites being
recorded. However, it has really only been in the
last two decades, and with the advent of oil ex-
ploration, that attempts at comprehensive archae-
ological recording have been carried out. This
work has largely been descriptive in nature, and
a lemporal framework for the evidence is still
lacking, although indications from nearby areas
would suggest a history of occupation of consid-

ROCKSHELTER TEST EXCAVATIONS

B - Bone
ача Dripline
aiy Bedrock С - Core
жа Roof fall F - Flake
== — Silcrete boulders

K14,P21 Excavation squares
FIG. 4. Plan of Murderers Bore rockshelter.

erable antiquity. Important issues now for south-
west Queensland archaeology are to obtain a
chronology of occupation and to identify ways
such a chronology might be obtained.

Elsewhere in Australia, archaeologists have
traditionally focused on rock shelters and caves
to obtain temporal information on human occu-
pation. In southwest Queensland, numerous
rockshelters occur in the sediments of the Winton
Formation where they are exposed as dissected
residuals, mesas and buttes. Most, however,
occur in rugged terrain well away from perma-
nent or semi-permanent sources of water, and
most appear to contain only shallow and dis-
turbed deposits which are likely to prove a poor
source of archaeological information, particu-
larly chronological information.

KB - Kangaroo bone

RF - Retouched flake
Form tines at 0.5m Intervals

647

These assumptions, however,
need to be addressed. Size or depth
of rockshelters are not necessarily
a good indicator of archaeological
potential. A characteristic of Aus-
tralian rockshelters is that old, or in
other ways significant sites, do not
have to be large or deep. A number
of sites with deposits of about a
metre in depth, such as Colless
Creek (Hiscock, 1984b, 1990),
Maidenwell (Morwood, 1986),
Platypus Rockshelter (Hall et al.,
1988), Bushrangers Cave (Hill,
1986), Fern. Cave (David, 1991),
and Birragai (Flood et al., 1987)
have all provided dates that have
made important contributions to ar-
chaeological knowledge at both a
regional and continental level,
Some test excavations of
rockshelters in southwest Queens-
land were therefore necessary to
establish the validity of the assump-
tion concerning their archaeologi-
cal potential. If the assumption was
upheld, alternative approaches for
obtaining chronological sequences
in this area must be sought.

SITES

Three rockshelters were selected
for investigation. The first, infor-
mally named Murderers Bore 1, is
located on Kilcowera Station,
which is located between the Paroo
and Bulloo Rivers, approximately
60km NW of the Queensland/New
South Wales border town of
Hungerford (Fig. 1). The rockshelter is located on
the eastern side of the Willies Range, in the
headwaters of Benanga Creek, which flows into
Lake Wyara (Fig. 2). It was excavated during the
1984 field season as part of a preliminary survey
of that area (Robins, 1993).

The remaining two rockshelters are located on
Kyeenee Station. Kyeenee Station lies approxi-
mately 70km NW of Eulo and 80km NW of
Thargomindah, midway between the Paroo and
Bullo Rivers (Fig. 1). It lies on a divide between
Yowah Creek, which flows into the Paroo River
60km to the SE, and the tributaries of Bundilla
Creek which flows into the Dynevor Lake sys-
tem, 40km to the SW. One of these was а rela-
tively large rockshelter recorded by the author in

TABLE 1. Summary of frequency, size and mass of
artefacts types in excavations P21 and Ki4, Abbrevi-
ations: XU-Excavation Unit; ML=Mean length; MW
= Mean width; MM = Mean mass.

pe age i ЫЫ
mm) |(mm) |(mm)

ppu pas о [зот [27.7 [э |125 |
an naa l lios Ina 12a lo
capom L L RN
[K14/2 | p 112 |юэз |32 jos |
Lc edges a t аг 22 за _
cia {mus — Пп jus [зао |n» з |
[mds ^ ai | 120 |14
иа То [B FCU 51 С TN
[kiga|Fake — [i [79 [iso ja2 [os |

1977 (Queensland Deparment of Environment
and Hentage Branch Site number EB:A16); the
other was a smaller shelter. These rockshellters
were informally Kyeenee rockshelters I and 11
respectively.

w

MURDERERS BORE ROCKSHELTER

Murderers Bore rockshelter is located on the
northwestern side of a finger of dissected residual
that marks the boundary of the headwaters of the
tributaries that flow into Benanga Creek. It is
located 2.5km S of Murderers Bore, and 4km SW
of Benanga Creek at its nearest point. Lake
Wyara lies 13km to the east (Fig. 2). The closest
point of the Paroo River is 40km to the southeast.
The decision to excavate this site was based on
the fact that of all the rockshelters in the stu,
area it was the largest and appeared to have the
deepest deposit. There were also obvious signs of
human occupation in the form of stone artefacts
on the floor, and a dead acacia tree at the entrance
from which a section of wood had been removed.

The rocksheltér is in a scarp retreat of a table-
land and has a northwesterly aspect. The scarp
consists of a thin capping of Glendower Forma-
tion silcrete covering the softer sandstones, silt-
stones and mudstones of the Winton formation
(Fig. 3). The rockshelter has formed through the
action of cavernous weathering on the softer sed-
iments of the Winton Formation. The scarp re-
treat itself is highly eroded and is no more than
5m high in the vicinity of the rockshelter. The
scarp abuts a talus slope with shallow red earths
and silcrete, mudstone and siltstone boulders and
gravel. The surrounding vegetation is predomi-
nantly bastard mulga (Acacia clivicola) and
mulga (Acacia aneura) low open shrubland with

MEMOIRS OF THE QUEENSLAND MUSEUM

forbs and short grasses (Dawson, 1974).
Waterholes in Benanga Creek would provide the
nearest source of semi-permanent water. Y oulain
Springs, 7km to the southeast, would probably
have been the nearest source of permanent water.

The rockshelter itself is small and has an irreg-
ular shape. Its mouth is approximately 10m long
and 4m deep at its deepest point (Fig. 4). Maxi-
mum ceiling height is 3m (Fig. 5). Much of the
surface of the deposit appeared scuffed by cattle,
rock wallabies and kangaroos. The desiccated
carcass of a kangaroo lay beneath a mulga tree at
the entrance to the shelter. Bedrock was outcrop-
ping throughout much of the floor and boulders
of roof spall and silcrete lay on the floor.

EXCAVATION PROCEDURE

After driving a nail into the bedrock to the north
of the shelter to act as a datum, a baseline and an
alpha-numeric 50cm horizontal grid was estab-
lished for the site. The site was then mapped using
90° offsets from the baseline. Heights were deter-
mined with an autoset level. Two 50cm squares
(K14 and P21) thought to have the greatest depth
and the least amount of disturbance were selected
for excavation. P21 was selected because it was
the highest point in the deposit. K14 was selected
in the hope that sediments eroding downslope

2)

=

а=

a

Зои А A

fieatlon b - B

FIG. 5. Topographic profiles across Murderers Borc
rockshelter.

ROCKSHELTER TEST EXCAVATIONS 649

| | | м
| | ка ( |
| | | | g
| с

Пр

№ c “a, т
- —. А 1
MTS
А ‚ TII Kveenes
r А kshelter
vem
hi |
ONS, oc j£ |
y d 2 dp | A | \,
‚Т L \ А
4 A } I
if
WM ai |
е УТ QI = III III
| ME * : Da
| |

;
А |

3 |

“7-27
И;
Pe vm
X 5
Ф| ү ili
2 р УАДЕ ҮҮ!
ҮШ Me / ||! ^ | l -S
D c j
А 2
РА Ur A Ц | \
i box ШШ; | ! ~
——— 7 2 fi Oui ҮР
20mm. \ » r |
ГА ` JK nee Il Vv " | || 4
l | | | |
N © Rockshelter / К, |
M 1
| Чүү 9 |
-- ; 4 тү |
74 | Ñ
4 и бл ры. \ | ҮҮ
fi e m Ы N A ) | i
н
| ^ 22 `N t WII
"d ^ N Т. | M i з -
j zy 2 Cree um 7 hc 77 i^ X. Mt м )
wc ey ч. == a MeL E |
"ur 4 9i ~ 3L TI l А |
_- - ! `
| | | ў |
3 - % ~ ^ i, = I J
E 7, "s = A "i | \ > A
s ШШ) ? 4
"HEN jon
le 0 5 km

Scarp
= ЇЙЇ 200m - 250m

FIG. 6. Location of Kyeenee Ї and Кусепее I rockshelters.

wo MA Rn aet daa roots of the Acacia at the А 10x 10m squa ing from the baselin
entrance. Exca n followed the procedures down the tals Hope was laid out рга additio n tó

and "rofa rma satin ed by Johnson (1979:145- basic елу ment da ta, the minimum and aver-
165) араат? material Wak sien ed thro ough ареалын denies, ravi material types, artefact
j 1 types, the seer Laney psi ne of flakes,
2mm qo eros oil cores and g indsto Were те.
obtained "A sell Soil Co iia г Cha М orded (Ro bin $1993: 364, ae yr aryl
(Munsell Colou i. 1990).
All Thé ma terial helio мян s Wa rem s

De EN
for laboratory sorting. For ile

P21 w ted in two XUS. It contained
(XU). a grab sample (approx atl 1005) 0 he 2175kg a of de epost an and had a t denti o of ferti, Thie
Босуе рие pen ough bo Vp. bezen a dry reddish-yellow
also alysis. (Munsell (75 SYR 6/6)) sa ШУ depus sit with s ome

650

FIG. 7. Kyeenee I rockshelter.

gravel and well rounded pebbles. Beneath this
surface layer is a dry, compact, strongly brown
layer (7.5 YR 5/6) with increasing amounts. of
decaying bedrock to the base. The deposit has
been disturbed by a tree root growing through it.
Small amounts of charcoal occurred throughout.

K14 was excavated in five XUs. It contained
41.75kg of deposit and had a maximum depth of
40cm. The top 1-2cm is similar in texture to that
encountered in P21; a dry, coarse, sand with
gravel and cobbles, presumably derived from the
sandstone bedrock. Beneath this surface layer the
deposit is finer and more friable, but contains the

MEMOIRS OF THE QUEENSLAND MUSEUM

same amount of coarse material. The colour of the
deposit, reddish brown (5YR 4/4), remains the
same throughout. In parts of the square, bedrock
was encountered within 2cm of the surface while
in other parts of the square the deposit was up to
40cm in depth, indicating a very uneven bedrock
base. The deposit contains tree roots and wood,
leaves and small amounts of charcoal. A radio-
carbon date of 450+70уВР (Beta 11593) was
obtained from charcoal in the basal unit.

LABORATORY PROCEDURE

The retained sieve fractions were sorted for
cultural material, bone, charcoal and organic ma-
terial. Munsell Soil Colours were taken on the

ROCKSHELTER TEST EXCAVATIONS

Dripline

55 Bedrock

Rocks

-— Erosion wash

E b S Tree

sb 29,412 Excavation Stuaree
orm lines at 0.511 inturvat:

Wargrniale

FIG. 8. Plan of Kyeenee I rockshelter.

air-dried fine fraction. All the sorted 2mm and
4mm sieved material was retained, bagged sepa-
rately and registered Queensland Museum
QMS829.

Allidentified artefacts were weighed and mea-
sured for length, width and thickness. Attempts
were made to identify retrieved bone to species
or genera level and the bone was examined for
evidence of burning.

CULTURAL MATERIAL

Twenty five stone artefacts were recovered
from both pits; 5 from P21 and 20 from K14. All
the artefacts recovered from P21 were flakes
while K14 contained 14 flakes, 5 flaked pieces
and 1 tula (Table 1). Sixty five percent of the
artefacts recovered from K14 were from XU 2.

The small number of artefacts from these pits
restricts detailed analysis. All the artefacts re-
corded are silcrete, Flake lengths range from 3-
49mm. In general, there is not much difference
between average flake length and width in either
pit (Table 1). In K14 mean length was 9,4mm and
the mean width 11.2mm. In P21 mean length was
18.4mm and mean width 17.9mm.

651

Two small fragments of unidentifiable mam-
mal bone were found in XU 1 of K 14 which could
be the result of natural death or introduction by
humans into the site.

A dead tree (probably Acacia aneura) al the
front of the shelter retained a scar where a section
of wood had been removed from the trunk. This
scar was 850mm long and 250mm wide at its
widest point.

ASSOCIATED ARCHAEOLOGICAL EVIDENCE

In the 10m square laid out from the baseline
down the talus slope, 166 artefacts were recorded
from the surface, including 141 flakes, 14 re-
touched flakes, 6 cores, 2 multi-platform cores, 2
tula slugs and 1 tula. Maximum flake length was
70mm and minimum Imm. Maximum length of
cores was 220mm and minimum 30mm. Sixty
percent of flakes were between 60- 100mm long.
Fifty percent of cores were less than 100mm long

Section A- A

Erosion wash

Section B - B'

Section C - C'

2
Scale | ———À
0

em
ASS Rocks
Bedrack

FIG. 9. Topographic profiles across Kyeenee I
rockshelter.

652

West North

=3/
S ! с.

3 |
'

а 15 cm

1 - Dry grey to raddish grey layer 3 - Dry, dark gray layer

2 - Brown to dark brown tayar 4 - Brown to dark brown layer

(burrow?)
Excavation pit J9

Scala

1 - Loose grey ashy layer

7 - Thin arey ashy layer
2.- Compact weakly red layer - Dark grey ashy layer

9 - Reddish yellow layer with
course sediment

3 - Ashy grey layer

4- ГАРШЫ weakly rad
layer 10 - Course reddish brown
layer with charcoal
5 - Fine soll grey ashy layer
11 - Dark grey sediment with
;8- Reddish yellow layer plo leaves and
igs

Excavation pit J12

FIG. 10. Cross-sections of two faces in excavations J9
and J12, Kyeenee I rockshelter,

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2. Carbon content, colour and soil reaction for
selected in excavation J12, Kyeence I rockshelter.
Abbreviations: XU-Excavation unit
SR= Soil Reaction

xu % Carbon ur moi SR
MET —

- 7 1:5. YR 4/2
7.5 YR 3/2
7.5 YR 3/2 ERUIT.

while 30% were between 160mm-300mm long.
This figures illustrate the relatively small nature
of the artefacts. It is probable that the cores were
obtained from the locally available silcrete in
gibber form.

Distribution of artefacts across the surface was
patchy with artefact densities ranging from be-
tween 0-15/m?. Mean artefact density was be-
tween 1 and 5/т2. No knapping floors were
observed.

SUMMARY

This rockshelter contained a shallow cultural
deposit of recent age. There was evidence of
disturbance and it is possible that much of the
deposit had been moved downslope from the
Shelter. The shelter had a low density of cultural
material in the form of stone artefacts. Charcoal,
faunal and floral remains were also found in the
deposit but their presence may be attributed to
either human or non-human causes. The small
number and limited range of types represented by
the artefacts both in the deposit and on the slope
reinforce that the impression that site use was
infrequent and involved a limited number of ac-
tivities.

KYEENEE I ROCKSHELTER

Kyeenee I rockshelter is located in ап escarp-
ment at the headwaters of Burnt Creek, 6.5km
northwest of Kyeenee homestead. Burnt Creek
flows into Bundilla Creek 7km to the northwest
(Fig. 6). The rockshelter has formed at the head
of a steep-sided gully system fingering into a
scarp of Winton Formation dissected residuals.
The surface of the plateau has very shallow red-
dish brown to red, acid, loamy lithosols mixed
with boulders and gibber of silcrete and the Win-
ton Formation deposits. The vegetation consists
of areas of rockgrass (Eriachne mucronata),

ROCKSHELTER TEST EXCAVATIONS

Excavation Unit

4
2
3
a
5
в
7
e
$
10
11
2
1$ ^" ы ч
0 20 40 60
weight retained (%)
gleve size
ШШ somm MY ѕо-атт | .l4-2mm

FIG. 11, Gross texture of XU sieved in the field, excavation T1

Trockshelter,

mulga (Acacia aneura), western bloodwood (Eu-
calyptus terminalis) open tussock grassland, and
mulga, lancewood (Acacia petraea), bastard
mulga (Acacia clivicola) tall open shrubland
(Dawson, 1974). Within the gully system, large
river red gum trees (Eucalyptus camaldulensis)
grow. Semi-permanent water can be found in a
series of watetholes along the gully floor, The
rockshelter is located at the head of a small,
sloping bench which is above, and to the southern
side of, the termination of the main gully system
(Fig.7). It has a north-easterly aspect. The escarp-
ment plateau is within 1-1.5m of the rockshelter
vault, Bedrock walls extend out from the
rockshelter to box in the bench on its western and
eastern sides lo à maximum 6m width. A dropoff
to a small semi-permanent waterhole restricts its
length from the rockshelter overhang, to 10m
(Figs 8,9)

The entrance to the rockshelter is approxi-
mately 8m wide and 2m high, Eight metres in
from the entrance the rockshelter divides into two
chambers thal both extend back approximately
4m, giving a total depth of 12m. Within 4m of the
entrance the rockshelter height is about 1m (Fig.
9),

A shallow depression 1n the bedrock above the
shelter collects surface water and directs it over
the lip of the shelter, onto the bench and then into
the rock pool below. This runoff has resulted in
erosion and compaction of parts af the deposit on
the bench (Fig. 8).

Bo 100

653

The shelter is host to a colony of
bats and the surface of the deposit
is covered with bat dung. The des-
iccated carcasses of two wallabies,
sheep fleece and some bones of a
sheep were also in the shelter. Wal-
laby and kangaroo dung was also
evident in the deposit. Parts of the
surface of the deposit, particularly
towards the rear, appeared to have
been extensively scuffed by ani-
mals. Charcoal and two artefacts
were noted in the deposit prior to
excavation.

EXCAVATION PROCEDURES

A nail hammered into the south-
ern bedrock wall was used to de-
fine datum. A levelled baseline
was then nin at 355° across the
bench. From this line a series of 90°
offsets were run to various points in
the site (о outline its shape and to define an
alpha-numeric grid. An autoset level was used to
obtain heights at yarious points on the bench,
Where use of the level proved difficult, particu-
larly in the cave, a water level substituted. Two

profiles were run the length of the shelter and
bench.

TABLE 3. Faunal remains in excavation J12, Kyeenee
I Rockshelter. Abbreviation: XU=Excavation Unit

112/3 Unidentifiable

Snake vertebrae (python)
Mammal vertebrae (rat size)
Petrogale occipital (Possum

Rodent dentary. Шакай.
Small Rodent size fragment
Snake cranial fragment

Fragment, small rat size vertebrae

Snake vertebrae, rodent incisor, size
scapula fragment, macropod scapula (M.
rufus size)

2, Kyeenec

312/4 |0.47

bs. 8.49
Rodentincisor, rodent maxillary fragment,
possum size humeral fragment, macropod
dentary fragment (M. rufus size),

Ё 5.25
unidentifiable fragment
Maxillary fragment (large macropod),
J12/9 snake vertebral al fragment, suit ard
vertebral fragment

Rodent incisor, dentary fragment of
Perameles sp. (bandicoot), mammal

ае
De

Апей
JI2/L |201 |Unidentinable _

nt

654

weight retained (^5)
зо =

MEMOIRS OF THE QUEENSLAND MUSEUM

ysis, Organic carbon content, wet

25
20
15r
i
10) P ^.
FX
5 — PP D, r
= рі Mena
мй. ШЫ; ad E 2 П r1 — 1 | 1 -
1 2 28 5.5 Е m 63 125 250 500

particle size (microns)

== Excavation Unit 1 = Excavation Unit Y

FIG. 12. Particle-size distribution in the <2mm field sieved Fractions
from XU 1, 7 and 13, excavation J12, Kyeenee I rocksheller.

