Memoirs
of THE
Queensland Museum
Brisbane
31 July, 1975
Volume 17
Part 2
Volume 17
Part 2
Memoirs
OF THE
Queensland Museum
Published by Order of the Board
Mem. Qd Mus. 17(2): 195-235, pis. 7-26. [1975]
THE GENUS MACROPUS SHAW (MARSUP1ALIA: MACROPODIDAE) IN THE
UPPER CAINOZOIC DEPOSITS OF QUEENSLAND
Alan Bartholomai
Queensland Museunn
ABSTRACT
The genus Macropus Shaw, 1 790, is shown to comprise three subgenera, M. ( Macropus) Shaw,
M. (Osphranter) Gould, 1842, and M. ( Prionotemnus) Stirton, 1955, on the basis of
morphological and palaeontological evidence. M. ( Macropus) is known only from Pleistocene
sediments, whereas the other subgenera are also well represented in the late Pliocene Chinchilla
Sand. Only two species of M.( Macropus) are recorded, the most commonly encountered being M.
titan, while M. rama is described as a new species. Within M. (Osphranter), four species are
recognized, M. altus and M.ferragus from the Pleistocene deposits and M. pan and M. woodsi sp.
nov. from the Chinchilla Sand. M. ( Prionotemnus) possibly comprises six species, four of which,
M. agilis siva , M. gouldi, M. piltonensis sp. nov., and M. thor, are restricted to Pleistocene
sediments. M. dryas and M. palankarinnicus are present in the Chinchilla Sand. Palaeontological
evidence suggests that M. ( Macropus) was derived late in the geological history of the group and
that within the other subgenera, general and occasionally particular relationships can be
suggested for derivation of the recent fauna. Possible phylogenetic relationships within the M.
(Osphranter) group are suggested back to the late Pliocene. In many species, statistical evaluation
of most of the cheek teeth was possible, and comparisons with data from recent macropodids have
been made.
Representatives of the genus Macropus Shaw are
very abundant in the Upper Cainozoic sediments of
Queensland. Although some specimens referable to
the genus came from widespread localities in
Queensland, most have been derived from the
Pleistocene fluviatile deposits and the Chinchilla
Sand of late Pliocene age, both in the Darling
Downs area, southeastern Queensland.
Considerable diversity of opinion has existed
regarding the generic limits of the genus Macropus
and the taxonomy and temporal relationships of
referred species. Bartholomai (1967, 1973a*,
1973b) has examined progressively aspects of the
overall problem, and the present study clarifies the
bulk of the remaining problems. Some of the results
of the present study were foreshadowed in Bartho-
lomai, 1972, while preliminary work on part of the
older type material was included in a study by
Bartholomai (1966). The availability of large
samples now enables reassessment of the species to
be made, especially aspects of intraspecific vari-
ation, utilizing evaluation of the populations by
statistical and comparative morphological means.
Results contribute to a better overall understand-
ing of the taxonomy of the genus Macropus, and
suggest that species are of potential value in
correlation of Upper Cainozoic continental de-
posits.
Less detailed investigation has been made of the
continental distribution of fossil species of Mac-
ropus, and until revisionary work has been com-
pleted on collections in other Australian museums,
conclusions in this area are largely tentative. Exact
temporal relationships are frequently difficult to
establish away from the type areas.
All measurements are in millimetres.
Genus Macropus Shaw, 1790
Type Species: Macropus giganteus Shaw, 1790
(validated under the plenary powers of the In-
ternational Commission on Zoological Nomencla-
* Erratum: In Bartholomai (1973a) the illustrations of fossils comprising Plates 21 and 23, but not the captions to these
plates, were inadvertently transposed during printing. Thus the caption to Plate 21 refers to the illustrations in Plate 23
and vice versa.
196
MEMOIRS OF THE QUEENSLAND MUSEUM
ture, Opinion 760, 1966, by monotypy, as in-
terpreted by the neotype, Queensland Museum
specimen, J10749, designated by Calaby et al.,
1962, and restated by Calaby and Ride, 1964).
The first large, modern macropodids were col-
lected in 1 770, some sixty years before the first fossil
representatives of the family were discovered. A
party from Captain James Cook’s ship, the ‘En-
deavour’ secured three specimens from the vicinity
of the Endeavour River, near the present position
of Cooktown, and from these Muller (1776)
described Macropus canguru. The apparent holo-
type was destroyed during the Second World
War through bombing of the Royal College of
Surgeons. The identity of the species, however, was
in doubt, and Iredale and Troughton (1925) had
previously suggested that the specimen described
may have been a wallaroo rather than a kangaroo.
Further complication was added when these auth-
ors (Iredale and Troughton, 1937) suggested, that
the specimen may have been a Whiptail Wallaby
( Macropus parryi). Both suggestions were based on
unpublished work of Solander (1768-71). Raven
(1939) contended that the first description was of a
Grey Kangaroo, and this was strongly supported
by Morrison-Scott and Sawyer (1950). In an
attempt to stabilize the nomenclature, Calaby et al
(1962) also reaffirmed this position and selected a
neotype, Queensland Museum specimen J 10749.
This specimen is a juvenile Grey Kangaroo col-
lected from the Cooktown area. Ride (1963)
commented further on the nomenclatural problem,
and this was followed by a revised application by
Calaby et al. (1963) which resulted in considerable
comment. Kirkpatrick (1963) and Woods and
Kirkpatrick (1964) indicated that the original
specimen was a Wallaroo. Mayr (1964) and
Lemche (1964) also commented on details of the
case, numbered Z.N.(S.) 1584. New proposals were
then outlined by Calaby and Ride (1964), and these
were supported by Morrison-Scott (1964) and
Finlayson (1964). Another complication was the
designation by Troughton and McMichael (1964a,
b) of an additional neotype, this being a Whiptail
Wallaby. Voting on the case favoured the pro-
position by Calaby and Ride (1964) and involved
the use of the plenary powers, stabilizing the
nomenclature under Opinion 760, as indicated
above.
Considerable diversity of opinion still remains
regarding the generic limits to be applied to the
genus Macropus, both in terms of its neontological
and palaeontological usage. Ride (1962) concluded
that ‘no confusion would result from the perfectly
valid action of any author who writes about a
species of wallaby or kangaroo (living or extinct)
and prefers to remain non-committal about his
generic concepts. Such an author may simply
employ Macropus ... or he may follow a stated
taxonomic list’.
The problem is not a simple one and the species
which may be referred to Macropus have been
relegated to the following genera (or subgenera)
depending on the limits placed by individual
taxonomists — Macropus, Halmaturus (a junior se-
condary homonym of Macropus but used for the
purpose of separation), Osphranter, Fissuridon,
Megaleia, Wallabia, Thylogale , Petrogale , Per-
adorcas, Lagorchestes, Onychogalea, Lagostrophus,
Dorcopsis, Dorcopsulus, Dorcopsoides, Den-
drolagus, Setonix, Synaptodon , Prionotemnus, Pro-
temnodon and Troposodon.
Ride (1962) has presented an historical summary
of various usages suggested by major contributors
to taxonomic interpretation of the group. Genera
from the above list which are apparently nomencla-
turally stable in recent literature include Fissuridon,
Thylogale, Petrogale, Peradorcas, Lagorchestes,
Onychogalea, Lagostrophus, Dorcopsis, Dorcop-
sulus, Dorcopsoides, Dendrolagus, Setonix and
Troposodon.
Bartholomai (1973a) defined the generic limits
of Protemnodon , indicating its distinction from
other macropodids. Earlier work by Stirton (1963)
was supported in the contention that Protemnodon
is distinct from living kangaroos and wallabies,
including the Swamp Wallaby, making available
the generic name, Wallabia, for at least part of this
group.
Progression of the cheek tooth row was con-
sidered by De Vis (1895) to be of extreme
importance in separation of the Queensland fossil
sample into the genera Halmaturus and Macropus.
The process of progression is particularly evident in
kangaroos, as shown by Kirkpatrick (1963, 1964)
for M. giganteus, and later (Kirkpatrick, 1965)
restated for this species, and shown for Megaleia
rufa and Macropus robustus, where the extent of the
progression has been employed in ageing of
specimens. Information on molar progression in
M. rufa has also been presented in Calaby (1968).
Although rate of progression is less pronounced in
brush wallabies it is nevertheless significant, as
indicated for M. rufogrisea by Kirkpatrick (1965),
and would appear to represent differences in
function in the two groups rather than generic
distinction. The degree of movement in Wallabia
bicolor has not been investigated to the same extent
but appears to be generally comparable with that in
the brush wallabies.
Cytological investigation of recent macro-
podines by Sharman (1961) has shown the availab-
BARTHOLOMAI : MACROPUS IN UPPER CAINOZOIC
197
ility of characters previously disregarded and this
has been followed by Kirsch (1968), who has
examined marsupial haemoglobin and has pre-
sented preliminary information for many living
macropodines. According to details provided by
Sharman (1961), chromosome number and sex
chromosome morphology indicate the generic
distinctness of Megaleia, Setonix and Lagos-
trophus. In addition, middle-sized wallabies, kan-
garoos and wallaroos are diphyletic with the type
species of Wallabia, W. bicolor , differing markedly
from the rest of this group. Serological studies by
Kirsch (1968) suggest that species of wallabies and
kangaroos, including W. bicolor, Megaleia and
Lagorchestes are closely associated, a conclusion
which cannot be verified by the fossil record
because of general deficiencies in fossil samples yet
available. Sharman et al. (1966) have recently
investigated reproductive physiology of W. bicolor,
showing it to be unique among the Macropodidae.
Further, Calaby (1966) states that this species
differs in behaviour and has distinctive dental
characters' from the rest of the group. Calaby
considers that Wallabia should be recognized as a
monotypic genus, while the remainder of the
wallabies, the Grey Kangaroo and the wallaroos
should remain within Macropus. This action is
supported by the present study.
All members of Macropus are characterized by
16 (2N) chromosomes. While differences are evid-
ent between species in this grouping, they are
nowhere as marked as those enabling separation of
Wallabia, which has a diploid complement of 10 in
the female and 1 1 in the male. Ride (1970) applies
this terminology, but Frith and Calaby (1969)
again revert to use of Macropus for the Grey
Kangaroo and wallaroos and Wallabia for all the
brush wallabies.