One 50cm square, J12, was laid out just inside
the mouth of the rockshelter and another, J9, was
laid just outside the dripline. The squares were
positioned in areas thought to have a combination
of the greatest depth of deposit with the least
amount of disturbance. The squares were then
excavated following the procedures outlined by
Johnson (1979:145-165). The excavated material
was sieved through 2mm and 4mm wire mesh and
weighed using a spring balance, All the material
held in the sieves was retained for laboratory
sorting. A grab sample of approximately 100g of
the «2mm residue was retained from each exca-
vation unit for further analysis.

LABORATORY PROCEDURES

The 4mm sieved fraction was sorted into its
constituent parts: stone, stone artefacts, charcoal,
bone and organic material. The 2mm sieve frac-
tion was wet sieved before sorting into the same
categories. Afteranalysis, the sorted material was
bagged according to category, and registered into
the collections of the Queensland Museum under
the registration number QMS828.

All identified artefacts were weighed and mea-
surements of length, width and thickness taken.
Recovered bone was sorted into individual frag-
ments and identified to family or genus where
possible, Organic fragments were sorted and at-
tempts to identify individual fragments to genus
or species were made.

Soil analysis of the fine fraction was under-
taken on selected Excavation Units (J9/1/8/16
and J12/1/7/13). This included particle-size anal-

—*- Excavation Unit 13

and dry Munsell soil colours and
soil reaction (pH).

DEPOSIT DESCRIPTION

The character of the deposit in-
side the shelter is different to that
outside the shelter. The deposit in-
side the shelter exhibited a light,
dry, complex character while the
deposit outside the shelter is more
compact and homogeneous.

J12 PROFILE DESCRIPTION

112 contained deposit to a depth
of 55cm. Eleven stratigraphic units
(SU) were identified in the southern
and eastern profiles of the test pit
(Fig.10).

SU 1 is a thin, even surface layer
approximately 2cm thick. It is a

grey (5Y 6/1), loose, disturbed de-
posit with a бвр amount of bat and macropod
dung mixed with some roof spall.

SU 2 is a weak red (10YR УЗ) layer between
5cm and 8cm thick. It contains kangaroo and bat
dung, leaves and twigs and charcoal. This is a
more compact, less disturbed layer than SU 1.

SU 3 is an uneven, grey (3YR 6/1), ash rich
layer varying in thickness between 2cm and
10cm. This layer still contains bat dung, leaves

TABLE 4. Organic Remains recovered from excavation
J12, Kyeenee 1 Rockshelter. Abbreviation: XU=Exca-
vation Unit

E
| 712/4 Коди зр. (leaves, fruit) Acacia sp.
112/5 ига ури sp. (leaves) Acacia sp.
m (leaves) Acacia s,
Modes) Abutilon sp. (fruit 5
rich Ius 5р. (leaves, fruit) Acacia sp.
H27 115.6 |n rues e SP P

Eucalyptus sp. (fruit) Acacia sp.
12% phyllodes) Е

(лою jos | Eucalvplus sp. (leaves) Acacia sp.
1112/9 |96 | Eucalyptus sp ) P

112/10 Киса урты sp. (leaves) Acacia sp.

ЧЫН:

Eucalyptus sp. possibly tessellaris
(leaves) Acacia sp. (phyllodes)

Eucalyptus sp. (leaves) Acacia s
(ph Ides) E 5

bn |а

12/13

ROCKSHELTER TEST EXCAVATIONS

655

FIG. 13. Wood shavings, XU 5, J12.

and twigs but increasing amounts of both char-
coal and roof spall were noted.

SU 4 is a thin, discontinuous, finely textured
weakly red (10YR 5/4) deposit overlying SU 5
and 8.

SU 5 is a fine, very soft, grey (5Y 6/1) ash rich
layer with increasing amounts of roof spall and
large charcoal fragments. This layer has a thick-
ness of between 1em and 9cm and is restricted to
the southern side of the pit. It dips sharply at the
southeast corner suggesting that it had been dug
into SU 6.

SU 6 is a finer textured reddish yellow (5YR
6/6-6/8) layer with less charcoal that SU 5. It
varies in thickness from between 6cm and 13cm.
Like SU 5 above, it also dips sharply at the
southeast corner, suggesting that it has been dug
into SU 8.

SU 7 is a thin, discontinuous, grey (5 YR 6/1),
ashy layer that intrudes into SU 6.

SU 8 is a thick, dark grey (5 YR 4/1), ash rich
layer with large amounts of charcoal as well as
leaves and twigs. It has uneven upper and lower
boundaries.

SU 4, 5 and 6 directly overly SU 8 which in tum
lies on Stratigraphic Units 10 and 11 as well as
bedrock. It varies in thickness from between 5cm
and 25cm.

SU 9 is a small yellowish red (SYR 5/6) lens of
coarse sediment within Unit 8.

SU 10 is a reddish brown (5YR 4/4) lens con-
taining coarse sediments and charcoal. It lies both
on bedrock and SU 11 and has a maximum thick-
ness of 10cm.

SU 11 is a discontinuous unit lying on bedrock.
It is absent in the southeast corner of the pit,
suggesting that SU 8 has been excavated into SU
11. Itis a fine, dark gray (SYR 4/1) sediment with
charcoal, leaves and twigs throughout.

The profile reveals a complex depositional his-
tory. Stratigraphic Units 11, 8, 6, 5, 3, and 2
represent a series of occupation events indicated
by large amounts of charcoal and roofall as well
as disturbance of previous deposits. The occupa-
tion represented by these events alternates with
discontinuous SUs 10, 9, 7, 4, which contain less
charcoal and finer sediments. These may repre-
sent substantial breaks between episodes in the
rockshelters’ use or, alternatively, disturbance of
the deposit. However, quantification of deposi-
tion rate to investigate rockshelter use is difficult
on the basis of this small pit.

J12 SEDIMENT ANALYSIS

J12 was excavated in 13 Excavation Units and
produced ] 10kg of sediment. Laboratory analysis

656

Excavation Unit

© Фф - 0m ольш № „

MEMOIRS OF THE QUEENSLAND MUSEUM

there are peaks at 2.5 (180um)
and 4.0 phi (634m). This anal-
ysis indicates that the deposit
has a particle size distribution
similar to that obtained from
sediments in Holocene
sandplains of aeolian origin
found to the south in the
Currawinya National Park to
the south (Robins, 1993). The
similarities suggest that much
of the sediment in this deposit
may also be aeolian in origin,
although further work is re-
quired to test this proposition.
Munsell soil colours are con-

13 EE T
[^] 50 100 150 200
mass(g)

250

sistently brown to dark brown
(7.5YR 3/2-4/2) (Table 2). Soil
reaction is 6.4 in XU 1 and XU

300 350

— > J12 charcoal *2mm = —> H2 organic »2mm

FIG, 14. Charcoal and organic remains from excavation J12, Kyeenee 1

rockshelter

of the excavated sediments indicated that al-
though the profile is complex, the similarities
between stratigraphic units as measured by parti-
cle-size analysis, percentage carbon, wet and dry
Munsell colours and soil reaction, are greater than
the differences,

Only three Excavation Units; 9, 10, and 11,
contained stones >50mm in diameter. Through-
out the deposit the proportions of
the sieved fractions remained rela-
lively constant. The percentage re-
tained in the 4mm sieve for each
excavation unit varied between 5%
and 17%, in the 2mm sieve be-
tween 20% and 3%, while the <
2mm fraction varied between 61%
and 90%. The greatest variation in
percentages occurs near the base of
the pit where weathered bedrock 10
makes a greater contribution of
coarse material, in XU З and atthe 5
surface where the proportions of 44
finer sediments are greater 15
(Fig.11). 16

A half-phi particle-size analysis
of <2mm fraction from three se-
lected Excavation Units (1, 7 and
13) was undertaken. It revealed
similar particle-size characteristics
fer each of the Excavation Units
(Fig.12). For each of the samples

D Ü~ D Nn e ON ao

I rocksħelter,

Excavation Unit

7 but declines to 4.4 in XU 13.
Loss on ignition indicates a
high carbon content ranging
from 18.5% in XU 1 to 12.096
in XU 13.

This deposit has a complex
history that is the result of roof fall combining
with aeolian deposits blown in from the surround-
ing plateau. To this has been added a significant
human contribution in the form of ash and char-
coal from fires and other cultural material. The
relative contribution of these sources has varied
throughout time.

40 60 80 100
weight retained {%)
ШШШ 50-4mm C 4-2mm E comm

FIG. 15. Gross texture of XUs sieved in the field, excavation J9, K veenee

ROCKSHELTER TEST EXCAVATIONS

TABLE 5. Frequency, size and mass of artefacts recov-
ered from excavation J12, Kyeenee I rockshelter. Ab-
breviations: XU=Excavation Unit; ML=Mean length;
MW = Mean width; MM = Mean mass.

MW
Bu [eerte е ы ы шы G
222 ПЕ

mna 12/1

Core Shee TEE
[ns [Frae — [1 |321 [osi Поз |35 |
za [make —— [r [425 1222 [imo [вв |
ma jo |78 їз ов |
pon mae — à fioa hoa s fee
ШЕШТИ a ШЕШ
Үрер eat
[meer шин иан нна за
pixnmo|Fake — — [4 [223 [i94 [es [51 |
mangas | ees в fis [e |

FAUNAL REMAINS

A small amount, 17.7g, of bone was recovered
from the deposit, The individual pieces were un-
burnt, small and fragmentary. Animals repre-
sented include bird, rodents, macropods, possum,
and snake, Due to the type and condition of the
bone, identification to species level was generally
not possible. A list of remains is presented in
Table 3. The presence of these bones in the rock
shelter does not necessarily indicate human pre-
dation. Such remains could, and probably did,
result from natural deaths or were
the prey of other animals that used
the shelter from time to time.

657

recovered including 21 flakes, 5 flaked pieces and
i core. A summary of artefact types is given in
Table 5. This small number of artefacts restricts
detailed analysis. Almost half the artefacts oc-
curred in XU 7 and XU 8.

In general, the flakes tend to be as wide as they
are long. The mean length to width ratio of flakes
identified is 1:1,2 (Table 5).

The assemblage is generally small in size. For
example, the maximum flake length is 42.5mm
and the minimum 4mm. Despite the large amount
of charcoal in the deposit there was little evidence
of heat damage to artefacts. Only the core showed
evidence of exposure to heat in the form of pot-lid
scars on the cortex.

Wood Shavings. A total of twelve wood shav-
ings were found in XU 4 to XU 7. Two types were
identified. Ten shavings were from a dark wood
and had a pronounced curl. Two were from a
lighter wood and were shorter in length (Fig.13).

Charcoal. Charcoal was found in abundance
throughout the deposit. A total of 2,2kg was re-
covered in the 2mm and 4mm sieves from all
Excavation Units (Fig. 14). The charcoal was in
large pieces, firm and in very good condition.
XUs 5, 6, 7 and 11 each had over 250g. Only
Excavation Units 1, 2, 3 and 12 had less than 100g
(Fig. 14), Charcoal retained in the 4mm sieve
from XU 13 provided a date of 1580+60 BP (Beta
11591) and from XU 5 a date of 90450 BP (Beta
61794).

FLORAL REMAINS aor |
One hundred and twenty two |
grams of organic remains were \\
found throughout the excavation po) |!
(Table 4). The majority of remains \
were Acacia spp. phyllodes ог Eu- 157 (|
calypt spp. leaves and fruit. As \
these species are common to the \
site today it is not possible to de- 5
termine if they were introduced by
the human occupants of the o

weight retained (44)
35

rockshelter or were blown in.
None of the remains were charred
or burnt.

CULTURAL REMAINS
Stone Artefacts. Twenty-seven
stone artefacts (all silcrete) were

—— Excavation Unit 1

H 51 B 125
particle size [microns)

250 24С0

== Excavation Unii à * Excavation Unit (6

FIG. 16.Particle-size distribution in the <2mm field-sieved fractions from
XU 1,8 and 16, excavation J9, Kyeenee I rockshelter,

658

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG, 17. Pebble core, XU 5, Excavation J9, Kyeenee I rockshelter.
J9 PROFILE DESCRIPTION

J9 contained 65cm of deposit. It was excavated
in 16 XUs and produced 162kg of deposit. Four
Stratigraphic Units were identified in the profile
(Fig. 10).

Stratigraphic Unit 1 consists of a thin, fine, dry,
grey to reddish grey layer (5YR 5/1-5/2) with
loose gravel and humic material. Some forbs
were growing on the surface and rootlets were
evident throughout. It is 1-2cm thick.

SU 2 isa moister, more compact brown to dark
brown layer (7.5 YR 5/3-4/3) with an increase in
finer sediment. The deposit is slightly drier and
lighter with depth. This Stratigraphic Unit has a
diffuse and irregular boundary that blends into
SU 3. Its maximum thickness is approximately
25cm.

SU 3 is a dry, dark grey, (7.5 YR N4) unit that
made up the bulk of the deposit. It lies on an
uneven bedrock base and is coarser and less co-
hesive than SU 2. A number of large stones are
present throughout. À small burrow is evident in
the southwest corner. It has a maximum thickness
of approximately 45cm.

SU 4 is a small lens that intrudes into SU 3 in
the southeast corner. It is similar in colour and

texture to SU 2 and may represent a burrow from
SU 2 into SU 3.

J9 SEDIMENT ANALYSIS

The J9 deposit is more compact, and does not
exhibit the complex stratigraphic detail of J12.
(Fig. 12). However, while not evident in the pro-
file, analysis of the deposit reflects a more subtle
complexity of depositional history.

Increase in clays by weight with depth and the
decrease in sands by weight with depth may be
attributable to postdepositional weathering and
erosion, particularly as J9 lies near the edge of an
erosion gully. Deposits of J12 inside the shelter
are not as compact and show a more consistent
relationship in particle-size distribution with
depth. Further work is needed to test this expla-
nation.

Overall, sieving data indicate a similar pattern
of deposition to that of J12 (Fig.15). However, the
lower XUs have a higher percentage of coarse
fractions in them compared to J12. The propor-
tion of stones >5cm is up to 23.5% by weight in
XU 10. J9 has four peaks in the coarse fractions,
at XUs 1, 10, 13/14 and 16 (Fig.15). The finer
fractions («2mm )j, vary from between 75.3% and
52.396 by weight, with a general diminution in

ROCKSHELTER TEST EXCAVATIONS

FIG. 18. Pebble core, XU 14, Excavation J9 Kyee

representation in the lower XUs. There is a
marked increase in the weight representation of
the 50-Amm and 2-Amm fractions at the surface -
the only place where their increased contribution
is not accompanied by a similar increase in the
250mm fraction.

The particle-size analysis at half-phi intervals
of samples from selected XUs (1, 8, and 16)
reveals a particle-size distribution similar in pat-
tern to that of J12, with two important exceptions.
The first is the segregation and subduing with
depth of the 125um and 63 рт sizes (Fig. 16). The
second is the increase in the clay fraction with
depth; from 1546 at the surface to 3196 at the base.

There is minor colour change with depth. The
surface wet and dry colours are brown to dark
brown changing to very dark brown and dark
greyish brown at the basal unit (Table 6). Soil
reaction (pH) varies from 5.15 to 3.49 (Table 6).
The carbon varies from between 13.28% to
11.0% and does not show any obvious depth-re-
lated trend.

FLORAL REMAINS

Five unidentified seeds were recovered from
J9. Three were from XU 8 and two from XU 10.

659

nee I rockshelter.

CULTURAL REMAINS

Stone artefacts. Eighty five silcrete stone arte-
facts were recovered including 60 flakes, 14
flaked pieces, 6 retouched flakes and 5 cores. The
distribution of artefacts throughout the deposit is
relatively even. No artefacts were found in XU
15 or XU 16 (Table 7). No evidence of heating
was noticed on the artefacts although a pot-lid
flake and pot-lid scars were noticed on non-cul-
tural stones in XU 9.

As with J12, the flakes tended to be small and
as long as wide. The longest flake was 83mm and
the shortest 4.4mm. Mean width-to-length ratio
forall flakes was 1:1.24. Retouched flakes tended
to be larger than the flakes, with a mean length of
20.6mm, although the width to length ratio re-
mained similar at 1:1.4. Only one core, with a
maximum length of 132mm and a mass of 1,338g,
can be considered large compared to the rest of
the assemblage.

More artefacts were recovered from J9 (85)
than J12 (27). This may be attributed to one or
more of four factors. These explanations are not
mutually exclusive:

1)There was greater discard of artefacts on the
terrace than in the rockshelter (cultural factor).

Excavation Unit

0 20 40 50 60
mase (д)

—*— J8 charcoal «2mm

FIG. 19, Charcoal remains from excavation J9, Kyeenee I rockshelter

2) Artefacts discarded in the shelter were
eroded downslope onto the terrace (taphonomic
factor).

3) Artefacts on the surface above the rockshel-
ter had been eroded downslope and onto the
terrace outside the shelter (taphonomic factor);
and

4) The sample of deposit is not representative
wilh respect to artefact distribution (sampling
factor).

А more extensive excavation and detailed ex-
amination of artefact distributions of the site
would be needed to test these.

The two pebble cores are of same interest. They
are the only artefact of their type observed during
archaeological investigation’s phase either in the
vicinity the Currawinya Lakes or on Kyeenee
(Robins, 1993). Both were recovered from this
square J9; one from XU 5 and one from XU 14.
The former is formed on an ovate, waterworn
silcrete pebble. It has been substantially reduced
from one end through a series of flakes removed
along one margin (Fig. 17). A number of small
flake scars with step terminalions emanate from
the platform. This flaking has formed a dentated,
arcuate platform margin. The second pebble core
has also been formed on a small, ovale, water-
worn pebble and exhibits similar characteristics
to the previously described specimen (Fig.18).
This example has a straighter, although more
dentated, platform margin. One large flake has
been removed from the end opposite the flaked
edge. The platform also has one large and several
small flakes removed from it. These cores occur
in an area with an abundance of raw material. The
steeply dentated and step-flaked margin is remi-

MEMOIRS OF THE QUEENSLAND MUSEUM

niscent of the edges produced for
adzing tools, such as tulas, and may
imply that the cores were used as
adzes (although it is to be noted
that dentated and step-flaked mar-
gins can be produced as a result of
flaking cores).

Charcoal. A total of 1.4kg of
charcoal was recovered in the 2mm
and 4mm sieves from all XUs
(Fig. 19), Although J9 only had just
over half the amount of charcoal
recovered from J12, a similar pat-
tern of distribution emerged. In
both cases there is little charcoal at
the surface, an increase in the top
one third of the deposit, followed
by adiminution in representation to
the respective bases of the excava-
tion (cf. Figs. 14,19). Charcoal re-
tained in the 4mm sieve from XU 15 provided a
date of 1280+70уВР (Beta 11592), and from XU
8 a date of 660+60yBP (Beta 61793).

Faunal Remains. Two small fragments of un-
identifiable burnt bone were recovered from XU
9 and XU 12.

OTHER ARCHAEOLOGICAL EVIDENCE

On the plateau surface above the site, the sur-
face is bedrock which is devoid of vegetation but
covered with silcrete gibber varying in size from
1-20cm in diameter, Nodules of silcrete embed-
ded in the bedrock have been comprehensively
flaked. There is also an extensive scatter of arte-
facts on this surface. A 10x10m quadrat was laid
out on this surface, directly above the rockshelter
where a shallow depression directs surface water
over the lip of the shelter onto the terrace below.
Two thousand and eighty two silcrete artefacts
were recorded in this quadrat, including 1,807
flakes, 146 cores, 40 multi-platform cores and 89

TABLE 6.Carbon content, colour and soil reaction for

selected Excavation Units in Excavation J9, Kyeence
1 Rockshelier. Abbreviations: XU-Excavation Unit;
SR=Soil Reaction; C, (Av.)2Carbon (Av.).

% C. |Munsell Colours

SR
Avi [wa [by — oh

7.5 YR 3/2

j 7.5 YR 4.2
brown to dark
bro’

XU

э |126

dark brown

wn

hon | 10 ҮЕЗЛ 10у 42 40
very dark grey | dark greyish brown |“

Y 10 YR 4/2

|1916 10 ук? dark greyish brown |48 |

ROCKSHELTER TEST EXCAVATIONS

FIG. 20. Kyeenee II rockshelter

retouched flakes. Artefact densities within the
quadrat varied from 10-30/m?.

Ninety percent of the flakes are between 1-5cm
in length, the remainder are between 6cm and
8cm in length. Maximum flake size is 8cm, min-
imum is 13mm. Seventy five percent of the cores
are 60mm-75mm long, the remaining 25% are
30mm-50mm long. Maximum core length is
75mm, minimum is 30mm.

A small, indistinct stone arrangement is located
on the gibber strewn surface plateau surface op-

661

ы

posite the rock shelter. The arrangement has de-

teriorated to the point where the only identifiable
feature remaining is a small mound with a cleared
centre.

SUMMARY

Sedimentary deposits in the Kyeenec I
rockshelter contain evidence pointing to late
Holocene human occupation commencing al
about 1500y BP. It does not necessarily imply the
earliest human use of the rockshelter. The depos-
its formed through a combination of aeolian ac-
tivity which bought sand and dust inside the
rockshelter, there to combine with roofall and

652

TABLE 7:Frequency, size and mass of artefacts recov-
ered from excavation 19, Kycence I Rockshelter. Ab-
breviations: XU=Excavation Unit; ML=Mean length;
MW - Mean width; MM — Mean mass.

a ia i |

пол |Fake — 2 | вэ [us |
ааа аа 0а 90

Prep eho be |

[Flaked piece [3 | | |
E m m а E
P Emm pe je e
| jFakdpice |: | | | — [201
[9 [Flake {5 [i21 [us |ss jos |

naar
[urbane ||. har бо аз |
[196 Flake а [m3 [igs ja? јав |

Retouched
| (еее п [ез pes [ias [o7 |

рэл [Fike — — |5 [125 i26 |44 [jia |
135 [soo |az9 [145.1 |
|| o2 |

Ira a ра га
jo [m2 зэ аз 142 |

Imm

Core — |! |s$3 |560 [182 [606 |
sna [Pake — |4 |272 |223 |74 |247 |
|j/]3 |Flake | 87 [16 |

= Lu
ona [rake — isa fu из |
BEN TO ав |05 јав ри

occupationa] material including stone and
wooden artefacts and charcoal. The faunal and
floral material found in the deposit has an equiv-
ocal history and may derive from human and/or
non-human sources,

This shelter contains evidence of a range of
human activities including stone and wooden ar-
tefact manufacture and maintenance, and burn-
ing. The large amounts of charcoal in the deposit
indicate major episodes of buming. These epi-
sodes of burning appear to be unrelated to those
of woodworking or stone artefact manufacture,
maintenance or discard as the bunung has not
affected the other forms of evidence. The inter-

MEMOIRS OF THE QUEENSLAND MUSEUM

pretation of episodic burning events is reinforced
by the preservation of floral and faunal remains
that also appear to be unaffected by the burning.
The infrequent and episodic human occupation is
indicated by the generally sparse nature of the
evidence of occupation at the shelter, The argu-
ment for episodic use of this site by people is
further strengthened by the faunal evidence. If the
faunal remains are non-humanly derived it impl-
ies that visits by humans were sufficiently infre-
quent for other animals to have occupied the site
in their absence.

The area immediately surrounding this site ex-
pands the evidence of usage. The presence of the
stone arrangement opposite the shelter indicates
that the area was probably used for ceremonial
activities. On the plateau surrounding the site, the
extensive and dense artefact scatter argues for the
secular use of stone, and if density of artefacts can
be related to intensity of use this area was used
more frequently than the rockshelter. Artefact
maintenance and manufacture represents a signif-
icant proportion of that use.

This site can be interpreted as one that has
played host to people who have carried out a
variety of tasks at the site during short-term,
episodic visits.

KYEENEE П ROCKSHELTER

Kyeence Il rockshelter is located on the castern
side of a small outlier which is in a pocket of
gently undulating mulga plains (Fig. 6). At ap-
proximately 2km from the rockshelter, Dissected
Residuals encircle this plain on its eastern, north-
ern and western sides. The ephemeral, dendritic
drainage channels of Werie Ella Creek, à tribu-
tary of Benanga Creek, have formed approxi-
mately 1km the west.

The outlier, formed in Winton Formation bed-
rock, is approximately 250m long on its north-
south axis and 74m wide at its widest point.
Resistant bedrock has resulted in a small 3m high
escarpment in which three rockshelters have
formed (Fig. 20). The steep escarpment in which
the rockshelter is located gives way to a gentle
scree slope. This scree slope consists of boulders
of weathered mudstone and siltstone, lithosols
and shallow red earths supporting gidgee (Acacia
cambagei tall open mulga (Acacia aneura)
shrubland (Dawson, 1974),

The rockshelter entrance is low (1.6m) and
wide (6.4m) and 2m from the entrance opens out
to 7.6m. Three metres from the entrance this
chamber constricts to a small, 80cm wide hole

ROCKSHELTER TEST EXCAVATIONS

Inner chamber

/

/
Entrance hole

E a

—1

—

s MENT
wf Datum
=

ü 2m

ae m"———

En ds

** Bedrock
Ca + Exeavation square
Foen lines ah б 5m immessi

FIG. 21. Plan of Kyeenee II rockshelter

that provides access to an inner chamber that is
4m wide and 2m deep (Fig. 21). The roof of the
cave is low throughout, with a maximum height
of 1.6m near the dripline (Fig. 22). Swallows nest
on the ceiling, bats roost in the inner chamber and
kangaroos and wallabies use the outer chamber.

Although other rockshelters in the outlier con-
tained deposits with charcoal and artefacts on the
surface, they had been extensively disturbed by
animals. KRS II had a similar amount of deposit,
but appeared to be less disturbed. It was selected
forexcavation on this basis. The rockshelter floor
consists of fine grey sediment mixed with roof
fall. In parts of the floor, bedrock was exposed.

At the time of investigation, two flakes and
some charcoal were observed on the rockshelter
floor. A number of artefacts were also scattered
down the scree slope. Although the topography
of the rockshelter indicated that the floor deposits
were unlikely to be thick, it was hoped that its
excavation would provide some in situ, dateable
evidence of occupation and add to knowledge of
the character of deposits in the region.

EXCAVATION PROCEDURE
After positioning a datum point outside the
shelter, an alpha-numeric 50 x 50cm horizontal

663

grid was established for the floor using 90" offsets
from the baseline. Heights were determined with
autoset and water levels, One 50cm square, C3,
was selected for excavation to obtain the maxi-
mum depth in the least disturbed deposit. It was
then excavated following the procedures and pro-
forma outlined by Johnson (1979). The excavated
material was sieved through 2mm and 4mm wire
mesh. All the material retained in the sieves was
sorted on site, The >4mm material was sorted a
second time on site and returned as backfill, The
2-4mm fraction was wet sieved, A grab sample
(100g) of the material that had passed through
both sieves was retained for further analysis. The
excavated pit was lined with plastic and back-
filled with rocks and excavated deposit.

Two 10m squares were laid out from the base-
line to extend down the talus slope. All the arte-
facts occurring within these quadrats were
recorded in a manner similar to that undertaken
for the surface and other excavated sites,

DEPOSIT DESCRIPTION

СЗ had a mean deposit depth of 11cm. Thirty
seven kilograms of deposit was excavated in
three XUs. The deposit consisted of fine grey
sediment mixed with roof fall. The >2mm sieved
fraction represented between 30% and 35% of the
mass of the deposit throughout, Charcoal is pres-

Entrance hola —

Section B - B:

Saw

n

Boni ck

FIG. 22. Topographic profiles across Kyeenee Il
rockshelter

ent only in minor amounts but a trend of decreas-
ing content towards the top was detected. The
base of the deposit rested on uneven and weath-
ered bedrock. à

Detailed analysis of the deposit did not proceed
because no artefacts пг other cultural matenal
were recovered.

ASSOCIATED ARCHAEOLOGICAL EVIDENCE

On the talus slope, the 10m square laid out from
the baseline contained 38 silcrete artefacts. The
artefact types recorded included 25 flakes, 9 re-
touched flakes, 2 single platform cores, 1 multi-
platform core and 1 tula, The maximum artefact
density was 3/m?. All the flakes had lengths of
between 20mm and 65mm. The cores ranged in
size from 15mm to 75mm.

The adjoining quadrat contained 87 silerete
artefacts including 59 flakes, 7 retouched flakes,
13 cores and 8 multi-platform cores. All were
made on silcrete. Flake lengths ranged from be-
tween 25mm and 90mm. Eighty percent of these
were between 25mm and 50mm in length. The
cores ranged in size from 30mm to 75mm.

SUMMARY

The sediments in and around this rockshelter
contain limited evidence of human occupation.

The deposit is shallow and in parts disturbed.
Numerous artefacts litter the scree slope in front
of the shelter indicating the possibility that either
the slope was the preferred area for human activ-
ities or that the deposit in the rock shelter has been
purged, perhaps a number of times, and the con-
tents of the shelter strewn over the slope.

DISCUSSION

Although only a smali sample, the excavation
of these three rockshelters provides some basic
evidence about the nature of archacological de-
posits in rockshelters in southwest Queensland.

In each case, the rockshelters had been well
formed, ranging from 4m to over 12m in depth.
All were formed in the sediments of the Winton
Formation which were capped with tertiary
silerete bedrock or gibber. The depth of deposit
in each case was relatively shallow, ranging from
6-80cm. The rockshelter directly associated with
a source of temporary water had the deepest and
culturally richest deposit. The deposits of the
rockshelters that were 2 considerable distance
from water sources (Murderers Bore 1 and
Kyeenee Rockshelter 2) were shallower and had
little cultural evidence associated with them, al-

MEMOIRS OF THE QUEENSLAND MUSEUM

though the talus slopes in front of each were
littered with stone artefacts, indicating some cul-
tural use. The two deposits that were dated were
late Holocene in age (450+70уВР and
1580+60yBP).

These investigations confirm the impression
that the rockshelters of southwest Queensland are
unlikely to contain deposits that are old, cultur-
ally rich or deep. Rockshelters that are consider-
able distances from sources of permanent or
semi-permanent water will be less likely to con-
tain deposits than those that are close to a water
source, However, Kyeenee I rockshelter had ex-
cellent preservation conditions and a range of
organic material had been preserved in it. While
these rockshelters may nol represent а good
source for obtaining long chronological se-
quences, under the right circumstances they may
well provide a useful source of information for
late Holocene Aboriginal occupation of this land-

scape,

ACKNOWLEDGEMENTS

This project would not have been possible with-
out the support and assistance of Greg Sherwin
from Kilcowera and Colin Vagg from Kycence.
Their co-operation and assistance for what must
have seemed a never ending saga is much appre-
ciated. This project would not have been possible
without thc help in the field of Julia Findlay, Rob
Neale, Andrew Border and Richard McKellar,
The forbearance and support of my colleagues
Michael! Quinnell, Tina Baum and Judith Bartlett
is appreciated, Steve Van Dyke assisted with
mammal identification, Patrick Couper with rep-
tile tdentification, and the Queensland Herbarium
with plant identification. Grant McTainsh and
Errol Stock provided useful comments and ad-
vice on the soil samples, Peter Hiscock, Peter

‘eth and Mike Smith kindly provided unpub-
lished data and references. Natalie Franklin pro-
viced access to information from DEH records.
Tracy Adams undertook the soil analysts. Grant
McTainsh, J. Hall, Glen Ingram, Errol Stock and
Bruno David read drafts and provided comment
and advice. John Richter produced the
illustrations. Vedette MeGuire and Jody Niethe
typed various drafts of the manuscript. 1 would
like to thank the Queensland Museum Board and
Peter Jell (Assistant Director, Research) for their
generous support for this project. The Australian
Heritage Commission, through the National Es-
tate Grants Programme, contributed funds.

ROCKSHELTER TEST EXCAVATIONS

LITERATURE CITED

ALFREDSON, С. 1987, "Repon ом the Archaeological
Survey of the Proposed Pipeline from Cooroo to
Jackson’. (Unpublished Report for Delhi Petro-
lean: Brisbane).

DAVID, В, 1991. Fern Cave, rock art and social forma-
tions: Rock art regionalisation and demographic
models in southeastern Cape York Peninsula. Ar-
chaeology in Oceania 26:41 -57.

DAWSON, N.M. 1974. Land Systems, Pp,74-83. In
"Western arid landuse study, Part I’, Queensland

rtment of Primary Industry, Division of Land
Utilisation Technical Bulletin 12.

ELKIN, A.P. 1949 The origin and interpretation of
petroglyphs in south-east Australia, Oceania
20(2):119-157.

FLOOD, J.. DAVID, B., MAGEE, J. & ENGLISH, B.
1987. Birriigai: A Pleistocene sile in Ше south-
eastern Highlands. Archaeology іп Oceania
22(1):9-26.

HALL, J, 1986. Exploratory excavation at Bushrangers
Cave (Site LA:B75), a 6000-year-ald campsite in
Southeast Queensland: Preliminary results. Aus-
tralian Archaeology 22:88-103.

HALL, J., GILLIESON, D.S. & HISCOCK, Р. 1988,
Platypus Rock Shelter (KB:A70), S.E, Queens-
land: Stratigraphy, chronology and site formation,
Queensland Archaeological Research 5:4-24.

HISCOCK, Р. 1984а. Archaeological survey, Jackson
to Noccowlah Coridor and associated develop-
ment. (Unpublished Report, ANU Archaeological
ponas for Hollingsworth Consultants; Bris-

ane).

1984b, Preliminary report on the stone artefacts
from Colless Creek Cave, northwest Queensland,
Queensland Archaeological Research 1. 120-151.

1985, Naccowlah Block Baseline Archaeological
Study. (Unpublished Report to Kunja Aboriginal
Consultants: Brisbane).

1986, A Technological analysis of stone artefact
assemblages from the Hunter River Valley re-
gion: NSW National Parks and Wildlife Service
Hunter River Valley region archaeology project
stage ] (Unpublished Report to NSW National
Parks and Wildlife Service: Sydney),

1990. A study in scarlet: Taphonomy of inorganic
artefacts. Pp,34-49, In Solomon, S., Davidson, І,
& Watson, D, (eds), Problem solving in tapho-
nomy, Tempus Vol.2 (Anthropology Muscum,
University of Queensland; St Lucia).

HISCOCK, P. & HALL, J. 1988. Technological Change
at Bushrangers Cave (LA:A 11) S.E. Old. Queens-
land Archaeological Research 5:90-1 12.

HISCOCK, P. & HUGHES, P.J, 1983, One method of
recording stone artefact scatters during site sur-
veys. Australian Archaeology 17:87-98.

HUGHES, PJ, & LAMPERT, R.J. 1980. Pleistocene
occupation of the and zone in south-east Australia:
Research prospects for the Cooper Creck-

Strzeleckà Desen region. Australian Archaeology

10:52-67,

JOHNSON, 1. 1979. “The Getting of Data’. Unpub-
lished Ph.D, thesis, (Australian National Univer-
sity, Canberra).

LANCE, A. 1990. An archacological survey of the
proposed Yanda-Noccowlah pipeline. (Unpub-
lished Report to Santos Pty Ltd; Adelaide).

1992, ‘An archaeological survey of the Patroclus
pipeline southwest Queensland’. (Unpublished
Report to Santos, Pty Ltd, Adelaide).

LANCE, A. & HUGHES, P, 1983. Archaeological sur-
vey of the revised route of the Jackson Oilfield
access road, (Unpublished Report tù
Hollingsworth Consultants; Brisbane),

LEWIS, J. 1966. Cuttaburra campfires, Walkabout
32(2):35-38,

LILLEY, LA. 1983, An Archaeological Study of the
Jackson Oil Project. (Unpublished Report to
Hollingsworth Consultants; Brisbane),

McKELLAR, H. 1984. Marya-Mundi- A history of the
Aboriginal people of soulh west Queensland,
{Cunnamulla Native Welfare Association}
Cunnamulla).

MORWOOD, М.Ј, 1986, The Archaeology of Art:
Excavations at Maidenwell and Gatton Shelters,
SE Queensland, Queensland Archaeological Re-
search 3:88-132.

MUNSELL COLOR COMPANY 1990. Munsell soil
color charts (Revised Edition), (Munsell Color
Company: Baltimore, Maryland).

RICHARDSON, N. 1983, An Archaeological Survey
of the Jackson Oilfield Access Road, ety
M Repon to Hollingsworth Consultants: Bris-

апе),

RIDDELL, Р.О. 1928. Ordered arrangement of stones
by Aborigines at Durham Downs, Cooper's
Creek, Queensland, Transaction of the Royal So-
ciely of South Australia 52:165-166,

ROBINS, R.P, 1977, A discussion of two engraving
sites in southwest Queensland. Anthropology So-
cicty of Queensland Newsletter 111:1-2.

1981, Four Aboriginal dwelling sites in south- west
Queensland, Australian Archaeology 11:79-90.

ROBINS, R. 1993. Archaeology and the Currawinya
Lakes: Towards a Prehistory of Arid Lands of
Southwest Queensland. Unpublished Ph.D. the-
sis. (Griffith University: Brisbane).

ROWLANDS, RJ, & ROWLANDS, J.M. 1976. Report
af a visit to north-western New Souih Wales and
south-weslem Queensland. (Unpublished report
to the Australian Institute of Aboriginal Stunbes:
Canberra).

1978, Visit to western New South Wales and south-
western Queensland, June, 1966. (Unpublished
Repost, Australian Instirute of Aboriginal Stud-
ies: Canberra).

SMITH, MA. WILLIAMS, E. & WASSON, R.J.
1991, The JSN site: some implications for the
dynamics of human occupation in the Strzelecki

666 MEMOIRS OF THE QUEENSLAND MUSEUM

Desert during the late Pleistocene. Records of the Creek. Records of the South Australian Museum
South Australian Museum 25(2):175-192. 24:43-66.

STOCKTON, E.D. 1977. Pipeline survey of Aboriginal WHITE, P.J. & O'CONNELL, J.F. 1982. A prehistory
relics in northwestern New South Wales and of Australia, New Guinea and Sahul. (Academic
southwestern Queensland. Australian Archaeol- Press: Sydney)
ogy 7:7-17. н ,

VETH, P. & HAM, C. 1989. The archaeological signif- WILLIAMS, E. 1988. The archacology of the Cooper
: : Basin: Report on fieldwork. Records of the South
icance of the Lower Cooper Creek. (Unpublished À 22:53-62
Report to the Division of Anthropology, South Australian Museum 22:53-62.

Australian Museum: Adelaide). WITTER, D.C. 1992. Regions and resources. (Unpub-

VETH, P., HAMM, G. & LAMPERT, R.J. 1990. The lished Ph.D. Thesis: Australian National Univer-
archaeological significance of the Lower Cooper sity: Canberra).