Equal area grid diagrams, as proposed by
Thompson (1959), have been shown by Bartho-
lomai (1973a) to be of value in illustrating gross
relative size and displacement differences present
between skulls of species of Macropus, Wallabia
and Protemnodon, based on a skull of W. bicolor as
a standard. Apart from differences between the
genera noted in Bartholomai (1973a), the figures
indicate a remarkable uniformity in modification
of the grid to M. giganteus and the brush wallabies
which possess the diploid chromosome number of
16.
Separation of Macropus is further supported by
the anatomy of the cheek teeth, particularly lower
molars. All species referred to Macropus have high-
crowned lower molars with strong links and with
near vertical, lingual lophid margins and non-
vertical labial margins. If ornamented, the strongly
curved, posterior hypolophid surface is, with rare
exception, grooved or pocketed. Protemnodon has
more rectilinear and generally relatively lower
lophids, and has the posterior surface of the
hypolophid considerably less curved. Ornamen-
tation of this surface is restricted to development of
a posterior cingulum. The permanent premolar is
not lost through progression. Wallabia has low
crowned molars with rectilinear lophids and low
links. Lateral margins of the lophids are bulbous,
while the posterior hypolophid surface is not
ornamented. These characters are of paramount
importance in consideration of the fossil material
and support the generic distinction of these forms
at least. The position of Synaptodon, described by
De Vis (1889), cannot be resolved at this time,
because it is based on inadequate and completely
undiagnostic material. The holotype of its type
species, S. aevorum De Vis, F811, from the late
Pliocene Chinchilla Sand at Chinchilla, was stated
by De Vis (1895) to be distinguished by peculiar
anterior and posterior abutting processes of the
molars. Examination has shown that crowns of the
molars are almost totally devoid of enamel except
for the ‘processes’ and a small patch on the trigonid
basin of the posterior molar. This suggests that the
‘processes’ may have resulted from weathering,
particularly as remaining enamel is soft and chalky,
whereas the dentine is hard and mineralized.
Certainly no other specimen in the Queensland
Museum collections duplicates this condition and
no adequate reason apart from abnormal weather-
ing can be suggested for the state of preservation of
the holotype.
Megaleia is not represented by fossil material in
Queensland. Tedford (1967) records Megaleia in
the Lake Menindee deposits in western New South
Wales but applies the name in the subgeneric sense
within Macropus. Megaleia is distinct from Mac-
ropus on the basis of modern species but this
distinctness becomes difficult to apply to fossils
because of general similarity in cranial morphology
between species of both genera. The last molar in
the cheek teeth series in Megaleia tends to be much
larger than that preceding it, a feature not common
in species of Macropus.
Although the description of the fossil genus
Prionotemnus Stirton, 1955, is largely inadequate,
an investigation of the referred sample in the
University of California, Berkeley, indicates that
the species is valid and that the name is available for
use within the Macropus complex.
For the purpose of this study, the genera
Macropus, Megaleia, Wallabia and Protemnodon
are recognized. Within the Macropus group, sev-
eral distinct species groups are apparent and these
198
MEMOIRS OF THE QUEENSLAND MUSEUM
are recognizable as far back as the late Pliocene.
Broadly speaking, these correspond to the kan-
garoos (excluding Megaleia), the wallaroos and the
brush wallabies (excluding Wallabia ). These
groups are regarded as subgenera.
Generic Diagnosis: Medium to large macro-
podines; cranium with rostrum markedly deflected
downwards; diastema relatively elongate; lower
molars with high lophids, strong links and with
lingual margin of lophids near vertical and labial
margins markedly divergent; posterior hypolophid
surface strongly curved, and where ornamented
this comprises a groove, fossette, or very rarely a
posterior cingulum.
Subgenus Macropus Shaw, 1790
Type Species: Macropus ( Macropus) giganteus
Shaw, 1 790 (validated under the plenary powers of
the International Commission on Zoological No-
menclature, Opinion 760, 1966 by monotypy).
Diagnosis: Medium to large macropodines
with palate entire; foramen ovale unhooded and
with alisphenoid only slightly grooved at margin of
foramen; upper incisors form V-shaped series in
occlusal view; I 3 long with labial surface marked by
two deep, vertical grooves; basioccipital slightly
keeled; postglenoid foramen well-developed; dias-
tema elongate; palate narrow anteriorly; per-
manent premolars reduced, rapidly lost during
progression; upper molars with strong forelink;
anterior ridge from paracone usually reduced or
absent; infraorbital distance between foramen and
anterior rim of orbit short; rostrum not greatly
inflated; lower molars with strong posterior
groove; mandible broad below anterior cheek
teeth.
Subgenus Osphranter Gould, 1842
Type Species: Macropus (Osphranter) anti-
lopinus Gould, 1842 by monotypy.
Diagnosis: Palate entire; foramen ovale hooded,
with alisphenoid deeply grooved at margin of
foramen; upper incisors form U-shaped series in
occlusal view; I 3 quadrangular with labial surface
marked by one vertical groove at anterior one-
third; basioccipital moderately keeled; postglenoid
foramen very reduced; diastema variable but
frequently short; palate broad anteriorly; per-
manent premolars relatively robust, but rapidly
lost during progression; upper molars with neglig-
ible to weak forelink; anterior ridge from paracone
usually reduced but occasionally stronger; in-
fraorbital distance between foramen and anterior
rim of orbit long; rostrum moderately to greatly
inflated; lower molars with strong posterior
groove; mandible excavated and narrow below
anterior cheek teeth.