GLOSSARY

Definitions for Stone Artefacts

Flake: A flake was defined as a piece of stone struck from a core and which exhibited characteristics which
indicated that it had been struck. These included features associated with the point of force application (PFA)
manifest principally in the presence of a ringcrack, a bulb of force on the ventral surface and other surface
deformities including ripples, and an eraillure scar (Hiscock, 1986).

Core; A piece of stone with no positive flake scars but with one or more negative flake scars (Hiscock, 1986).
Retouched flake: A flake which has had other flakes removed from it leaving scars onto the ventral face and/or
deriving from the ventral surface (Hiscock, 1984).

Flaked piece: A chipped artefact which is missing the defining attributes of a core, flake or retouched flake.
Some weathered, fractured or heat shattered artefacts are identifiable only to this level. This category is only used
when the artefact had definitely been chipped, but its identification was uncertain (Hiscock, 1984).

Single platform core: A core from which all flakes have been struck from the same surface (the platform).
Multi-platform core: A core which has been rotated in the course of flaking so that flakes are struck from two
or more surfaces. Commonly, flakes will be struck from negative flake scars on the core,

Snapped Flake: A flake which has been fractured transversely, These usually are elongate flakes (before fracture).
The fracture surface is approximately perpendicular to the natural surface and often has a S-shaped undulation.
Tula: A flake with a convex ventral face and a large angle between the ventral surface and the platform. It has
steep retouching at the distal end that extends around to the margins. The platforms are generally large and the
flake is thick ventrally/dorsally (Hiscock & Hughes, 1983:96).

OBSERVATIONS ON THE EARLY LIFE HISTORY STAGES OF NOTADEN
BENNETTII IN THE CHINCHILLA AREA OF SOUTHERN QUEENSLAND

M. SHARMAN, 1, WILLIAMSON & D.S.L. RAMSEY

Sharman, M., Williamson, I. & Ramsey, D.S.L. 1995 12 01: Observations on the сапу life
history stages of Nofaden bennettii in the Chinchilla area of southem Queensland. Memoirs
of the Queensland Museum 38(2):667~-669. Brisbane. ISSN 0079-8835.

Epgs, larvae and metamorphs of Notaden bennettii were observed in natural pools in the
Chinchilla area, southeast Queensland between February and March 1994. The observations
suggest that V. bennetrii has some interesting behavioural and morphological characteristics
that are absentorrare іп other Australian anurans. [ ] Notaden bennettii, life history, southeast
Queensland.

М. Sharman, Í, Williamson & D. S. L. Ramsey, School of Life Science, Queensland University

of Technology, GPO Box 2434, Brisbane, Qld 4001, Australia; 27 September 1995.

Notaden bennettii is a common frog found over
a large area of inland eastern Australia (Cogger,
1992). However there is little information on the
early life history stages of this species. We report
here on observations of the eggs, larvae and
metamorphs of №. bennettii made in the Chin-
chilla area of southern Queensland.

Observations were made at a site within the
Barakula State Forest, approximately 40km north
of Chinchilla іп southern Queensland. Vegetation
in the area is mainly open woodland dominated
by white cypress (Callitris glauca), Observations
were made in situ, or on live individuals trans-
ported to the laboratory. Descriptive terminol-
ogy for tadpoles follows Altig (1970) and the
staging system used is from Gosner (1960). Mea-
surements of tadpoles were made to the nearest
0.5mm under a dissecting microscope, and
me s were measured using dial calipers.
Water temperatures were recorded with maxi-
mum-minimum thermometers.

A number of permanent and temporary water
bodies were monitored for calling males, eggs,
larvae and metamorphs approximately once per
week from November 1994 to March 1995.
Notaden bennettii were heard calling on January
30, February 8 and February 14. Amplexed pairs
were located on February 14 following 70mm of
rain in the previous 48 hours. Amplexus was
pelvic and pairs were seen in shallow temporary
pools (10-20cm deep). Egg laying was not ob-
served, but 15 egg masses were noted 1n four
separate pools the following moming. Each mass
was oval in shape and measured approximately
10x6cm. Masses were free floating, possibly due
to a number of air bubbles distributed throughout
the mass, and contained approximately 500 eggs
in individual jelly capsules. In 13 of the 15 egg

masses a 2-3cm wide band of vegetation (mainly
dead Cypress needles) was found around the pe-
rimeter of the egg mass (Fig. 1). The remaining
two masses had only small amounts of vegetation
around their perimeters. Because egg laying was
not observed it is not clear whether each mass
represents one clutch, and if adults placed the
vegetation around the egg mass or if vegetation
accumulated around the egg mass as it drifted in
the pool. However, the small interval between
observing amplexus and resulting egg masses
(«12h) and the consistent arrangement of vegeta-
tion suggest that the vegetation was placed
around the egg mass, perhaps to act as a barrier
against egg predators.

Embryos in natural pools had hatched by 1200
on February 17 (approximately 60h). Embryos
transported to the laboratory had reached stage 16

FIG. 1, Typical egg mass of Notaden bennestii, Masses
measured approximately IOx6cm-

FIG. 2. Lateral view of a stage 37 Notaden bennettii
tadpole. Scale bar = 10mm.

after approximately 36 hours, and stage 20 after
approximately 60 hours. Temperature was not
controlled during this period, but ranged between
20° and 30°С. Stage 26 tadpoles ranged in length
from 7.5 to 10.0mm, and maximum length at-
tained was 38.0mm (at stage 40). A stage 37
tadpole is illystrated in Fig. 2. Ratios of body
proportions were body length : total length = 0.42
(п. = 20), body width : body length 20.63 (n= 10),
and body depth to body width = 0.61 (n = 10),
Eyes were dorsal and the mouth was orientated
ventrally. There were three upper and three lower
rows of labial teeth with the formula 3(2-3)/3,
with the A2 gap being narrow and the A3 gap
wide (Fig. 3). Marginal papillae had an anterior
gap. The anus was median and the spiracle was
sinistral, located ventrolaterally and orientated
posteriorly. Tadpoles between stages 27 and 38
generally had a small pale spot mid-dorsally. The
typical N. bennettii back pattern became evident
at stage 37 to 38. A darkened posterior tail section
developed from a slight difference in the distri-
bution of dark pigment at stages 27 to 30, to a

gendi PRETI gs
a ~
yt umma mme"
Lan MO A
P. (NS "m ; S mea, em
fe D / Ma 5
Den ENG
а ' 3
c SUSU ger rre tmm sta а 9 »
= Д rset mm navem rat >
ER FH eal FT

МАА T US M PIT

FIG. 3. The oral disc of a stage 37 Noladen bennettii
tadpole. Scale bar = 1mm,

MEMOIRS OF THE QUEENSLAND MUSEUM

distinct dark posterior section in tadpoles beyond
stage 36 (Fig. 2). Dark tail tips, with adjacent light
areas, may function to misdirect the attack of
predators (Altig & Channing, 1993), especially
odonate naiads (Caldwell, 1982). Many tadpoles
from natural pools had tail damage consistent
with attack by odonates, and odonates were com-
mon in pools with N. bennettii,

In the field tadpoles were noted in the shallow
water at the edge of pools at all times of the day,
Recently metamorphosed individuals ranged in
size from 11.0 to 14.9mm (mean = 12.7, п = 18).
Metamorphs were noted near pools from Febru-
ary 2810 March 14. The length of the larval period
in natural ponds is difficult to determine because
pools with tadpoles of known age dried before
individuals metamorphosed. Water temperatures
in these pools ranged from 19° to 36°C. Some
larvae in an experimental pond reached metamor-
phosis in 50 days (temperature range 19° to
36°C.). However, egg and larval period may be
as short as 28 to 30 days if eggs were laid when
frogs were first heard calling (January 30) and the
metamorphs emerging from those eggs were the
ones noted on February 28.

Metamorphs were active dunng the day and
appeared to bask in sunny positions on the damp
substrate. The meat ant, Iridomyrmex purpureus,
was seen preying On some metamorphs. Other
anuran species, noted as either calling males or as
larvae, that used the same water bodies as N.
bennettii were Litoria alboguttatus, L. fallax, L.
latopalmata, L. peronii, L. rubella, Cyclorana
cultripes, Limnodynastes ornatus, L, tasmanien-
sis, L, terraereginae, Crinia parinsignifera and
Uperolea rugosa.

Slater & Main (1963) described the tadpole of
Notaden nichollsi. However, information on the
life history of the genus Notaden is limited. The
observations presented here suggest that Notaden
bennettii has some interesting behavioural (egg
laying behaviour) and morphological (tàil colour
pattern) characteristics that are absent or rare in
other Australian anurans.

Observations were made during field work con-
ducted for the CSIRO - ANZECC Cane Toad
Research Committee as part of a grant to T, Wil-
liamson. Steve Richards and Gien Ingram pro-
vided useful comments.

LITERATURE CITED

ALTIG, К 1970. A key to the tadpoles of the continental
tes States and Canada. Herpetological 26:
1 2

van i.

NOTADEN BENNETTII 669

ALTIG, В. & CHANNING, A. 1993. Hypothesis: func- GOSNER, K.L. 1960. A simplified table for staging
tional significance of scaler and pattern of anuran

tadpoles. Herpetological Journal 3: 73-75. anuran embroys and larvae with notes on their

CALDWELL, J.P. 1982. Disruptive : a tail scaler poly- identification. Herpetologica 16, 183-90. _
morphism in Acris tadpoles in response to differ- SLATER, P. & MAIN, A.R. 1963. Notes on the biology
uy рота, Canadian Journal of Zoology 60: of Notaden nichollsi Parker (Anura:

COGGER d HG. 1992. ‘Reptiles and Amphibians of Leptodactylidae). Western Australian Naturalist

Australia’ 3rd Edn. (A H & A W Reed: Sydney). 8: 163-166.

670

FIRST RECORD OF THE LONGFIN MAKO (ISURUS
PAUCUS) FROM AUSTRALIAN WATERS. Memoirs of
the Queensland Museum 38(2): 670. 1995:- The longfin mako
Isurus. paucus Guitart Manday, 1966 is a poorly known
epipelagic tropical and warm-temperate shark which has been
most frequently documented from the western North Atlantic
(Compagno 1984). It has also been recorded from the eastern
North Atlantic, central, western and western North Pacilic and
the western Indian Ocean off Madagascar (Fourmanoir &
Laboute, 1976; Compagno, 1984; Taniuchi, 1984) but proba-
bly has a more extensive distribution In warm seas than these
records suggest. Compagno (1984) noted that it is probably
taken regularly in the Japanese tropical pelagic longline fish-
ery. Last & Stevens (1994), in their guide to the sharks and
rays of Australia, commented that this species 'still remains
unrecorded locally but almost certainly occurs in oceanic
waters off northern Australia'.

The apparent ranty of this species may in part be due to
confusion with the shortfin тако Isurus oxyrinchus Rafin-
esque, 1809 from which it differs principally in its longer
pectoral fins (about as long as head length), ventral head
colaration and slimmer body.

The longfin тако has been reported lo reach a maximum
length of 417cm total length (TL); the size at birth is about
97cm and males and females are mature by 245cm TL.

Between 24 July and 27 August 1994, observers who
regularly monitor the catches of Japanese longliners Inside the
Australian Fishing Zone reported several longfin makos as
follows: 24 July at 32S, 154 E, $ , approx. 230cm TL, surface
water temp.(WT) 19,8-21.1'C; 14 August at 14'S, 149'E,
approx. 200cm TL, WT 23.8-24.0'C; 16 August at 14'S,
148" E, approx. 200cm TL, WT 23.7-23.9°C:27 August, а!
15" L6'S, L51 16 E, at a hook depth of 50-145m, WT 24.6'C;
9 September at 13°9'S, 150°18'Е, 9, 201em TL, 56,5kg, hook
depth 60- 190m, WT 24 (У, Full proportional measurments are
available from the senior author.

Photographs of one specimen confirmed their identifica-
Gon (D. Heran, Foreign Fishing Observer Station, pers.
comm.), Or 9 September 1994, в 201em TL specimen (Fig.
1) was caught by a longliner and retained by the observer
(M.Scott) onboard at the time, This shark was subsequenily
sent to Hobart, photographed and preserved in the collection
of the Tasmanian Museum and Art Gallery (Registration
Number TMD 2241), Proportional dimensions of this speci-
men (afler Compagno 1984) have been compared with those
from the Indo-Pacific and central Pacific (Garrick 1967) and
eastern North Atlantic (Moreno & Moron 1992). The only
significant differences are that an (he Australian specimen, the
pectoral fins were 103.7% of head length (measured to the

MEMOIRS OF THE QUEENSLAND MUSEUM

pectoral fim drign) compared to а mean value of 102,096 for
Garrick's (1967) specimens and 95.5% for Moreno and
Moron's (1992) fish. Furthermore the Australian individual
appeared to have a shorter snout than other compared speci-
mens, however, this could be due to differences in measuring
technique.

Acknowledgements

We are grateful to the Master and crew of the 'Vision' for
transporting the longfin mako from the ‘Kinsie Maru 5° to
Cairns, Bob Lameson for arranging freight of the specimen
from Cairs to Hobart, Wade Whitelaw and Noel Kemp for
assistance with logistics, Grant West for help with the mea-
surements and, together with Peter Last, for comments on the
manuscript, Thor Carter for taking the photograph, the Com-
monwealth Observer Program for data on additional longfin
mako specimens and the Master and crew of the "Kinsei Maru
5", who caught the shark.

References

Compagno, L.J.V. 1984. FAO species catalogue. Vol.4,
Sharks of the world. An annotated and illustrated cata-
logue of shark species known to date, Part ] -
Hexanchiformes to Lamniformes: viii, 1-250. FAO
Fisheries Synopsis 125: 1-250,

Fourmanoir, P. & Laboute, P. 1976. Poissons de Nouvelle
Caledonie et dés Nouvelles Hebrides. (Les Editions du
Pacifique: Papeete, Tahiti) 376pp.

Garrick, J.A.F, 1967. Revision of sharks of genus Isurus with
description of a new species (Galeoidea, Lamnidae).
Proceedings of ihe United States National Museum 118
(3537):663-690.

Last, Р.Е. 1994. Sharks and rays of Australia. (CSIRO: Aus-
tralia),

Mareno, J.A. & Moron, J. 1992. Comparative study of the
genus Isurus (Rafinesque, 1810), and description af a
form (‘Marrajo Criollo’) apparently endemic to the
Azores, Рр, 109-122. In Pepperell, Ј.С. (ei) Sharks:
Biology and fisheries. (CSIRO; Australia).

Taniuchi, T, 1984, Distribution of elasmobranchs in Choshi,
Tzu Islands, and Ogasawara Islands. Report of a prelim-
inary investigation on sharks and rays in the western
Pacific Ocean. Japanese Group for £lasmobranch Stud-
ies Special Publication No.1; 14-24.

JD, Stevens, CSIRO Division of Fisheries, Marine Labara-

tories, GPO Box 1538, Hobart, Tasmania 7001, Australia; M.

Scoll, Foreign Fishing Observer Section, Queensland Depari-

ment of Primary Industries, GPO Box 2454, Brisbane,

Queensland 4001, Australia; DI July 1995

Vig. |. Ieurus paucur, 2, ler TL, captured at 15*09*S, T0" 187B off eastern Australi,

RE-APPEARANCE OF THE BLUE MUDHOPPER, SCARTELAOS HISTOPHORUS
(PISCES: GOBIIDAE) IN THE GREATER BRISBANE AREA

К.А. TOWNSEND & 1А. TIBBETTS

Townsend К.А. & Tibbetts LR. 1995 12 01; Re-appearance of the blue mudhopper,
Scartelaos histophorus (Pisces: Gobiidae) in the greater Brisbane area. Memoirs of the
Queensland Museum 38(2):67 1-676. Brisbane. ISSN 0079-8835.

Mudskippers have not been reported іп the greater Brisbane area since 1919 (McCulloch &
Ogilby, 1919), Recently, large populations of these animals were observed along the banks
of the Brisbane River, A survey of the mudskippers in the greater Brisbane arca was initiated
and only the blue mud hopper, Scartelaos histophorus was identified. It has a discontinuous
distribution from the mouth of the Brisbane River upstream to the suburb of Fig Tree Pocket
and as far south as the suburb of Lota, Preliminary information on seasonal change in
abundance for S. histophorus was obtained during this study. Future research areas based
upon mudskippers as biological indicators are outlined. [ ] Scartelaos, ecology, distribution.

К.А. Townsend, Department of Zoology. University of Queensland 4072, Australia; LR.
Tibbetts, School of Marine Science, University of Queensland, Brisbane 4072, Australia; 27

September 1995.

Mudskippers arc amphibious gobiids found in
the muddy intertidal zones of tropical and sub-
tropical coasts and estuaries. They are distributed
from northern Australia to as far north as Kuwait
Bay, including the east and west coasts of Africa,
Melanesia, Micronesia. India, Polynesia and the
West Indies (Stead, 1906; McCulloch & Ogilby,
1919; Marshall, 1966). The members of the
Gobiidae that are commonly known as mudskip-
pers are divided into the Periophthalminae and
the Apocrypteinae; the former being more terri-
torial. Representatives from both subfamilies
have been recorded in Australia (McCulloch &
Ogilby, 1919; Weber & DeBeaufort, 1953; Mar-
shall, 1966; Milward, 1974; Grant, 1985).

The first record of mudskippers tn the Brisbane
area was of Periophthalmus vulgaris Eggert.
1935 (given as P. koelreuteri (Pallas)) in 1877
(Castelnau, 1878). Since that time no other study,
including the comprehensive study conducted by
Milward between 1964 and 1974 along the
Queensland coast, had noted this species in the
greater Brisbane area. Milward (1974) found the
southern limit of P. vulgaris distribution to be
Burnett River Heads (latitude c. 24245'5).

In 1919 another species (Scartelaos
histophorus (Cuvier & Valenciennes, 1837)) was
noted in Moreton Bay and along the Brisbane
River; identified as 5, viridis by McCulloch and
Ogilby (1919). This was the only report of these
mudskippers in Australia until 1966 when a sur-
vey which included all of the eastern Queensland
coast, noted 5. histopliorus to have à distribution
no further south than Bundaberg (lat. 24°53°S)
(Marshall, 1966). Milward's (1974) study ex-

tended the species’ range to Urangan (lat. 25°15"
S) in Queensland (Milward, 1974). Despite an
extensive search for mudskippers in the greater
Brisbane area, none were found (№. Milward,
pers comm.).

The objectives of this study were two fold,
firstly to identify the species of mudskippers
found in the greater Brisbane area and secondly
to survey the distribution and zonation of these
species.

METHODS AND MATERIALS

Surveys of the distribution were carried out by
visiting river and bay shores of western Moreton
Bay, south east Queensland (27°22'S, 153? 10"
E). Additional information was obtained through
interviews and reports supplied by members of
the public, Brisbane Transport and university
staff in response to newspaper articles. Written
correspondence from the public was followed up
by an interview by the principal investigator to
confirm the identification of the animals, as often
intertidal fish such as the peacock blenny
(Istiblennius meleagris (Valenciennes)), family
Blenniidae, were identified as mudskippers. This
interview was then followed up with a visit to the
reported location to confirm the distribution re-
port. Through this process rhe general habitat of
the animals was observed and potential sites for
mudskipper populations identified.