Subgenus Prionotemnus Stirton, 1955
Type Species: Macropus ( Prionotemnus) palan-
karinnicus Stirton, 1955.
Diagnosis: Palate with extensive post-palatine
vacuities; foramen ovale hooded, with alisphenoid
deeply grooved at margin of foramen; upper
incisors form V-shaped series in occlusal view; I 3
triangular with vertical lateral groove in median
position or even in posterior moiety; basioccipital
markedly keeled; postglenoid foramen well-
developed; diastema moderately elongate; palate
anteriorly narrow; permanent premolars robust,
retained until very old age but occasionally lost
through progression; upper molars with minimal
forelink; anterior ridge from paracone strong;
infraorbital distance between foramen and anterior
rim of orbit long; rostrum not greatly inflated;
lower molars lacking posterior groove but some-
times with reduced posterior cingulum; mandible
broad below anterior cheek teeth.
Macropus (Macropus) titan Owen, 1838
(Plate 7, Figs. 1-2; Plate 8, figs. 1^1; Plate 9, figs.
1-4; Plate 10, figs. 1-3)
Macropus titan Owen, 1838, pp. 359-60, pi. 29, figs. 3-5;
1840-1845, 1, p. 392, pi. 101, figs. 1-2; 1845a, p. 236;
1845b, pp. 324-5; 1873, p. 128; 1874a, pp. 248-60, pi.
21, figs. 6-17, pi. 22, figs, 10-8, pi. 23, figs. 2-3, 12-4,
pi. 26, figs. 9-15; 1874b, pp. 783-4, pi. 76, figs. 1-6;
1876, pp. 204-9, pi. 25, figs. 1, 4, pi. 26, figs. 1-2;
1877, pp. 400-11, 435-9, pi. 76, figs. 1,'4, pi. 77,
figs. 1-2, pi. 78, figs. 1-2, pi. 79, figs. 1-2, pi. 8 1, figs.
6- 17, pi. 82, figs. 10-8, pi. 83, figs. 2-3, 12- 14, pi. 86,
figs. 9- i 5 ; Waterhouse, 1846, pp. 58-9; McCoy,
1862, p. 145; 1867, p. 191 ; Daintree, 1872, p. 274;
Etheridge Jun., 1878, pp. 183-4; 1892, p. 673;
Lydekker, 1887, pp. 225-30; Anderson, 1929, pp.
35-9, pi. 17, figs. 1-3, pi. 18, figs. 1-7; Simpson,
1930, p. 73.
Macropus magister De Vis, 1895, pp. 120-4, pi. 18, figs.
11-16; Bartholomai, 1966, pp. 123-4, pi. 19, figs. 1-3.
Macropus faunus De Vis, 1895, pp. 127-9, pi. 18, figs.
3-6; Simpson, 1930, p. 72; Bartholomai, 1966, pp.
122-3, pi. 18, figs. 1-3.
Material: F3738, cast of holotype, partial right
mandibular ramus with M : broken, M 2 , P 3 removed by
fenestration, juvenile, original in British Museum (Nat-
ural History), No. 10777, Wellington Caves, N.S.W.,
from Pleistocene cave deposits (figd Owen, 1838, pi. 29,
figs. 3-5; 1874a, pi. 22, figs. 17-8; 1877, pi. 82, figs. 17-18).
F2924, holotype Macropus faunus De Vis, partial right
maxilla with P 3 -M 3 , juvenile, Darling Downs (figd in
BARTHOLOMAI : MACROPUS IN UPPER CAINOZOIC
199
part, De Vis, 1895, pi. 18, figs. 4-5; figd Bartholomai,
1966, pi. 18, figs. 1-3).
F645, lectotype Macropus magister De Vis, partial
cranium containing P 2 -M 2 , P 3 removed by fenestration,
juvenile, Ravensthorpe, Pilton, SE.Q., (figd in part, De
Vis, 1895, pi. 18, figs. 13-14; figd Bartholomai, 1966, pi.
19, figs. 1-3).
Additional material referred to Macropus titan Owen
comprises 74 juvenile mandibular rami, 126 adult mandi-
bular rami, 4 isolated lower teeth, 1 1 cranial fragments, 25
juvenile maxillae and 84 adult maxillae from the follow-
ing localities in the eastern Darling Downs: King Creek;
King Creek, at M.R. 045455 Clifton 1-mile map; King
Creek, at M.R.037455 Clifton 1-mile map; King Creek,
near M.R.039454 Clifton 1-mile map; King Creek, near
M.R. 047452 Clifton 1-mile map; King Creek, Man-
apouri, at M.R. 099465 Liverpool Range 1-mile map;
King Creek, at M.R. 098465 Liverpool Range 1-mile map;
King Creek, at M.R. 048457 Clifton 1-mile map; King
Creek, between Pilton and Nobby; Ravensthorpe, Pilton;
Clifton; ?Pilton; Pilton; Spring Creek; Freestone Creek;
Westbrook Creek, near Kingsthorpe; Hirstglen; Gowrie;
Gowrie Creek; well at depth of c. 2 m, between Gowrie
Creek and radio station 4AK, Oakey; in sewerage drain at
c. 10 m, Dalby; near Dalby, Condamine River at Spring-
vale; Jimbour Creek near Dalby; Jimbour Creek, about
3 km south of Jimbour; bank of Condamine River, at
M.R. 043426 Dalby 1-mile map; Jimbour District; Cam-
booya, and from the eastern Darling Downs (particular
localities unspecified).