Walking through the mudflats caused some
difficulties as the animals were efficient at detect-
ing movement which caused them to quickly
reireal down their burrows. This made positive

672 MEMOIRS OF THE QUEENSLAND MUSEUM

A Reef Point

Redcliffe

27°15

Bramble Bay

A
A

Nudgee Beach A Mud Island

хо
А St. Helena Island
SQ fisherman
4 Island
*

Brisbane River
o

Wynnum Green Island

; o Doboy Creek Waterloo
Botanical Gardens
Bulimba Bay Moreton Bay
Orleigh Park South Bank | Lota
Dutton Park Ferry Terminal
A
o Oxley Creek
Fig Tree Pocket
Key:
Confirmed sightings..
Potential habitats Redland Bay

Surveyed areas and none found.
Location of zonation study...
Site of animal capture.

Albert River
>

FIG. 1. Study sites and distribution of Scartelaos histophorus in the greater Brisbane area.

REAPPEARANCE OF BLUE MUDHOPPER

identification of the animals in the field difficult,
specimens therefore had to be secured, The extent
of the local distribution may have been under
estimated as a result of disturbance to populations
caused by the observer before the animals could
be detected.

Many methods were employed in the attempts
10 capture specimens for identification. Рам au-
thors have noted this difficulty. D.G. Stead
(1906) wrote "the capture of the little fish seems
at first to be an easy task, but woe betide the
reckless enthusiast who ventures on the treacher-
ous ooze in its vain pursuit. He wil] emerge from
the enterprise with bemired raiment and a much-
chastened spirit!". Milward (1974) also tried
many methods and was reduced to "patience and
guile” to capture the fish, where as Stebbins and
Kalk (1961) stunned the creatures using "rubber
bands cut from automobile tires and fired from
the thumb", For this study, attempts were made
using a throw net, a nylon noose, a dip net and by
Simply allempting to chase and pounce upon
them. The method that proved most successful
consisted of stalking the animals at night during
low tide with the aid of a spot light. Once dazzled
the fish were scooped up using a bucket. The
animals used for identification m the laboratory
were captured by spotlighting in the Lota area.

The identification of the captured animals was
done using a taxonomic key prepared by
Milward, 1974. An ccological key developed
from Milward (1974) and plates from Milward
(1974) and Grant (1993) assisted with long range
identification,

RESULTS

Spatial distribution: Only one species of mud-
skipper, Scartelaos histophorus (^pocrypteinae),
was found in the Brisbane area during this study,
Morphological characteristics of captive speci-
mens were in agreement with the description
given in the taxonomic key devised by Milward
(1974). The body of the animal is elongate with
very small scales. The dentition consists of sub-
horizontal teeth in the lower jaw with caninoid
teeth on each side of the symphysis. The second
dorsal fin is elongate with 20 rays. The bulbous
eyes contain a free lower eyelid and rows of short
barbules are found on each side under the lower
jaw. The colour of the dorsal surface in life 15 very
similar to the substrate on which they live, with
the ventral surface being white. When preserved,
the animal takes on a slate blue colour from which

673

the common name blue mud hopper (Grant,
1993) may have arisen.

S. histophorus are found in areas containing
substantial amounts of thixotropic mud (with à
depth to the compact layer ranging from approx-
imately 0.1-1m deep) and are usually associated
with mangrove areas which undergo cyclic tidal
emersion. Their distribution extends from the
mouth of the Brisbane River, inland to the suburb
of Fig Tree Pocket (Fig. 1). Visual searches did
not uncover any animals from Nudgee Beach 1o
as far north as Reef Point (Fig. 1). These arcas,
although containing apparently suitable habitats
associated with mangroves, did not support com-
munities of S. histophorus. The tidal flats associ-
ated with the suburbs of Lota supports a large
colony (c.21000) with potential distribution cx-
tending at least as far south as Waterloo Вау.

Temporal patierns - long term: Interviews were
conducted with two employees of the Brisbane
Public Transport service who have worked on the
Dutton Park - University of Queensland ferry
service for mány years. They reported that they
had not noticed the mudskippers until the summer
of 1992. They kindly interviewed their co-work-
ers, many of whom have worked on the Brisbane
River for more than ten years, and they too con-
firmed this observation.

The owner of а local aquanum shop captured
some of the animals when she first noted them,
again the date reported was 1992. Information
solicited from the Brisbane community via arti-
cles placed in local newspapers indicates that the
mudskippers may have been present as early as
the early 1970's in isolated pockets in the
Wynnum arca (M. Peart, pers. cornm.). The areas
of Doboy Creek, Dutton Park Ferry terminus,
Orleigh Park (J. Thomson, pers. comm.), Oxley
Creck (D. Miller, pers. comm.) and the mud flats
at Bulimba (К. Киеу, pers, comm.) were also
identified as mudhopper habitats.

Temporal patterns - seasonal: Preliminary in-
Formation on apparent seasonal change in abun-
dance was obtained during this study. Animals
were first observed by the authors followrng the
first sighting of 5. histophorux at Dutton Park
Ferry terminus in the summer of 1992-93 and
subsequently disappeared in the winter of 1993,
They reappeared in October 1993 (late Spring).
The number of animals noted during the present
study (c.60) reduced sharply in March 1994 (late
summer) until their disappearance in May 1994
(early autumn). The numbers of adult animals
substantially reduced in numbers from the Dutton
Park study site c. April 11-15.

674

KEY:

Adult mangrove 2опе.................. i, 1
Sapling mangrove 20Ппе.................. 2
Pneumatophore 2опе................... 3

High tide level
Low tide 1е\уе!.......................

MEMOIRS OF THE QUEENSLAND MUSEUM

Sree ry

FIG, 2. Zonation pattern present on the shore of the Dutton Park ferry terminal.

At the start of the distnbution observations
only adult animals were sighted. The first juve-
nile (c.50mm total length) was recorded from
Dutton Park study site on March 4. Juveniles
were found in a different zone (the pneumato-
phore zone) from the adults and were still present
at the study site after adult numbers had reduced
substantially. By mid August 1994 no mudskip-
pers of any age group were found to be inhabiting
the site. All evidence of territories such as holes
or shallow pools were no longer present.

Zonation: Animals at the Dutton Park ferry
terminus are distributed in a distinct band across
the shore. The shore at the Dutton Park Ferry
terminus is divisible into four distinct zones (Fig.
2). The first zone consists of adult mangroves
(Avicennia sp.) which grew close to the high
water mark. The second zone adjacent to the first
is the mangrove sapling zone which contains
many fiddler crabs (Uca spp.). The third zone was
the mangrove pneumatophore zone which con-
tains small unidentified macropthalmid crabs. Ju-
venile mudskippers were also found in this area
during the months of April-and May. Adult mud-

skippers are found only in the fourth zone from
the edge of the pneumatophores (c. mid-tide
level) to the low tide mark. Scartelaos only be-
comes active once the tide level had fallen below
the pneumatophore zone. The "Scartelaos zone"
(i.e. below the pneumatophore zone) was also
apparent at the Lota study site. The Lota site also
includes a fifth zone below the mudskipper col-
ony to the low tide mark in which no animals are
observed and thixotropic mud and seagrass dom-
inates. This fifth zone was not observed at the
Dutton Park site.

Patterns occurred in horizontal distribution as
the animals were not uniformly distributed within
the Scartelaos zone at the Dutton Park site. The
area closest to the jetty contains several mating
pairs with all solitary animals being excluded
from the area. The area with many lone animals
contained mostly displaying animals with very
few pairs apparent and many agonistic interac-
tions (Townsend, unpublished data).

REAPPEARANCE OF BLUE MUDHOPPER

DISCUSSION

This is the first detailed study of mudskippers
in the Brisbane region, It reveals that either his-
torical records and some detailed studies are less
than accurate or that mudskipper range varies
remarkably. Periophthalmus vulgaris, which had
been previously reported in the greater Brisbane
area (Castelnau, 1878) was not found. Interviews
with ferry staff and members of the Brisbane
community indicates that S. histephorus has only
recently reappeared in substantial numbers in the
greater Brisbane area. Seasonal activity was re-
corded with a complete absence of the animals
from the study site during the winter months of
August through to mid October. The only other
report of seasonality ts for Boleopiihalmus
pectinirostris from Japan (Fukuda, 1994), The
greatest activity of the animals occurring during
the summer months from April to November and
only moving to catch food during warm days of
winter (Fukuda, 1994). The extreme seasonality
of S. histophorus may be unique to the Brisbane
area as this complete "disappearance" has not
previously been reported.

It is hypothesised that the animals undergo
either aestivation or migration away from the
sites during the Winter months, Berti et 21. (1992)
demonstrated the "homing" abilities of a Peri-
ophthalmus sp. from a delta in the Tana River
(Kenya) and, although this ability was only tested
al a maximum distance of 250m away from the
territories, similar abilities would be necessary if
migration does occur in S. histephorus. Alterna-
tively, mudskippers may avoid extremes in tem-
peratures by retreating into deep water-filled
burrows (Tytler & Vaughan, 1983) therefore the
animals may be aeslivating during the winter
months. Further study, via tag and recapture or
burrow excavation, needs to be undertaken lo test
these hypotheses.

Mudskippers may prove to be convenient bio-
indicators of estuarine health, Further research to
this end may investigate the relationship between
the benthic diatom community and the density
and abundance of mudskippers. For example,
subtle modifications of the benthic diatom com-
munity, due to anthropogenic effects, may lead to
changes jn the abundance and distribution of 5.
histophorus. In Japan, reduction of estuarine
health due to polluted rivers has been pinpointed
as one of the causes of reduced numbers of
Boleophthalmus pectinirosrris (Fukuda, 1994).

675

FUTURE RESEARCH

In addition, future research may be aimed at
answering the following questions, Have the $.
histophorus communities "returned" to the
greater Brisbane area or were they isolated to
pockets of fragmented habitat? Can the popula-
tion change be attributed to pollution levels, sub-
stratum changes, increase in mean temperatures
or other physical factors? Research should be
aimed al identifying if the factors resulting in the
changes of distribution of S. histophorus are due
toenvironmental quality such as diatom numbers,
sediment changes, chemical changes or due to
anthropogenic effecis or physical changes such
as temperature and climate.

Unique to S. histophorusinthe greater Brisbane
area, 1$ the disappearance during the winter
months. Research into the activities of the ani-
mals during the cold winter periods may be inves-
tigated. Are they aestivating or do they under go
migration? Winter searches and tag and recapture
methods may be employed to answer this.

Little is known about the factors resulting in the
age structure, setllement and recruitment pra-
cesses of S. Aistophorus. The vulnerability of the
population needs to be identified. This paper
highlights the need for further study to fully un-
derstand the role of S. histophorusin the Brisbane
estuarine ecosystem.

ACKNOWLEDGEMENTS

We thank the following people for their help:
Kevin Townsend, Melanie Venz, Peter Kind, all
the kind responses from the Brisbane public, lan
Johnstone and Leigh Henderson from Brisbane
City Transport, Coral McGregor from the
Clayfield aquariums, computer help from Narelle
Hall and Petter Fugelli and for the kind advice
from Dr. Norman Milward.

LITERATURE CITED

BERTI, R, CHELAZZI, L, COLOMBINI, 1. &
ERCOLINI, A.. 1992. Direction-findine ability in
a mudskipper from the delta of the Tana river
(Kenya). Tropical Zoology 5(2); 219-228,

CASTELNAU, F., 1878 Australian Fishes: New or little
known species. Proceedings of the Linnean Soci-
ety of NSW Vol IT: 225-248.

FUKUDA, T., 1994, Endangered species in Japan:
Mutsugoro (Mudskipper) Boleophthatmus
pectinirostris, Nature Productions, Japan 24: 12.

GRANT, E.M., 1993. Grants Guide to Fishes. Wilke
printers, QueensiIand:597, 602.

676

MARSHALL, T.C., 1966. Pp. 368-373. Fishes of the
Great Barrier Reef and coastal waters of Queens-
land. (Angus & Robertson Ltd: Sydney).

MCCULLOCH, A.R. & OGILBY, J.D. 1919. Some
Australian fishes from the family Gobiidae. Re-
cords of the Australian Museum 12: 193-291.

MILWARD, N.E., 1974. Studies on the taxonomy,
ecology and physiology of Queensland mudskip-
pers. PhD Thesis, Department of Zoology, Uni-
versity of Queensland, Brisbane.

STEAD, D.G. 1906. Pp. 186-187. Fishes of Australia:
A popular and systematic guide to the study of the

MEMOIRS OF THE QUEENSLAND MUSEUM

wealth within our waters. (William Brooks and
Co.: Sydney).

STEBBINS, R.C. & KALK, M. 1961. Observations on
the natural history of the mudskipper, Peri-
ophthalmus sobrinus. Copeia, 1: 18-27.

TYTLER, P. & VAUGHAN, T. 1983. Thermal ecology
of the mudskippers, Periophthalmus koelreuteri
(Pallas) and Boleophthalmus boddarti (Pallas) of
Kuwait Bay. Journal of Fish Biology 23(3): 327-
337.

WEBER, M. & DEBEAUFORT, L.F. 1953. The Fishes
of the Indo-Australian Archipelago. Volume X
(E.J. Brill, Leiden: Netherlands).

DEVONIAN THELODONT SCALES (AGNATHA, THELODONTI) FROM
QUEENSLAND

SUSAN TURNER

Turner, S. 1995 12 01: Devonian thelodont scales (Agnatha, Thelodonti) from Queensland.
Memoirs of the Queensland Museum 38(2). 677-685. Brisbane. ISSN 0079-8835.

Turiniid thelodont scales from the Emsian-Eifelian Cravens Peak Beds ofthe Toomba Range,
Toko Syncline, western Queensland and the undifferentiated Broken River up (late
Emsian serotinus Conodont Zone to Givetian) of the Broken River Province of north
Queensland are referred to Turinia gavinyoungi sp. nov. and Jesslepis johnsoni gen. et sp.
nov. respectively. [ ] Devonian, Thelodontt, Turiniidae, Queensland.

Susan Turner, Queensland Museum, P.O. Box 3300, South Brisbane, Queensland 4101,

Australia; 12 October 1995.

Early to Middle Devonian limestones from the
Broken River Province, near Greenvale, north
Queensland have yielded microvertebrate assem-
blages which include rare thelodont scales (Jell et
al., 1983; Turner, 1991b, 1993), Turinia sp. cf. T.
australiensis has been reported from the Martin's
Well Limestone of Magpie Creek, Shield Creek
Group at Broken River (Turner, 1993), and one
possible turiniid scale has been found within the
Givetian part of the Broken River Group. Within
the Burges Formation (late Emsian-Eifelian) of
the Broken River Group a series of thelodont
scales have been found at various localities in the
Burges and Wando Vale districts. In the Toko
Syncline, western Queensland, a limestone mem-
ber of the Cravens Peak Beds has also yielded
abundant thelodont scales from shot point local-
ities (Turner et al., 1981; Young, 1984, 1995).
The scales from both locations are sufficiently
dissimilar to those already known to warrant as-
signment to new taxa. They are referred to the
Turiniidae which comprises 3 genera; Turinia,
Australolepis and Boreania (Turner, 1991a).
Turiniid thelodonts are now known from all con-
linents in the Lower Devonian and are most
longlasting in Gondwana, becoming extinct only
in the early Frasnian.

Broken River samples were collected by J.S.
Jell (The University of Queensland UQ Locality
numbers, UQL) or during sampling for conodonts
by John Talent and Ruth Mawson (Mawson et al.,
1988; Mawson & Talent, 1989; Macquarie Uni-
versity, Centre for Elostratigraphy & Palaeobio-
logy samples from SAG section); the type
specimens are designated with Queensland Mu-
seum numbers (QMF). For recent summaries of
Broken River geology see Withnall & Lang
(1993) and Sloan el al. (1995), The Toko Syncline
limestones were collected by John Draper and

samples specifically for fossil fish were obtained
by Gavin C. Young (type specimens designated
in the Commonwealth Palaeontological collec-
tion, CPC, based at the Australian Geological
Survey Organisation, Canberra).

SYSTEMATIC PALAEONTOLOGY

Superclass AGNATHA
Class DIPLORHINA
Subclass THELODONTI
Order THELODONTIDA
Family TURINIIDAE Obruchev, 1964

Jesslepis gen. nov.

TYPE SPECIES
J. johnsoni sp. nov

DIAGNOSIS

Small to medium-sized scales; large open oval
to slit-like pulp cavity. Crown rises steadily from
anterior to sharp posterior point at an angle of
30-45'. Sharply angled posterior point. Crown
narrow, base low, neck virtually absent. Two to
four deeply dissected crown ribs meet the crown-
base interface at a steep angle. Crown ribs can
extend from the anterior rim to the posterior point
in a straight line.

REMARKS

Differs from Turinia, Australepis and Borania
(eg., Turner, 1991а) in the general morphology of
the crown, having arelatively smooth and steeply
sloping upper surface in most scale varieties:
elongate triangular or arrowhead-like outline;
and the low number of crown ribs.

678 MEMOIRS OF THE QUEENSLAND MUSEUM

DEVONIAN THELODONT SCALES

Jesslepis johnsoni sp, nov.
(Figs 1, 2A-M)

1991b turintid, Turner: 447.
1993 endemic thelodants, Turner: 183,

DIAGNOSIS
As for genus.

MATERIAL EXAMINED

HOLOTYPE: QMF33724,

PARATYPES: QMF33721-3, 33725-40. 20 scales from
UQL4703 youngest; UQLA704 (type locality);
LIQL4706 oldest; UQL4697; UQLA388; UQLA374;
014054, basal Lomandra Limestone, late Emsian
and early Eifelian Broken River Group. (stratigraphic-
ally below Fish Hill Limestone Member, sce Withnall
& Lang 1993); SAG 2 (3 scales); SAG B (2 scales), SE
of Arch Gorge section (Mawson & Talent, 1989; table
5) Chinaman Creek Limestone of Sloan et. al. (1995).

DESCRIPTION

Scales mostly 0.5-1mm long. Oral or head
scales (Fig. 2A) are rounded and cap-like in on-
togenetically young scales as in most thelodont
Benera but can have deeply dissected crown ribs
which can curve (Figs 1B,C, 2A,B).
Cephalopectoral scales are more elongated and
can have prongs midway on the lateral ribs (Fig.
1A,L). Lateral ribs can be curved (Fig. 1A, B,D,
Н) or straight (Fig. 1F-G,M). Postpectoral or
tnink scales are narrow and arrow-shaped and
generally longer than wide (Figs 1F, 2G,J). V-
shaped scales (Fig. 1M) might possibly be from
the pinnal or caudal regions. Bases are mostly as
large as the scale and about one-third the height
(Figs 1D,G-H, 2F,LL) but can be shallow in on-
togenetically young scales (Fig. 11), Basal cren-
ulations and lobules are common (Figs
1A,E,K-L, 2K.M) often mirronng the shape of
the crown, In lateral view the posterior base
seems d and forms a skirt-like extension
(Figs 1D,G, 2C,LL). The basal cavity ean be à
shallow central concavity (Fig. 2E) becoming
reduced as the base grows to a small central pulp
opening (Fig. 2H).

Histology, Internal examination by thin section
or by anise oil immersion has not been possible
because the blackened specimens are few in num-
ber. However, nearly all scales examined are

479

cap-like with a large wide-open pulp cavity but
older scales with mature bases are seen (see above
and Fig, 2),

AGE

The Burges Formation of Withnall & Lang
(1993) has been referred to "undifferentiated Bro-
ken River Group" by Sloan et al. (1995). Upper
and lower limits of the taxon are not well con-
strained but on the evidence of associated corals
(Jell, pers. comm.) and conodonts from SAG
section the possible range seems to be between
serotínus and basal ensensis Conodont Zones.