A juvenile mandibular ramus from Chinchilla is re-
ferred to M. titan , as is a juvenile mandible from the
Nogoa River, near Rawbelle, mid E.Q., a mandible from
Jimboomba, SE.Q., and an adult maxillary fragment
from Rubyvale, near Anakie, C.Q.
Specific Diagnosis: A large species. Diastema
elongate. P 2 relatively small with longitudinal crest
normally bifid and with strong, well-defined post-
erolingual cuspid, DP 3 and lower molars with high,
slightly curved lophids and strong, high links and
high anterior cingulum. Posterior surface of hy-
polophid with near vertical groove and well-defined
posterior fossette. P 3 small, usually with bifid
longitudinal crest, but occasionally trifid ; postero-
lingual cuspid present, close to posterior cuspid of
crest and united to this by high ridge. P 2 with well-
defined cusps and high bifid longitudinal crest;
protocone least well developed; cuspule present
labiad to metacone. DP 3 and upper molars with
high lophs and mid-links; forelink present, re-
latively well-defined. P 3 small, with longitudinal
crest normally bifid but occasionally trifid; hy-
pocone well-defined but low; protocone absent.
Description: Mandible moderately deep, re-
latively thick; base of symphysis deflected at lower
level than general base of ramus, near planar;
symphysis very elongate, shallow, not ankylosed,
rugose; geniohyal pit moderately deep, above
posterior symphysial limit; diastema very elongate,
with diastemal crest posteriorly acute, less acute
anteriorly; ventral margin of ramus rounded
between symphysis and extremely weak digastric
ridge and process. Mental foramen moderately
large, oval, usually set well below diastemal crest,
well anterior to anterior root of P 3 . Ramus with
relatively shallow lateral groove extending pos-
teriorly from just below posterior diastemal crest to
below anterior root M 2 , close to alveolar margin.
Digastric process separated from base of angle by
very shallow post-digastric sulcus, bounded above
by shallow digastric fossa; this fossa separated
above from broad depression opening posteriorly
into pterygoid fossa. Post-alveolar shelf short,
with angle not well-developed, leading to post-
alveolar ridge, ascending posteriorly to disappear
on mesial wall of coronoid process, above large
mandibular foramen. Masseteric crest raised to
about level of occlusion of cheek teeth; masseteric
foramen moderately large, with masseteric fossa
relatively deep. Angle of mandible markedly
inflected. Anterior margin of coronoid process near
vertical. Bulk of angle of mandible, coronoid
process and condyle not preserved in any specimen.
I L elongate, lanceolate, deeply rooted; slightly
curved in lateral view, markedly curved in occlusal
view, developing subhorizontal facet of wear with
upper incisors and mesial wear facet at tip by
approximation with other lower incisor; root
compressed, oval in section; crown subquadrantal
in section, tapering and blade-like anteriorly,
enamelled laterally, this produced dorsolabially
and ventrolingually into flanges; crown also en-
amelled ventromesially; distally, subhoF zontal
dorsal wear facet develops rapidly, but tip is not
rounded.
P 2 relatively small, short, approximately suboval
in occlusal view, with lingual surface slightly
convex and labial surface markedly convex. Ante-
rior cuspid with well defined anterolingual and
posterolabial ridges, the latter contributing to a
poorly defined longitudinal crest; poorly developed
cuspule present along anterolingual ridge; post-
erolabial cuspid with well-defined anterolingual
ridge curving and descending to unite with ridge
from anterior cuspid, as continuation of longitu-
dinal crest; crest usually markedly bifid with well
defined labial and lingual grooves present near
mid-point; occasionally two, well-defined labial
grooves are present, in close juxtaposition, cor-
responding with two lingual grooves, producing a
trifid longitudinal crest; grooves sometimes re-
duced; strong, ornamented lingual ridge oc-
casionally descends from posterior moiety of crest
towards posterolingual cuspule; posterior ridge
200
MEMOIRS OF THE QUEENSLAND MUSEUM
from posterolabial cuspule curves into mesial
posterior groove, sometimes uniting with labial
ridge descending from posterolingual cuspule.
Major posterior ridge from that cuspule curves
anterolingually to terminate in slight basin formed
between posterior cuspids; anterior ridge from
posterolingual cuspule descends slightly lingually
to base of crown towards lingual groove from
longitudinal crest. Lingual base of crown some-
times tumescent. Posterior of crown occasionally
with angular ridge present at lingual margin.
DP 3 molariform, subrectangular in basal out-
line, slightly constricted across talonid basin, with
lophids moderately high, convex posteriorly. Hy-
polophid much broader than protolophid. Tri-
gonid basin relatively broad, its length being less
than distance between lophids. Forelink high,
strong, abruptly curving anterolingually from
protoconid to point well labiad to mid-point of
high anterior cingulum; cingulum more anteriorly
extended at anterolingual margin than elsewhere.