Thelodont scales are rare in these marine lime-
stones and for now, because of the potential range
of variation within turiniid squamations, all but
one of the scales found are placed in J. johnsoni.
De Pomeroy (1994) recorded new shark genera
from the Broken River sequence, mainly from the
late Eifelian-Givetian interval (kockelianus-var-
cus Zones), associated with acanthodian, thelod-
ont, placoderm, and osteichthyan remains, She
has reported one thelodont scale from the Eifelian
SAG section (De Pomeroy, pers. comm.) which
has been referred to an indeterminate thelodont
(De Pomeroy, 1995, in press, fig. 6M, N). This
scale is referred to the new genus and most
closely resembles the scale illustrated in Fig. 3B.
The new scales descrihed here come from a series
of limestone samples, notably from the measured
section "SAG" which has conodont control
(Mawson et aL, 1988; Mawson & Talent, 1989).
There are now three scales from SAG 2, and two
from SAGS. These scales occur 26,8 and 67m
above the base of the sections respectively (Maw-
son & Talent, pers. comm.) SAG2 conedonts
include P. I linguiformis and P. parawebbi,
which co-occur in the australis Zone (Mawson &
Talent, 1989; fig. 9), Conodont elements, 7.
siruvet and Р. L klapperi appear at 55.6m above
the base of SAG section (Mawson & Talent 1989,
Table 5). SAG8 would, therefore, appear to be
close to the base of ensensis Zone. The new scales
are associated with onychodnnt teeth and unde-
termined acanthodian scales, Cheiracanihoides
ef. comptus (C. comptas s.s. is typical of latest
Emsian-Eifelian; Turner, 1993), cosmine scales,
shark scales (De Pomeroy, 1994) and possible
shark teeth.

FIG. 1. Jesslepis johnsoni gen. et sp. nov. from UQL4704, Broken River Group, Broken River Province,
Queensland, Australia. A, QMF33721, lateral view. B, crown view of A, C, QMF33722. lateral view. D,
QMF33723, lateral view. E, crown view of D. Е, Holotype QMF33724, crown view. С, lateral view at F. Н,
QMF33725, lateral view, J, crown view of H, К, QMFE33726, crown view. L. antero-lateral view of K. M,
QMF33727, crown view. №, lateral view of M. Bar = 0 5mm.

680 MEMOIRS OF THE QUEENSLAND MUSEUM

DEVONIAN THELODONT SCALES

REMARKS

In common with other Australian turiniid scales
(e.g. Turinia australiensis, Australolepis
teddont; Turner, 1991b), the new taxon shares
some general characters such as the high crown,
large ulp cavity and cap-like nature of most
scales. However, micro-omament is absent; this
latter character appears in many of the
Gondwanan species of Turinia and appears to
have phylogenetic sinificance (Turner, submit-
ted).

ETYMOLOGY

In honour of Dr J.G. ("Jess") Johnson (1932-
1994), Devonian biostratigrapher and palacontol-
ogist, and lepis; Greek; scale.

Turinia gavinyoungi sp. nov.
(Figs 2N?, 3)

I981 Turiniz cf, pages, Turner el al: 54, figs, 6-8.
IOF-L, HH.
1991h Turinia n. sp., Turner: 447, pl. 5D, Gg. 4B, E-G

DIAGNOSIS

Head and trunk scales large with high crowns.
In trunk scales anterior median crown section
narrow. Crown deeply dissected with ribs and
lobes leading to a high central ndge culminating
in sharp posterior point. Typically three pairs of
lateral ribs, Ribs can be split and upturned. Mul-
tiplication of riblets and lappets on crown. Micro-
ornament sometimes present. Concave neck.
Base usually wider than crown and expanded
anteriorly. Basal nodules common.

MATERIAL EXAMINED

HoLoTYPE: CPC 20079/3 in Turner ct al., 1981, fig.
7E, atypical trunk scale from a small limestone outcrop
in the southern part of Toomba Range, western flank
of Toko Syncline, Georgina Basin, western Queens-
land, central Australia, Approximate latitude and lon-
gitude 23" 23° 47"S, 138" 08" 10"Е (map and section
in Turner et al., 1981, fig. 2; Turner & Young, 1987,
fig. 1). Cravens Peak Beds: upper Emsian? - lower
Eifelian, top Lower? - basal Middle Devonian (see
discusssion in Young 1995).

PARATYPES: CPC20079/1-2, 4-20, 20080/1-4 from
sample GEO 65/28 (Turner et al. 1981); GB77.(loc. 11)
С.С. Young coll. V1649, QM F33742-97 from same

681

locality as the type specimens. QMF33741 from
UQL4374 Burges Formation of Broken River Province
might belong to this taxon.

DESCRIPTION

See descriptions and measurements given in
Turner et al. (1981: 55); average length 1.5mm.
Material from sample GB77 is figured here for
the first time. Head/oral (Fig. 3A-C),
cephalopectoral/transitional (Fig. 3D-L, Y?) and
trunk scales (Fig. 3M-X) are found. Turner et al.
(1981; figs 6-8) showed some of the range of
scale variation. Crowns are typified by the out-
wardly expanding triangular lappets which end in
many posterior points (Fig. 3M-N, R-W) The
crown can be divided into five or more compart-
ments by, typically, three pairs of deeply dis-
sected ribs and lappets (Fig. 3J-L,Y). The
posterior median part of the crown is sharply
pointed and can be recessed below the crown
ridge (Fig. 3R,W). Lateral ribs can expand into
an upward tuming projection at mid length or
near the crown-neck interface (Fig. 3L,N,P).
Multiple nbbing (Fig. 3M) and micro-omament
is seen especially on trunk scales (Turner ct al.,
1981; figs TE-G, 8D, 107), On the bases very long
anterior processes are typical (Turner et al., 1981;
fig. SB; Fig. 3R-T) and commonly can be double
but with one process longer than the other (Fig.
3U-V); this latter might represent an individual
variation. Multiple basal lobules (Fig. 3B) can
occur (Turner ef al., 1981; fig. ЗА).

Histology. Preparation in anise oil shows a typ-
ical turiniid dentine crown with thin durodentine
cap surmounting a large base of aspidine pene-
trated by the canals of Sharpey's fibres (Turner,
19913). Large pulp cavities are common (Fig.
3A), Although the dentine tubules seem relatively
sparse, they are straight with few branches except
al the proximal ends; no clearly specific attributes
can be seen except where the dentine tubules are
longer to reach into the expanded lappets and rib
extensions,

REMARKS

Turner (їп Turner et al., 1981) compared this
form with the type species, Turinia pagel Powrie
in its well-developed elongate high crown with a
slight anterior notch. The scales differ in the more

FIG, 2, Broken River Group thelodont scales, A-M, Jesslepis johnsoni . А. QMP33728 head scale from UQL 4054,
basal view. B, QMF33729 from UQL4704, crown view. C-E, QMF33720 from UQL4704. C, lateral view. D,
crown view. E, basal view, Р-Н, QMF33731 from UQLA704. F, lateral view, G, crown view. Н, basal view. 1-7,
QMF33732 from UQL4388, I, lateral view, J, crown view, K-M, QMF33733 from UQL4388, К, crown view,
L, lateral view. M, basal view. №, Turinia gavinyoungi?, QMF33741, broken scale from UQLA374. Bar = 0 5mm.

682 MEMOIRS OF THE QUEENSLAND MUSEUM

DEVONIAN THELODONT SCALES

extreme development of lappets and partitions of
the crown, seen particularly well in the trunk
scales. Closely spaced double ribs are compara-
ble. with those of Givetian Turinia hutkensis
Blieck & Goujet (1978) of Iran, whereas the
out-turned ribs resemble those of the Eifelian to
Givetian T. pagoda and 7. spp. A & B of West
Yunnan, China (Wang et al., 1986), The presence
of micro-ornament suggests close relationships
with the earlier Т. australiensis group of scales,
as well as to Givetian taxa, T. pagoda and T.
antarctica Turner & Young (1992), The scales in
the Cravens Peak Beds sample are robust and
although the crowns can be well scalloped they
do not exhibit the fragility and lightness seen in
scales of the early Frasnian turinid, Aus-
tralolepis. Onc possible scale from the marine
limestones of the Broken River Group with pos-
terior extensions and lappets (Fig. 2N) might be
à scale of Turinia gavinyoungi.

Thelodont scales outnumber acanthodian
scales in the assemblage by approximately 4:1.
The associated fauna includes climatiid spines,
Acanthodes type and Machaeracanthus scales;
scales and bones of an antiarch placoderm
(Young, 1984); shark teeth Mcmurdodus, (Turner
& Young, 1987) and various shark scales-
Onychodont teeth are common as well as scales
of a sarcopterygian. Lepidotrichia are also pres-
ent. The samples appear to be relatively imal-
tered, the colour ranging from orange to mse
quartz and, given the good preservation and lack
of abrasion, the scales are unlikely to have been
transported far,

ETYMOLOGY

For Dr Gavin C. Young, Devonian fossil fish
worker, who collected much of the Cravens Peak
material.

AGE

Based on the associated vertebrate assemblage,
the present consensus favours an Eifelian age for
the in situ limestones of the Cravens Peak Beds
(Young, 1995), In the absence of conodonts the

683

evidence is provided by fish and crustaceans. P.J.
Jones (in Turner etal., 1981) noted that the ostra-
cods and the eridostracan from shot point sam-
ples and from GEO 65/28 were most like those of
early Devonian age from the northern hemi-
sphere, The thelodont in the shot point samples is
T. cf. australiensis and thus could be as old as mid
Lochkovian-Pragian ог more likely early Emsian
(Turner, submitted). T. gavinyoungi occurs in
samples GEO 65/28 and GB77. Comparing this
taxon with others known from Australia, from the
nearby Ross River (Young et al, 1987) and
Hatchery Creek Formation (Young & Gorter,
1981) where there are scales comparable with
Turinia sp, of Khush-Yeilagh and T. hutkensis of
Iran (now thought to be of early Eifelian and late
Givetian age respectively (Lelièvre et al, 1993),
then an Eifelian age for T. gavinyoungi seems
most probable,

SIGNIFICANCE

Work in the last decade itas clarified the range
and distribution of thelodont scales jn Australian
Devonian sequences (Turner, 1995; Young,
1995), Their usefulness as zonal or age indicators
has been acknowledged especially in the absence
of conodonts. The taxonomy of Australian the-
lodonts 3s better understood now that compari-
sons can be made from a series of different facies
from. many horizons throughout the Lower to
early Upper Devonian and from several coun-
tries. With the aid of recent studies of condont-
dated sequences, the ranges of thelodont species
have been refined (Turner, 1995; submitted;
Young, 1995),

The new scales from north Queensland are
unlike any previously decribed in the Devonian.
In their simpilicity they resemble earlier Silurian
thelodont scales but the ubiquitous light-weight
of the base with a large pulp cavity is a feature in
common with the younger Australolepis of the
early Frasnian of Western Australia, Further stud-
ies from conodont-dated sections should help 10
refine the range of Jesslepis.

FIG. 3. Turinia gavinyoungi sp. nov, from the Cravens Peak Beds of the Georgina Basin, Queensland, Australia.
A-C, I? Head/oral scales. D-H, J-L, Y? cephalopectoral/transitional scales. M-X, trunk scales. A, QMF33742,
intemal sagittal section of broken scale. B, QMF33743, lateral view, C, QMF33744, lateral view, D, QMF33745,
lateral view. E, antero-lateral view of D. F, QMF33746, crown view. С, lateral view of F. H, QMF33747, crown
view, I, QMF33748, crown view of oral or special scale. J, QMF33749, crown view. К, QMF33750, crown
view. L, QMF33751, lateral view. M, QMF33752, crown view, N, QMF33753, crown view, О, QMF33754,
crown view. P, QMF33755, lateral view, Q, crown view of P, R, QMF33756, basal view. S, QMP33757,
Jatero-basal view. Т, basal view of S, U, QMF33758, lateral view. V, anterior view of U. W. QMF33759, anterior
view. X, lateral view of W. Y, QMFE33760, crown view. Bar = 0.5mm.

684

T. gavinyoungi is comparahle with Early
Devonian scales such as Turinia pagel, and T,
uustraliensis but is more like Middle Devonian
turiniids from elsewhere in Gondwana. As the
Cravens Peak Beds contains no conodonts, as-
sessment of the associated fauna has suggested
the age of T. gavinyoungi as Eifelian (Young,
1995). However, the possibility that one scale
from the conodoat-bearing Broken River Group
belongs to this taxon suggests that the age and
range will be clarified by future sampling of
measured sections,

ACKNOWLEDGEMENTS

1 thank Dr John S. Jell (The University of
Queensland) and Dr Gavin C. Young (A.G.S.O.,
Canberra) for allowing and enabling me to study
máterial collected by them. Professors John Tal-
ent and Ruth Mawson (M.U.C.E,P., Sydney)
gave freely of information regarding conodont
dating of Broken River samples; Alison De
Pomeroy allowed me to see unpublished data on
her thelodont scale. An anonyinous reviewer
helped to clarify some points in the manuscript.
Work on Australian microvertebrates has been
supported bv grants from the Australian Research
Committee and is currently supported by an Aus-
tralian Research Fellowship. | thank the late Gud-
run Sarne (Queensland Museum), Dora Aitken
(ARC funded) and Diane Phipps (The University
of Queensland) for taking the SEM photomicro-
graphs and Broce Cowell (Queensland Museum)
for providing photographic support. This is a
contribution to IGCP 328: Palaeozoic
Microvertebrates.

LITERATURE CITED

BLIECK, А. & GOUJET, D. 1978. A propos de non-
veau maleriel de Thélodontes (Verlébrés
Agnathes) d'Iran ct de Thailande: aperçu sur la
répartition géographique еї stratigraphique des
Agnathes des "regions gondwaniennes" au
Paléazoique moyen, Annals de la Société
péologique du Nord, Lille, 97. 363-72.

DE POMEROY ‚ А. 1994, Mid-Devonian shark scales
from the Broken River, north Queensland, Aus-
tralia. Memoirs of the Queensland Museum 37, 1:
37-114.

1995. Australian Devonian fish biostratigraphy in rela-
tion lo conodonl zonation, AUSCOS-I Sympo-
sium volume. Courier Forschungsinstitut
Senckenberg 182: 475-486.

IN PRESS, Biostratigraphy of Early and Middle
Devonian mucravertehrates from Broken River,

MEMOIRS OF THE QUEENSLAND MUSEUM

north Queensland, Western Australian Museum

Records

JELL, J.S., TURNER. S. & YOUNG, С.С, 1983, New
Palaeozoic fish fannas from Queensland. "Evolu-
tion and Biogeography of early Vertebrates", spe-
cialist symposium 6th A.G.C. 16-25 Feb., Sydney
& Canberra, Abstracts ; 15-16,

LELIEVRE, H., JANVIER, P. & BLIECK, A. 1993.
The Siluro-Devonian vertebrates of Western
Gondwana and related terranes (South America,
Africa, Armorica-Bohemia, Middle East). pp 139-
173. In ЈА, Long (ed.) ‘Palaeozoic Vertebrate
Biostratigraphy and Biogeography’ (Belhaven
Press: London),

MAWSON, R. & TALENT, J.A. 1989. Late Emsian-
Givetian stratigraphy and conodont biofacies -
carbonate slope and offshore shoal to sheltered
lagoon and nearshore carbonate ramp -Broken
River, north Queensland, Australia, Courier
Forschungsinstitut Senckenberg 117: 205-259,

MAWSON, R., TALENT, J.A., BEAR, V.C., BEN-
SON, D.S., BROCK, G-A., PARRELL, J.R., HY-
LAND, K.A.. PYEMONT, B.D., SLOAN, T.R..
SORENTINO, L.. STEWART, M.1., TROTTER,
J.A.. WILSON, G.A. & SIMPSON, А.б. 1988,
Conodont data in relation to resolution of stage
and zonal boundaries for the Devonian of Aus-
tralia. Pp. 495-527, In N.J, McMillan, А Е, Embry
& D.J. Glass (eds) “Devonian of the World", val ,
(Canadian Society of Petroleum Geologists; Cal-

Bary),

OBRUCHEY, D.Y, 1964. Osnovy Paleontologii v. 11,
Byeschyeliostniy i Ribi. Nauka, Moscow [English
translation 1967. ‘Fundamentals of Paleontology,
VoL XI. Agnatha & Pisces’. (Israel program for
scientific translahion: Jerusalem).

SLOAN, Т.К, TALENT, ЈА., MAWSON, R., SIMP-
SON, AJ., BROCK, G.A, ENGELBRETSEN,
M., JELL. 1.5. AUNG, A.K.. PFAFFENRIT-
TER, C., TROTTER, J. & WITHNALL, LW.
1995. Conodont data from Silurian-Middle
Devonian carbonate fans, debris flows, al-
lochthonous blocks and adjacent autochthonous
platform margins: Broken River and Camel Creek
areas, north Queensland, Australia, Courier
Forschungsinstitut Senckenburg 182, 1-77,

TURNER, S. 1991a. Monophyly and interrelationships
of the Thelodonti, Pp, 87-119. In Chang, M-m, Liu
Y-h & Zhang, G.R. (eds) ‘Early Vertebrates and
related problems of Evolutionary Biology’. (Sci-
ence Press: Beijing).

1991b, Palaeozoic vertebrate microfossils in Austral-
asia. Pp. 429-464. In P. Vickers-Rich, J.N.
Monaghan, Е.Е. Baird & Т.Н. Rich (eds) 'Verte-
hrate Palaeontology in Australasia’, (Pioneer De-
sign Studios with Monash University Publications
Committee: Melbourne).

1993. Palacozoic microvertebrate biostratigraphy of
Eastern Gondwana. Рр.174-207. In J.A. Long
(ed.) ‘Palaeozoic Vertebrate Bioatratigrpiy and
Biogeography’. (Belhaven Press: London),

DEVONIAN THELODONT SCALES

1995. Defining Devonian microvertebrate faunal as-
semblages or zones for Eastern Gondwana. Pp.
43-52. In Turner, S. (ed.) Ichthyolith Issues Spe-
cial Publication 1, IGCP 328 (J.J. Zidek Serv:
Socorro: New Mexico).

(submitted). Devonian thelodont scales from East
Gondwana. In Klapper, G., Murphy, M.A. &
Talent, J.A. (eds) J. С. Johnson Memorial vol-
ume, Geological Society of America Memoir.

TURNER, S., JONES, P.J. & DRAPER, J.J., 1981,
Early Devonian thelodonts (Agnatha) from the
Toko Syncline, western Queensland and a review
of other Australian discoveries. Bureau of Mineral
Resources Journal of Australian Geology and
Geophysics 6: 51-69.

TURNER, S. & YOUNG, G.C., 1987. Shark teeth from
the Early- Middle Devonian Cravens Peak Beds.
Alcheringa 11: 233-244.

TURNER, S. & YOUNG, G.C., 1992, Thelodont scales
from the Middle-Late Devonian Aztec Siltstone,
southern Victoria Land, Antarctica, Antarctic Sci-
ence 4: 89-109.

WANG, S-T., DONG, Z-Z. & TURNER, S. 1986.
Discovery of Middle Devonian Turiniidae (The-
lodonti:Agnatha) from west Yunnan, China. Al-
cheringa 10; 315-325.

APPENDIX

685

WITHNALL, I.W. & LANG, S.C. (eds) 1993, Geology
of the Broken River Province, North Queensland.
Queensland Geology 4, 292pp.

YOUNG, G.C. 1984. An asterolepidoid antiarch
(placoderm fish) from the Early Devonian of the
Georgina Basin, central Australia. Alcheringa 8:
65-80,

1995. Timescales Calibration and Development. 4,
Devonian. Australian Phanerozoic Timescales.
Biostratigraphic charts and explanatory notes.
Second Series. AGSO Record 1995/33, 1-47 +
chart.

YOUNG, G.C. & GORTER J.D. 1981. A new fish fauna
of Middle Devonian age from the Taemas/Wee
Jasper region of New South Wales. Bureau of
Mineral Resources Bulletin 209, Palacontological
Papers 1981: 83-147.

YOUNG, G.C., TURNER, S., OWEN, M., NICOLL,
R.S., LAURIE, І.К. & GORTER, J.D. 1987. A
new Devonian fish fauna, and revision of post-Or-
dovician stratigraphy in the Ross River Syncline,
Amadeus Basin, central Australia, Bureau of Min-
eral Resources Journal of Australian Geology and
Geophysics 10: 23- 42.

Broken River province (approximate latitude and longitude 19° 144°40'Е) sample locality details;

UQL4703 (youngest), UQL4704, UQL4706 (oldest), south-folded limestones, traverse to road from Digger’s
Creek Crossing (BURGES 1:100,000 Sheet 7859: 683.489) stratigraphically below Fish Hill Limestone Member
(see Withnall & Lang 1993).

UQL4697 limestone about 70m W of old road crossing at Digger's Creek, stratigraphically below Fish Hill
Limestone Member, sec above,

UQLA734 thin limestone in shale, north bank of Broken River upstream of Gorge (645.457);

UQL 4388 thin crinoidal debris limestone, north bank of Broken River upstream frorn UQLA374 (644,458);

UQL4054 western creek bank of Dosey Creek, 750m upstream from its junction with the Broken River (615.438),
basal Lomandra Limestone, late Emsian;

MUCEP samples SAG 2; SAG 8: SAG = a measured section through Chinaman Creek Limestone Member
(Mawson & Talent, 1989, fig. 1, 4, table 5) commencing SE of Arch Gorge (at BURGES Sheet, 717.539)
approximately 4.2km E of Jessey Springs Hut,

686

FIRST RECORD OF PARASITISM B'Y A TICK ON AN
AUSTRALIAN FRESHWATER CROCODILE. Memoirs
of the Queensland Museum 38(2):686. 1995:- 1 report an
uncommon occurrence of the tick, Amblyomma sp., after
encountering а nymph om а yearling freshwater crocodile,
Crocodylus johnstoni, Although leeches and dermal nema-
todes are commion ecto ites on this crocodile (Webb et al

1983), thas 18 the first record Of tick parasitism on any Austra-
Gan crocodile.

The crocodile was captured during an ongoing study of C,
Johnsioni $n the Lynd River of north central Queensland
(Tucker et al., 1994). The crocodile (#3057) was marked as à
hatchling on 18 December 1993 nnd recaptured on 30 Novem-
ber 1994, approx. 3, Rken upstream From the onganal nest site
The recapture site was U. lkm upstream from an area known
focally as the "Croc Hole” (near 144" 20" W, 17°45'S), The
crocodile was initially resting on а sandy hank, entered the
Water and was Ihen caught within a shallow pool. И was
emaciated and3ts length (19.7cm snout-vent length) and mass
(120g) were Һем average ja companson Ie conspecifics af
the same age (mean Jengtb =23,.5cm, SD =3.2cm, л = 100;
mean mass = 250g, SD = L10g,»=92) From the crocodile's
peck, I removed a nymphal tick which was placed inio a vial
of ethanol. The tick was idemified at the Deparimem of
Parasitology, University of Queensland and the specie
deposited ia the Queensland Musuem (QMS26 190, U.Q. Par-
asrtolagy reference B398), Giver Ihe limitations of existing
taxunomic keys, it is difficult to positively identify any
nymphal tick beyond the genus level, However, ihe distribu-
tion records for tick specimens in the Queensland Museum
strongly suggest either Ат. limbatum or Ami. moreliae.

No ficks are known to be obligate ectoparasites of croco-
diles (J, Keirans, pers. comm.) and there are few published
records of ticks found on crocodiles. Neuman ( 1899) recorded
Ambiyamma ? grossum (Pallas) oo crocodiles in Saninam but
the ldentity of the ick ss questionable. A pomamrma exerum
have been recorded on a crocodile (probably C. aifoiicus) in
the Congo (Schwerz, 1927), on С, nilorzur i Uganda
(Manliysee & Colb, 1987) and іп Mala (Villiers, 1995).
These records likely resulted from a crocodile preying ороп а
Gck-infested unimal or by transferral of a tick trom another
repüllan host. The latter source 19 more probably grven the
high frequency of tick infestations recarded оп Vararus niot-
cus from the same regions in Africa.

Over 100 species of Amblyomma exist worldwide and the
life histories of well known species indicate they are three-
hos! ticks (Roberts, 1970). Six Australian ticks are common
ectoparasites of reptiles (Amd, аатинаа. Amb, полете,
Amb, limbatum, Amb calabayi, Am Jitriatm and Ap.
hydrosauri) (Roberts, 1970; Heatwole Æ Pianka, 1993) and а
complete host-parassus list appears in Roberts (1970), Two
newly described ies, Amb. plaueri aud Ap ребра,
were added recenily for varsmid lizards (Keirans ct al, 1994.
Although а few semi-aqualic hosts have heen recorded, in.
cluding two chelid turtles amd 4 water dragon, reptilian hosts
Of ticks are inantly from and terrestrial habitats,

Distribution records of repulian hosts for thie tick gemis
(Queensland Мивеипк. unpubl- data) include many large
snakes and lizards with ranges that overlap the study site,
particularly Eastern brown snakes {Pseudonaja rexrilis), car-
pel pylhons (Morelia spilota variegata) and sand goannas (М.
gouldii Cogger, 1992), although other umdocuinented hoses
cannot be excluded, Cioanna ects are common on the sandy
Пос plain at the study site and their truils cross the aver near
the capture location (A. Tucker, pers. obs}. Although previ-
ous herpetofaunal surveys of (his region indicated a decline in
their numbers (ОРЕН, unpabl. data), recent observations
confirm that goannas are at Jessi focally abundant near
Frenchy's Crossing {0.7km upstream of the recapture Joca»

MEMOIRS OF THE QUEENSLAND MUSEUM

tion), Il cannol be verified that the tick was (гапа еттей Irom
these or other potential hosts but the tick certainly attached
dumne a terrestrial emergence by the crocodile, as hard ticks
do not оссигоп wholly xc Vertebrates. Freshwaler croc-
odiles emerge mainly to hask, or nest, but sometimes move
overland during the dry season. Parasite transferal by inges-
поп of a tick-infested host is unlikely as the diet of yearling
C johnstonisncludes no vertebrates other than tiny Frogs. This
hast-parasite relationship зв undoubtedly rare since this repre-
sents ihe only occurrence noted in aver 3300 crocodiles
examined indrvidually for ectoparasites during the эш.

Although the evidence |х circumstantial, it appears likely
that a local goanna population serves as a preferred host for
the tick, Large lizards can easily serve as potential vectors
since tropical tends or varanids are commonly parasited by
ticks and offen found near habitats used by crocodilians, № is
reasonable to presume that tick-parasitism nf crocodiles prob-
ably resulted from imlirect rransferral from a reptilian host
rather than through direct ingestion of parasitised prey. In
either case, the occurrence of tick parasitism on crocodiles
remains highly infrequent,

Acknowledgements

I thank B. Scott for tick identification; T. Jessop апи Т,
Dempster for field assistance, J. Kesrans for references; C.
Limpus апан, McCallum for coqatuichve comments, snd H-
Janetski and M. Bull for records of reptile or tick distributions.

Literature Cited

Cogger, H.G. 1992. Reptiles and Amplubians of Australia
(Reed Press: Sydney),

Heaiwole, Н. & Pianka, В.К. 1993, Natural history of rhe
Squamata. Pp. 197-209. In Glasby, C.L, Ross, G.J.B. &
Beesley, P L. (eds.) Раша of Australia, vol.2a, Am-
phibia and Reptilia ( Australian Government Publishing
Services Canberra).

Keirans, J.E., King, DR. & Shamad R.D. 1994. Aponowema
(Bothriocroton) glebopalma, n. subgen. n. sp. and
Amblyomma plauerti n Sp. (Acari Ixodida’ Ixodidae),
Parasites of monitor fizards (Varanidac) in Australia,
Journal of Medical Entomology 31:132-147,

Matihysee, Ј.С. & Calbo, М.Н, 1987 The Ixodid Ticks of
Uganda (Entomological Society of America: College
Park, Maryland).

Neumann, L.G, 1899, Révision de Ia famile des Ixodedés.
Mémoires de la Société Zoologique 12- 107-297.
Roberts, FHS. 1970, Australian ticks (Commonwealth Sci-
АНЕ amd Inifustrial Research Organisation: Aus-

кайа}.

Schweiz, J, 1927. Coninbution à l'étude de [xedidae (liques)
n Conge belge. Revue Zoologique Africaine 15: 73-

Tucker, A-D., McCallum, H.E, Limpus, C.J. & McDonald,
K.R, 1594. Cracadyfur johnstoni in the Lynd River,
Queensland: cannnuanon of a long term fiel study, Рр.
469-473. In Proc. 2nd Regional Meeting of the Croco-
dile Specialist Group. JUICN, Gland Switzerland.

Villiers, A, 1955, Note sur quelques Ixodidae el Gramasidae
parasites de veriébrés rencontrés en Afmque Oc-
cidentale Francais Bulleting de Где francaise
d' Afnque Noire 17А: 444-454.

Webb, С JW. & Manolis, S.C. 1983, Crocodylus johnstoni in
the McKinlay River Area, Nothem Terntory, V. AB-
normales apd injuries. Australian Wildlife Research
Ик 407-420.

Antan D. Tucker, Department of Zomogy, University of
Шен Brisbane, Queensland 4072, Ansirahin; 24 April,

A NEW CTENOTUS (REPTILIA: SCINCIDAE) FROM THE MITCHELL GRASS
PLAINS OF CENTRAL QUEENSLAND

STEPHEN K. WILSON AND PATRICK J, COUPER

Stephen K. Wilson & Patrick J. Couper 1995 12 01: A new Crenorus (Reptilia: Scincidae),
C. agrestis from the Mitchell Grass plains of central Queensland. Memoirs of the Queensland
Museum 38(2):687-690. Brisbane. ISSN 0079-8835.

Ctenotus agrestis sp. nov., from the black soil, grassland plains of the Aramac area of central
Queensland, is a member of the C. lesueurii group. It bears some resemblance to C. robusins
and С, joanae. Colour, pattern and scalation readily distinguish it from these species.

L ICtenotus, C. lesueurii group, black-soil, central Queensland,

Stephen K, Wilson & Patrick J, Couper, Queensland Museign, PO Box 3300, South
Brisbane, Queensland 4101, Australia; 14 June 1995,

Ctenotus is the largest genus of Australian rep-
tiles. Ninety species are now recognised. Diver-
sity is greatest in arid and seasonally dry areas,
yet few Ctenotus inhabit the deeply cracking clay
(="black-soil") plains with Mitchell grass (As-
trebla) of Australia's eastern interior. During a
herpetological survey in central Queensland
(1987), three specimens of a new Ctenotus, con-
forming to the C. lesueurii species group of Storr
ct al.,1981 (toes only slightly compressed, lamel-
lae under toes with a wide callus; second supra-
ocular wider than third; pattern well developed
with bold dorsal stripes and pale lateral spots)
were collected from grasslands near Aramac.

While many reptiles (e.g., Ctenotus robustus)
include black-soil plains in their broad distribu-
tions, only a small number appear to be confined
to this distinct habitat. C. agrestis joins a short list
of species (Pogona henrylawsoni (Agamidae),
Ctenotus schevilli and C. joanae (Scincidac),
Varanus spenceri (Varanidae), Pseudechis col-
letti, Pseudonaja guttata and Р. ingrami
(Elapidae) regarded as black-soil endemies.

All measurements were taken using Mitutoyo
electronic callipers. Supraciliaries, supralabials,
infralabials and subdigital lamellae on the fourth
toe were counted on both sides of specimens
examined. Abbreviations for body measurements
are as follows:- snout-vent Jength (SVL); axilla
to groin (AG); tail length, vent to tip (TL); fore-
limb, axilla to tip of longest digit (L1); hindlimb,
groin to tip of longest digit (L2); forelimb to
snout, from anterior limb insertion to tip of snout
(LI-S); head width, widest point (HW); head
length, tip of snout to posterior margin of pari-
etals (HL); snout, tip to anterior margin of orbit
(Sy; eye to ear, posterior margin of orbit to dorsal
anterior margin of ear (EE). Additional material
examined in this study 15 listed in appendices 1 &

2. Other abbrevialions used: Queensland Mu-
seum (QM), Australian Museum (AM), Muscum
and Art Gallery of the Northern Territory (NTM),

SYSTEMATICS

Ctenotus agrestis sp. nov.
(Figs 1,2)

Ctenotus sp. (2). Wilson & Knowles, 1988, p.277.

MATERIAL EXAMINED

HOLOTYPE: QMJ46694, Brendallan Stn, via Aramac,
central Queensland (22°57'S, 145" 14' E), coll, S.K,
Wilson and P. J. Couper, 06 March 1987. PARATYPES:
0М246489, 0M146695, collection data as forholotype
except QMJ46689, coll. 05 March 1987.

DIAGNOSIS

Ctenotus agrestis can be confused only with C.
robustus and, to a lesser degree, with C. joanae.
It is readily distinguished from the former by its
pale colour (dorsally pale grey-brown vs brown -
olive brown); size (max SVL 73.9 vs 110.0mm);
size and shape of ear lobules (inconspicuous and
rounded versus prominent, and pointed or
rounded); single supradigital scale row on the
fourth toe (extending along entire digit vs distal
portion of digit only, fig.3).

From C. joanae it is distinguished by the num-
ber of scales along the mid-line between the
mental and anal scales (77-78 vs 63-68); and hy
the upper lateral pattern (elongate pale dashes
versus plain, or sometimes with a series of small
pale dots).

DESCRIPTION

SVL(mm) 66.5-73.9 (mcan-71.2, N=3). Pro-
portions, (%SVL):- AG=49,2-53.3 (mean-51.2,
№3); TL=155.4-173.2 (mean=164.3, №2);

688

MEMOIRS OF THE QUEENSLAND MUSEUM

FIG. 1. Ctenotus agrestis sp. nov., (Holotype QMJ46694) in life (Photograph by S. Wilson).

L1=24.8-29,2 (теап=26.9, N23); L2=37.6-41.5
(mean=39.9, N23); L1-S=33.1-33.4 (mean-33.2,
N=3); HW=12.9-13.0 (mean=12.96, N=3);
HL=16.5-18.1 (mean=17.6, N23); S=7.3-8.3
(mean=7.9, N=3); EE=6.8-7.8 (mean=7.3, N=3).

Nasals in point to broad contact; nasal groove
absent; prefrontals very narrowly to moderately
separated; maximum length of frontal 1.7-2.1
times maximum width (mean=1.9, N=3); frontal
contacting frontonasal, prefrontals, first three
supraoculars and frontoparietals; supraoculars 4,
second the largest; supraciliaries 7-8 (mean=7.5,
N26), first or second the largest; frontoparietals
paired and distinct from interparietal; enlarged
nuchals 7-9 (mean=8, N=3), two-three in direct
contact with parietals; loreals two; presuboculars
1; preoculars 2, the lower being the largest; supra-
labials 7-8 (mean=7.5, N=6) with fifth or sixth
subocular; infralabials 6-8 (mean=7, N=6);
postmental contacting two infralabials on each
side; ear opening large, vertically elliptic with 3
-4 (mean 3.2, N 6) small lobules on anterior edge.

Midbody scale rows 30-32 (mean-30.7, №3);
number of scales in a direct line between mental
and anal shields 77-78 (mean=77.7, N23);
paravertebral scales, from anterior-most nuchal
to posterior margin of hindlimb 60-61
(теап=60.7, N23); lamellae beneath fourth toe

FIG. 2. Ctenotus agrestis sp. nov., Holotype
QMJ46694. Lateral and dorsal views of head.

NEW CTENOTUS FROM CENTRAL QUEENSLAND

FIG.3. Arrangement of supradigital scales on the fourth
toe. (Left) Crenotus agrestis sp. nov., Holotype
QMJ46694, (Right) Ctenotus robustus, AMR62272.

17-19 (mean=18, N=6) broadly callose; a single
row of supradigital scales present along almost
the entire length of the fourth toe.

The measurements and scale counts for the
holotype (QMJ46694) are as follows:
SVL=73.3mm; Аб=36.1пт; TL=114.0mm;
Ll-18.2mm; L2-27.5mm; L1-8224.29mm;
HWz9.5mm; HL-12.1mm; S-5.4mm;
EE=5.0mm. Maximum length of frontal 4.9mm;
maximum width of frontal 2.4mm; supraciliaries
8. first largest; enlarged nuchals 9, three in direct
contact with parietals; supralabials 8 (left side)
with sixth entering eye, 7 (right side) with fifth
entering суе; infralabials 7 (both sides); ear lob-
ules 4 (left side) 3 (right side); midbody scale
rows 30; number of scales in a direct line between

FIG. 4. Habitat of Crenotus agrestis, Brendallan Stn,
via Aramac, Central Queensland (22*57°$, 145" 14" E).

689

mental and anal shields 77; paravertebral scales
61; lamellae beneath fourth toe 17 (left side) 18
(right side).

Pattern. Holotype - Dorsal and upper lateral
ground colour pale-grey brown. Longitudinal
stripes (Fig. 1) are as follows: Vertebral black,
broad and prominent, from nuchals to base of tail;
paravertebrals diffuse and pale; dorsolateral
white, from above eye to tail tip, bordered dor-
sally by irregular black edge; upperlateral, a se-
ries of pale grey dashes, from eye to hindlimb;
midlateral white, from nostril to tail; lowerlateral
obscure grey-brown, from labials to tail; ventro-
lateral grey-brown, incomplete, between axilla
and hindlimb. Ventral surface white. Head with
obscure dark blotches. Limbs pale grey-brown
with pale stripes.

Variation in paratypes - Vertebral narrow and
obscure (QMJ46695). Irregular black upper edge
to dorsolateral, almost non-existent (QMJ46695).
Ventrolateral continuous (QMJ46689).

DISTRIBUTION
Known only from Brendallan Stn (22°57'S,
145°14’E), via Aramac, central Qld.

HABITAT

(Fig. 4) The type and only known locality is an
open black-soil plain vegetated with Mitchell
Grass (Asirebla sp.) and scattered low Acacias
(probably A. farnesiana).

ETYMOLOGY
Latin - agrestís relating to the fields, alluding
to the open, grassland habitat at the type locality.

ACKNOWLEDGEMENTS

We thank the Landers family for kindly allow-
ing us to work on Brendallan Station; Ross Sadl-
ier and Paul Homer for allowing us access їо
specimens in the collections of the Australian
Museum and Museum and Art Gallery of thc
Northern Territory respectively; Gordon Guymer
from the Queensland Herbarium for identifying
the flora in the habitat photograph; Lauren Keim
for her assistance in the laboratory; Greg
Czechura and Glen Ingram for their encourage-
ment and advice; Jeanette Covacevich for sug-
gesting improvements to the manuscript.