Trigonid slopes lingually, and more strongly labi-
ally and posteriorly from forelink and cingular
margin; well-defined anterolabial fossette present
while lingual moiety of trigonid somewhat broadly
V-shaped. Slight ridges descend anteriorly and
posteriorly from metaconid. Hypoconid with
strong, high ridge curving anterolingually across
talonid basin as midlink, uniting with strong
posterior ridge from protoconid, close to pro-
tolophid, labiad to mid-line; weak anterior ridge
from entoconid descends into talonid. Talonid
slopes labially and lingually from midlink; basin
broadly U-shaped in lingual moiety, sharply V-
shaped labially; slight accessory ridges occasionally
present near anterior of crest of hypolophid. Slight
ridge descends posteriorly from entoconid, while
stronger ridge descends from near mid-point of
posterior surface of hypolophid, curving to post-
erolingual margin well above crown base, uniting
with entoconid ridge to delimit well-defined pos-
terior fossette; fossette emphasised by near vertical,
broad groove in posterior surface of hypolophid.
Labial bases of lophids much more expansive than
lingual, giving crown appearance of flexure about
labial limit of talonid; base of crown occasionally
slightly tumescent at labial limit of talonid.
P 3 small, short, subtriangular in occlusal view,
with crown somewhat constricted at anterior one-
third; labial margin slightly convex, while lingual
margin markedly concave; anterior cuspid well-
defined, usually with moderately weak postero-
labial ridge descending as longitudinal crest; ante-
rior ridge poorly defined; posterolabial cuspid also
well-defined with anterolingual ridge normally
descending to unite with other portion of crest
above crown constriction; strong vertical labial and
lingual grooves usually present at anterior one-
third, giving crest a marked bifid appearance in
labial view; grooves sometimes poorly developed
with reduction in V-shaped appearance of crest;
occasionally second set of vertical labial and
lingual grooves present resulting in trifid ap-
pearance of crest; normally central area of crest
depressed, but occasionally near planar where trifid
condition prevails; broad, posterior ridge descends
from posterolabial cuspids towards crown base,
while high, well-defined lingual ridge unites cuspid
with well-defined, but lower posterolingual cus-
pid; this posterior crest usually concave pos-
teriorly, with broad, vertical, posterior groove
present; slight anterior and posterior ridges de-
scend from posterolingual cuspid. Base of crown
normally unornamented.
M x ., petrosal tympanic wing of
periotic; ps., presphenoid; pt., pterygoid; s.e.s., squamosal epitympanic sinus; s.f, sphenorbital fissure or foramen;
stf, stylomastoid foramen; sq., squamosal; t.a.r., tip of ascending ramus; t.c., transverse canal. A-P indicate
measurements given in Table 2.
C. Terminology of the left ear based on WAM M8041 ././., fold lines of pinna; h., helix; pi., pinna; st., supratragus.
D. Terminology of the left hind-foot, based on WAM M6181. a.m.g., anterior metatarsal granule; d2~4, digits 2
through 4; g. of i.p.2 , apical granule of 2nd interdigital pad; ha., hallux; he., heel; h.g., hallucal granule; i.p.2, 2nd
interdigital pad; i.p.3 + g., 3rd interdigital pad and apical granule; i.p.4 + g., 4th interdigital pad and apical granule;
ph.g., post-hallucal granule; ph.p., post-hallucal pad; p.m.g., posterior metatarsal granule; t.p.2-4, terminal pads 2
through 4.
240
MEMOIRS OF THE QUEENSLAND MUSEUM
periotic tympanic wing (from petrosal part) well-
developed; mastoid tympanic wing only just de-
veloped; ectotympanic develops very small tym-
panic wing in forms with smallest alisphenoid
tympanic wing (N. timealeyi); paroccipital does
not develop a tympanic wing although it encloses a
small non-auditory sinus; squamosal develops
small epitympanic sinus continuous with post-
glenoid cavity; an abrupt break occurs in the
periotic’s lateral surface at the level of the horizon-
tal semicircular canal; the mastoid part of the
periotic is swollen laterally, the result of a grossly
enlarged floccular fossa; foramen pseudovale long
and narrow; no true foramen ovale ever develops;
entocarotid foramen large, mesial to foramen
pseudovale, and leads via short canal to en-
docranium; no direct ventral observation through
entocarotid foramen possible; transverse canal
large and pierces basisphenoid mesial to anterior
ends of alisphenoid tympanic wings; transverse
canal foramen leads directly into endocranial
sulcus for entocarotid artery and does not appear
to pass transversely through basisphenoid; post-
glenoid foramen pierces postglenoid cavity im-
mediately posterior to postglenoid process and is
separated by thin transparent bone from the much
enlarged subsquamosal foramen; entire roof of
postglenoid cavity thin, transparent, and functions
solely to floor large postglenoid canal; posterior
surface of postglenoid process almost horizontal
and continuous with roof of postglenoid cavity;
rim of postglenoid foramen penetrated anteriorly
by postzygomatic foramen which leads into zygo-
matic arch; small branch of postzygomatic pene-
trates external surface of zygomatic arch; internal
jugular canal very well-developed between basi-
occipital and periotic with a very steep to vertical
basioccipital wall; internal jugular canal actually
penetrates basioccipital at anterior end; posterior
lacerate foramen passes between paroccipital and
periotic dorso-mesially to paroccipital process;
complete stylomastoid foramen bordered ante-
riorly by periotic with small slip of squamosal on
ventro-lateral rim; small foramen rotundum
floored by variably enlarged shelf of alisphenoid;
large sphenorbital fissure; basisphenoid with
median ventral keel; ectotympanic wide with
marked sulcus and crest associated with pars tensa
of tympanic membrane; anterior (dorsal) end of
ectotympanic helps enclose mesial wall of post-
glenoid canal; anterior end of ectotympanic poin-
ted, posterior end blunt; incus as in Sminthopsis
and other small dasyurids, articulates (not fused)
with malleus, with short incudal and long stapedial
processes, latter with oval lenticular process at
distal end; malleus with pronounced orbicular
apophysis, small anterior dorsal spine on head,
pronounced capitular crest and lamina, and non-
expanded distal tip on manubrium; tubular periotic
hypotympanic sinus and almost total lack of
development of mastoid epitympanic sinus; in-
cudal fossa of epitympanic recess of periotic large,
with well-developed lateral squamosal wall; no
sulcus or canal connects facial nerve canal or sulcus
with postglenoid canal; periotic horizontal antero-
lateral projection from epitympanic recess contacts
postglenoid foramen and almost excludes squa-
mosal from roof of postglenoid cavity; facial nerve
canal large and sulcus well-developed; small for-
amina sometimes (e.g. WAM M6181) pierce mas-
toid tympanic wing; fenestra ovalis subrounded
rather than oval; endocranial structure of periotic
largely unknown because specimens intact but
clear that floccular fossa very large and deep,
internal auditory meatus divided into widely sep-
arate passages for facial and auditory nerves;
squamosal reduced in all directions by surrounding
bones and squamosal hypotympanic sinus also
small; stapes as in other dasyurids, imperforate,
collumnar, and with small posterior process for
attachment of stapedial muscle and tendon; den-
tary shallow ventral to teeth; lateral surface of
ascending ramus wide; angular process long and
slender; mental foramen beneath Mj or M 2 .
Dentition : (Fig. 2 ; Plate 28A, B). 1 1 tallest upper
incisor and set off from I 2 by diastema; I 2 taller-
crowned or subequal to I 3 which is taller-crowned
than I 4 ; I 2-4 subequal in crown length; slight buccal
cingula developed I 1-4 ; I 4 lacks posterior cingular
cusp or lobe; diastema separates I 4 from C 1 ; C 1
taller-crowned than any premolar except some-
times P 4 which may be subequal in size; O-P 4 not
crowded in adult condition, small spaces separat-
ing all teeth; in adult condition P 1 just shorter-
crowned than P 3 which is markedly shorter-
crowned than P 4 ; buccal cingula complete C 3 -P 3
but incomplete P 4 and sometimes incomplete C 1 ;
lingual cingula complete P 1-3 but incomplete C 1
and P 4 ; posterior and anterior cingular cusps
present P 1-4 but only posterior cusp present C 1 ;
anterior cingular cusps may be small on C 1 , P 1 and
P 4 ; P 4 blade-like with convex postero-buccal flank;
DP 4 (WAM M8081) small with two very closely
approximated cusps, paracone and metacone, a
very small and low protocone, and no stylar cusps;
all cusps except protocone linked by longitudinal
buccal crest; M 1 narrower than M 2 which is
subequal to M 3 which is wider than M 4 ; M 1 longer
than M 2 which is longer than M 3 which is longer
than M 4 ; paracone increases in size posteriorly but
is reduced overall relative to most other dasyurids;
ARCHER: N INGA \JI, A NEW GENUS OF DASYURIDS
241
Fig. 2: The terminology of the dentition of Ningaui spp. A, lower right Pj-M 4 . B, upper right P^M 4 . C, lower right M 3 .
D, upper right M 3 .
a.c., anterior cingulum; a.prcr., anterior protocrista or preprotocrista; c.n., carnassial notch; c.o,, crista obliqua; e.,
ectoflexus; end, entoconid; hycd, hyporistid; hyd, hypoconid; hyld, hypoconulid; hyld.n., hypoconulid notch; me.,
metacone; mec., metacrista; meed, metacristid; meed.f, metacristid fissure or cranassial notch; med, metaconid; mst.,
metastylar corner of tooth; mstd, metastylid; pa., paracone; pac., paracrista; pacd., -aracristid; paed.f, paracristid
fissure or cranassial notch; pad, paraconid; p.c.c., posterior cingular cusp; pf, prefossa; p.prer., posterior protocrista
or postprotocrista; pr., protocone; prd, protoconid; prgd, precingulid or anterior cingulum; psgd, postcingulid or
posterior cingulum; pstd, parastylid; stA, stylar cusp A or position of stA; stB, stylar cusp B; stC, stylar cusp C or
position of stC; stD, stylar cusp D; stE, stylar cusp E.