LITERATURE CITED

STORR, G.M., SMITH, L.A. & JOHNSTONE, R.E.
1981.'Lizards of Western Australia. 1. skinks.

690 MEMOIRS OF THE QUEENSLAND MUSEUM

(Univerity of Western Australia Press with West- WILSON, S.K. & KNOWLES, О.С, 1988. ‘Australia’s
em Australian Museum: Perth), reptiles’. (Collins Australia; Sydney).

APPENDIX 1. Specimens of Ctenotus robustus examined in the current study. All localities are for Queensland
unless otherwise indicated.

Queensland Museum. QMJ12110, Mt Мону (16'41'S, 145°20°E); QMJ1686, Herbert Gorge (18°14'S,
145°32’E); QMJ27697, Hencamp Ck, 5km N, Ikm E Rollingstone (19*02'S, 146° 19 E); QMJ27646-47, Rowes
Bay, 3km N, 3km W Townsville (19*14'S, ` 14647 E); QMJ44567, Lolworth Stn., Blossom Hill (20"09'S,
144°59'Е); QMJ44572, Lolworth Stn.. (20° 12 S, 14458" E); QMJ5637, Lindeman Is. (20'27'S, 149°02'E);
QMJ44586-87, Campaspe Stn (20°30°5, 145"38' E); QMJ44706, Mt Cooper Stn. (20°30'S, 146°51'Е);
QMJ44622-23, Helenslee Stn. (20731'S, 145°42'Е); QMJ44588-89, Pajingo Stn. (20'46'S, 1467 10'E);
QMJ44850, Hanging Rock Stn. (21'09'S, 14647" E); QMJ44562-63, Natal Downs, Curtis Dam (21°11'S,
146'04'E); QMJ28332, Kynuna, 9.6km SE (21'40'S, 141°59’E); 0М731526, 0М731530, near Mt Flinders
(22°33'5, 150°46'Е); QMJ46682-83, 0М746688, 0М746692, Brendallan Stn via Aramac (22°57°S, 145°14'Е);
QMJ47152, Nth Keppel Is (23°10°S, 150"58'E); QMJ32586, ‘Lochnagar’, via Barcaldine (23°34’S, 145°39'Е);
QMJ46760, Kalapa (23'31'S, 15016 E); QMJ24218, Curtis Is, S епа (23°45'5, 151°18'Е); 0М727911,
Lowmead, Warro (24°32'S, 151°45'Е); QMJ47093, Dawson R. Crossing at Baroondah Stn (25'41'S, 149" 13'E);
QMJ12105, 17.6km from Goomeri (26'02'S, 152°02'Е); QMJ31520, Sandy Ck, via Ferndale (26'45'S,
151703'E); QMJ28625, Moreton Is., NE end (27^ 11'S, 153'24' E); QMJ22966, Virginia, Brisbane (27"23'5,
15309" E); QMJ40741, Brisbane (2728 S, 15301 E); QMJ6747, Toowoomba (27°34'S, 151°57'Е); QMJ16087,
Forest Hill (27°35°S, 152'21'E); QMJ26384, Dynevor Lakes, 44,5km E, 7km N Thargomindah (28'04'S,
144°10°E); QMJ35407, QMJ35427, Inglewood, old dump site (28'25' S, 151"05'E); QMJ35426, Brush Ck Stn,
21km S Inglewood (28'36'S, 151°06'E); QMJ40359, Ballandean, via Stanthorpe (28'48'S, 151750'E);
QMJ47095, Girraween NP via Stanthorpe (28°50°S, 151'55'E); QMJ12113, Wyberba, via Stanthorpe (28°52’S,
151°52'Е); QMJ30720, Texas Caves, via Texas (28'53'S, 15126' E); 1439, S Queensland; QMJ31860-01,
Condobolin, NSW (33*05'S, 147°09"E).

Australian Museum. AMR62275, 143km S Hughenden (22°00°S, 144°28'Е); AMR64334-35, 80.1km N
Muttaburra via Hughenden rd. (22"02'S, 149"29'E); AMR61500-13, 150km from Hughenden on Muttaburra rd.
(22713'S,144"16' E); AMR62271-72, 38km S of Muttaburra on Агатас rd. (2246 S, 14453'E); AMR62274,
64km S Muttaburra on Aramac rd. (22°51'S, 145*04' E); AMR62273, Aramac rubbish tip (22/58' S, 145" 14^ E).

APPENDIX 2. Specimens of Crenotus joanae examined in the current study. All localities are for the Northern
Territory unless otherwise indicated.

Queensland Museum. QMJ54383-84, vicinity of Widdallion Ck; NWQLD (18'26'S, 13829" E).

Australian Museum. AMR71363, Avon Downs (20°02’S, 137°30°E); AMR80360-61, AMR80531, 20km W
of QLD/NT border on Barkly Hwy, (19°58'5, 137°49°B),

Museum and Art Gallery of the Northern Territory. NTMR3636, Anthony Lagoon (19°59’S, 135'36'E);
МТМЕ5326, Anthony Lagoon (17^59'S, 135°32'E); NTMR8447, No. 6 bore, Rockhampton Downs (19°23'S,
135°24'Е); NTMR9573, No. 17 bore, Alroy Downs (19°06'5, 136°12°E); NTMR14628, Rocklands Stn.,
Barwidgee Ck (19*49'S, 137°55'Е); NTMR 16424, Brunette Downs, Racecourse (18"36'S, 136'06' E).

FIRST ENDEMIC AUSTRALIAN OECOBIIDAE AND NESTICIDAE (ARACHNIDA:
ARANEAE)

JÓRG WUNDERLICH

Wunderlich, J. 1995 12 01. First endemic Australian Oecobiidae and Nesticidae (Arachnida:
Araneae). Memoirs of the Queensland Museum 38(2): 691-692. Brisbane, ISSN 0079-8835.

Oecobius inopinatus sp.nov. and Nesticellachillagoensis sp.nov. are described from Queens-
land, and represent the first endemic Australian species of the Oecobiidae and Nesticidae
respectively. Both taxa are only known from females. [ ] Araneae, Nesticidae, Oecobiidae,

taxonomy, Australia.

J. Wunderlich, Hindenburgstr. 94, D-75334 Siraubenhardt, Germany; 1 September 1995,

Until now, no endemic species of Oecobiidae
and Nesticidae have been described from Aus-
tralia. It has not been discussed whether members
of these families never reached Australia to
evolve native species or whether such species are
extinct. These records show that there are indeed
endemics of these families in Australia: on the
bark of trees (Oecobiidae) and in caves
(Nesticidae). Such places are not well studied, so
these - and perhaps some more related and unre-
lated - species have been overlooked.

Family OECOBIIDAE

AUSTRALIAN SPECIES
Oecobius navus Blackwall, 1859, (= annulipes auct.)
cosmopolitan; Oecobius inopinatus sp, nov.

Oecobius inopinatus sp. nov.

(Fig. 1)

MATERIAL EXAMINED

HOLOTYPE: QMS26040, 9, near Rockhampton, E
Queensland, no exact location, on bark of trees. Coll.
J. Wunderlich. July, 1992.

PARATYPES: QMS26041, 29, 1 juv. same data.

DIAGNOSIS

Female prosoma medially largely black (Fig.
1A), epigyne/vulva (Fig. 1B,C) with small circu-
lar hole frontally which bears genital openings,
medially with strongly concave sclerotized struc-
ture, posteriorly with fissure; with large and thin-
walled receptacula seminis. Male unknown.

ETYMOLOGY
Inopinatus meaning unexpected.

DESCRIPTION
Measurements (mm): body 1.8-2.0 long, pro-
soma: 0.6 long, 0.7 wide, leg I: femur 0.52, patella

Fic, 1, A-C, Oecobius inopinatus sp. nov.: A, prosoma
dorsally. B, epigyne. C, vulva dorsally (receptacula
not shown). D, Oecobius navus Blackwall, 1859,
epigyne. Scale = 0.1mm.

0.2, tibia 0.4, metatarsus 0.38, tarsus 0.35, tibiae
II-IV 0.4.

Colour of sternum yellow, prosoma dorsally
yellow, marginally and largely medially black
(Fig. 1A), chelicerae frontally with black spot,
legs yellow with distinct black annulations, op-
isthosoma yellow to grey, dorsally with white and
dark grey spots, laterally with dark spots, venter
uniformly yellow, spinnerets yellow, posterior
ones dorsally darkened.

Prosoma (Fig. 1A) wider than long, with
large eyes, posterior median eyes reniform.
Cribellum and calamistrum of normal size.
Legs with some long bristles (most bristles
broken off), longest under tarsus IV. Op-

692

isthosoma ovally, dorsoventrally depressed, cov-
ered with short hairs.

RELATIONSHIPS
This species differs from other known species
in the unique conformation of the 9 genitalia.

ECOLOGY & DISTRIBUTION
The spiders were found on the bark of trees near
Rockhampton, Queensland.

Family NESTICIDAE
Nesticella chillagoensis sp. nov.
(Fig. 2)

MATERIAL EXAMINED

HoLorYPE; QMS20910, 9, deep inside Royal Arch
Cave, e.g. entrance of St Bernard, humid places under
stress, near Chillagoe NEQ. Coll. D. Flett & J.
Wunderlich, July 1992.

DIAGNOSIS

Female colour pale, legs with long hairs, eyes
of medium size (posterior margin of posterior
median eye lenses indistinct, Fig. 2A), epigyne
(Fig. 2B) slightly sclerotized, ducts and
receptacula seminis clearly visible; vulva (Fig.
2C). Male unknown.

DESCRIPTION

Measurements (mm): body 2.8 long, prosoma:
1.3 long, 1.15 wide, leg I: femur 2.0, patella 0.6,
tibia 1.95, metatarsus 1.95, tarsus 0.8, tibia II 1.5,
tibia Ш 1.0, tibia IV 1.7, length of some hairs: on
tibia I 0.35, on the opisthosoma dorsally 0.33.

Colour pale, prosoma and legs orange, op-
isthosoma medium grey. Prosoma wide, with
large and low grove.

Eyes (Fig. 2A) of only medium size, posterior
margin of posterior median eye lenses indistinct,
posterior row straight, posterior median eyes sep-
arated by slightly more than their diameter.
Chelicerae large, anterior margin with 3 large
teeth, posterior margin with some tiny teeth and
with tiny teeth in the furrow. Labium fused to
stemum, wider than long, claw of pedipalpus
long, slightly bent, with numerous long teeth. -
Legs of medium length, covered with long hairs,
tibiae with two hair-shaped bristles(?). All meta-
tarsi with a trichobothrium, its position on I in

MEMOIRS OF THE QUEENSLAND MUSEUM

н

Fic. 2, A-C, Nesticella chillagoensis sp. nov. A, eyes
dorsally. B, epigyne. C, vulva dorsally. Scale=0.2mm.

0.43. Opisthosoma ovally, dorsally covered with
long hairs, colulus basally with a couple of hairs.

RELATIONSHIPS

I do not know a closely related species. This
species is distinguished by the unique conforma-
tion of the female genitalia. Only few Nesticella
species from New Guinea have been described,
compare e.g. Lehtinen & Saaristo (1980). A new
Nesticella species from north Queensland was
hinted in Davies (1986: 47, fig. 76) (д). In this
epigeic (?) species, the posterior median eyes are
not reduced. It is a pity that we do not know its
male.

ECOLOGY & DISTRIBUTION

Considering the cave locality in north Queens-
land, Australia, the pale body colour and the
slightly reduced posterior median eyes, the spi-
ders seem to be trogloditic.

ACKNOWLEDGEMENTS

Тат grateful to the Deutsche Forschungs-
gemeinschaft (DFG) who supported this work.

LITERATURE CITED

DAVIES, V.T. 1986. Australian Spiders. Queensland
Museum booklet No. 14. 60pp.

LEHTINEN, P.T & SAARISTO, M.I. 1980. Spiders of
the Oriental-Australian region. II. Nesticidae. An-
nales Zoologici Fennici 17: 47-66.

CONTENTS (continued)
MACKNESS, B.S, & VAN TETS, С.Е,

Status of the late Гюрт» fossil darter Anhinga laticeps (De Vis 190бу......,....,,.. +з... 611
PATERSON, R.A & VAN DYCK,

A blue whale Esos musculus (Linnaeus, 1758) from St Lawrence, Queensland .,..,..,..‚.. 615
RAVEN, RJ. & SCHWENDINGER,P.J.

Three new mygalomorph spider genera from Thailand and China (Araneae) ...................... 623
ROBINS, R.

The results of test excavations in three rockshelters in southwest Queensland ..................... 643

SHARMAN, M., WILLIAMSON, I. & RAMSEY, D.S.L.
Observations on the early life history stages of Notaden bennettii in the Chinchilla area of southern
Queenpsfátid i, iiie i ао Ашин» eani] ERE CRI Pr rh md ку bmi gla Pali „667
TOWNSEND, К.А. & TIBBETTS, LR.
Re-appearance of the blue mudhopper, Scartelaos histopharus (Pisces: Gobiidae) in the greater
Brisbane ай. eiusd Ыы caa potit eda eH eee hele e ER Hte peus 671
TURNER, S.

Devonian thelodont scales (Agnatha, Thelodonti) from Онеепзїала............................. 677
WILSON, S.K. & COUPER, P.J.
A new Ctenotus (Reptilia: Scincidae), C. agrestis from the Mitchell Grass plains of centra)

Оневая аяй, 22.225 20а аа Аа рдЕ, 5522.600. af edi eL Es ien, 687
WUNDERLICH, J.
First endemic Australian Oecobiidae and Nesticidae (Arachmda: Araneae) ....................... 691
NOTES
COUPER, P J.
Communal nesting in the small tud Lampropholis adonis , ,....... ss. еа 382
COUPER, P.J,, COVACEVICH, J.A., WILSON, S.K.
Sap feeding by the Austraban gecko Gehyra dabi. ы... етв а р-р ОБ SY 396
KUTT, A.S
Е А Mictocpiecpsimn bat mortality in a harp trap due to green tree ant Oecophylla smaragdina attack . .. . . 437
отмо significant vertebrate fauna records from mid-altitude wet tropical rainforest, Lamb Kange
lale Forasti li soin dine ete nhi tae a eread tatto rod us ез er iba te 440

KUTT, A. & SKULL, S.
Degraded riparian habitat at Charters Towers and its poteritial to decrease local diversiry and/ or

cause extinctions ..... Fu Mele fe hide te shies tte eh de pleads ИРИРЕК 446
LAWLESS, Р.В.
Brush-footed trapdoor (Ozicrypia sp.) burrows and geckos at Wando Vale, МЕО .................. 470
LOOP, К.А.
Observations of the amethyst python (Morelia amethystina) feeding on rainbow bee-eaters
(Merops ornatus) cessisse sham he] tham ahh eden ree 504
MITCHELL, D., JONES , A. & HERO, J-M.

Predation on the cane toad oe ds s a) by the black kite (Milvus migrans). i2... o 512
SEYMOUR, J., VOLSCHENK, E. & SCOTT, B.
Record of the scorpion Liocheles karschii (Keyserling, 1855) from north east Queensland

(Scorpionida: Ischnuridae). .. 5)... 2 ee eee eee TETE tec eee en Se TS LOR Tee 532
SHEA, G.M.
RTT The holotype and additional records of Pogena henrylawsoni Wells & Wellington, 1985,..,........ 574
A, G.M.
Ойун dubia (Macleay, 1877) confirmed as senior synonym of Perochirus mestoni de VIs, 1890. .. . 610
SHORT, J
SLM be record of Palaemon concinnus Dana, 1852 (Crustacea: Decapoda: Palaemonidae) from Australia , 622
UYS, R.
An alien flatworm in Australian waters... 2.00.20) cee nee) aha d naa 642
STEVENS, J
First record of the longfin mako (Isurus paucus) from Australian waters ...... "greet oa 2 670
TUCKER, A.D.

First record of parasitism by a tick on an Australian freshwater стосоййс.........................686

CONTENTS
BAEHR, M.

Revision of Philipis (Coleoptera: Carabidae: Bembidiinae), a genus of arboreal tachyine beetles
from the rainforests of eastern Australia. Taxonomy, phylogeny and biogeography........... 315
BURROW, CJ. & SIMPSON, AJ.
A new ischnacanthid acanthodian from the Late Silurian (Ludlow, ploeckensis Zone)
Jack Formation, north Queensland as. Кы ы seg ees et Eia kr kr ee roa € est Dele 383
CANNON, L.R.G. & SEWELL, K.B.
Craspedellinae Baer, 1931 (Platyhelminthes: Temnocephalida) ectosymbionts from the branchial

chamber of Australian crayfish (Crustacea: Рагаз!ас1Чае)................................ 397

CLIFFORD.H.T.

A permineralized cupulate fructification from Ошеепв1апа................................+з+-+ 419
COOK, A.G.

Gastropods from the Ukalunda Beds and Douglas Creek, Early Devonian, north Queensland ........ 429
COOK, A. С.

А new Silurian megalodont bivalve genus from Bungonia, New South У/айез..................... 437
DAVID, B., WATCHMAN, A., GOODALL, R. & CLAYTON, E.

ТЕЗУ OPUS BOUNCES os ole voir a Fas Hceris wr Polk He phenol pee pace Pe gee de ang paces ys 441
DAVIE, P.J.F. & TURNER, P.A.

New records of Cryptopodia (Crustacea: Decapoda: Parthenopidae) from Australia ............... 447
DAVIES, V.T.

A new spider genus (Araneae: Amaurobioidea: Amphinectidae) from the wet tropics of Australia. .. . . 463
FRITH, C.B. & FRITH, D.W.
Hybridization between the Great and Spotted Bowerbird Chlamydera nuchalis and C. maculata:
an authenticated hybrid bowerbird (Ptilonorhynchidae) ........................ prever 471
KELLY-BORGES, M. & VACELET, J.
A revision of Diacarnus Burton and Negombata de Laubenfels (Demospongiae: Latrunculiidae)
with descriptions of new species from the west central Pacific and the Red Sea ,............. 477
GILL, P.C., EYRE, EJ., GARRIGUE, C. & DAWBIN, W.H.
Observations of humpback whales (Megaptera novaeangliae) on a cruise to New Caledonia and the

RRSP ROK Te РРО pulsi Казиле. Geko decr dr SAO LET URS 505
GYNTHER, I.C. & O'REILLY, P.S.
A new locality for the Hastings River Mouse, Pseudomys oralis, in southeast Queensland. .......... 513
HANCOCK, D.L.
The butterfly types of W.H. Miskin in the Queensland Museum (1 ерідоріега).................... 519
HANNAH D.S. & SMITH, G.C.
Effects of prescribed burning on herptiles in southeastem Опеепв1апй........................... 529
HAYWARD,P J. & RYLAND, J.S.
Bryozoa from Heron Island, Great Barrier Reef. 2........................................... 533
JAMIESON, B.G.M.
New species and a new genus of earthworms in the collections of the Queensland Museum
(Megascolecidae: Oligochaeta): 22. аена ehh Hm arre кка е наа 575

LAURANCE, S. G. W. & LAURANCE, W. F.
A ground-trapping survey for small mammals in continuous forest and two isolated tropical

ао oae Ли, dein bid eese Bagh de hein als ie Oden EIE Fa Tore Ae Rar. muera 597
MACKNESS, B.S.
Palorchestes selestiae, a new species of palorchestid marsupial from the early Pliocene Bluff Downs
Local Fauna, northeastern Queensland... .......... 0. ccc ele esc een аана 603

(continued inside cover)