242
MEMOIRS OF THE QUEENSLAND MUSEUM
paracone and metacone approximated; metacone
M 2 taller than subequal metacones M 1 and M 3 ;
metacone M 4 absent; protocone small overall and
decreases in size posteriorly; very slight pro-
toconule development M 1 3 ; no metaconule de-
velopment; protocone basin opens posteriorly;
anterior cingulum complete M 1 only antero-buccal
cingula well-developed M 1 ^ 4 ; no posterior cingula;
stD present M 1 3 and decrease in size posteriorly;
stB present or position at end of paracrista clear
M 1 4 ; stB M 1 variably developed and may even be
absent in some specimens (e.g. WAM M6181)
where stB and stA may be indistinguishable; stE
variably present M 1-3 and if present tiny; stA
indistinguishable M 2-3 and sometimes M 1 ; para-
crista increases in size posteriorly M 1 ' 4 ; metacrista
M 1 just shorter than that of M 2 which is longer
than that of M 3 ; ectoflexus increases posteriorly
from M 1 to M 3 ; Ij taller-crowned and longer than
I 2 which is taller-crowned and larger than I 3 ; I ]_ 3
with slight postero-buccal cingula; poorly-
developed posterior cingulum 1 3 ; no diastema
between I 3 and Cu C, with buccal and lingual
cingula; Cj premolariform but just taller-crowned
than any premolar; Pj shorter-crowned than P 3
which is longer-crowned than P 4 ; Q-P 3 wide and
contrast with relatively narrow P 4 ; small anterior
but well-developed posterior cusps present P, _ 4 ;
well-developed buccal but slightly-developed lin-
gual cingula P^; premolars contact one another
antero-posteriorly but are not crowded; dP 4 two-
rooted with one main cusp, the protoconid, and
possibly a very small posterior cingular cusp; M 2 _4
subequal in length and just longer than M,; talonid
wider, M 2 subequal to, and M 3 _4 narrower
than talonid; protoconid increases in height pos-
teriorly from Mj to M 3 ; protoconid M 4 subequal to
that of M*; metaconid Mj just shorter than
subequal metaconids M 2 paraconid M 1-4 in-
creases in height posteriorly; hypoconid subequal
in height M x 3 but just shorter in M 4 ; entoconid
miniscule and variably absent M 4 ; extremely
tiny metastylid sometimes present (e.g. WAM
M8080) M 2 ^ 3 ; notch for hypoconulid or its hom-
ologue in P 4 occurs in anterior cingulum Mj_ 4 ;
anterior and posterior cingula well-developed
M^; slight buccal cingula formed only on Mu no
lingual cingulum M x _ 4 or posterior cingulum M 4 ;
paracristid M 3 shorter than and directed more
anteriorly than that crest in M^; paracristid M 4
shorter than subequal paracristids M 2 3 ; metacris-
tids Mj 3 increase in length posteriorly; metacristid
M 4 shorter than that of M 3 ; carnassial notch
metacristids well lingual to midpoint of crest;
hypocristid longest in M 2 , subequal in M l and M 3 ,
and shortest or absent in M 4 ; crista obliqua longest
in M 4 and subequal in M t _ 3 ; crista obliqua
intersects trigonid well buccal to point below
metacristid carnassial notch on all molars except
M 4 where this crest intersects trigonid below
metaconid; hypocristid links hypoconid and hy-
poconulid and does not contact very small en-
toconid; metacristid and hypocristid M 2 _ 4 almost
transverse to long axis of cheek-tooth row; differ-
ence in height of smaller paraconid and larger
metaconid decreases posteriorly from to M 4 ,
two cusps being subequal in M 4 ; hypoconulid locks
into gap (hypoconulid notch) of antero-lingual
cingulum of each posterior molar; no cristid links
hypoconulid and entoconid; well-developed fissure
in paracristid and slight fissure in metacristid M 2 _ 4 ,;
these fissures in Mj very small; tiny parastylid
variably developed on M 3 _ 4 .
Discussion
In diagnosing Ningaui comparison has been
made with Sminthopsis and Planigale. There is little
risk of confusion with larger forms such as
Antechinus or Phascogale , or with the long-limbed
form Anteehinomys which in a number of charac-
teristics is very similar to some species of Smin-
thopsis. The species of Anteehinomys share the
same characters with species of Sminthopsis that
enable those species to be distinguished from
species of Ningaui, i.e. long narrow hind feet (in
Anteehinomys there is additionally a total loss of
the hallux) without enlarged post-interdigital gran-
ules, and a broad contact on the outside of the skull
between the squamosal and frontal bones.
The characters used in the diagnosis of Ningaui
are not all absolute because some are not ex-
pressions of exclusive presence in or absence from
Ningaui. Ningaui must be related to other taxa and
it is to be expected that even the diagnostic
characters of the genus will reflect these re-
lationships.
The use of the character of the post-interdigital
enlarged granules in the diagnosis should not be
taken to imply that these granules are invariably
small in Sminthopsis. Some species of Sminthopsis
possess slightly or variably enlarged post-
interdigital granules, but none have all four post-
interdigital granules conspicuously enlarged. For
example in the four known specimens of S.
longicaudata, the three which have feet preserved
show an enlarged hallucal and a single enlarged
metatarsal granule. However, this species is at once
distinguishable from species of Ningaui by its
possession of the other Sminthopsis characters
noted above as well as by the extremely long tail
(twice the length of the head and body). Sminthopsis
murina and S. leucopus are species which do not
ARCHER: NINGAUI, A NEW GENUS OF DASYUR1DS
243
normally possess enlarged metatarsal granules, but
one and even two metatarsal granules, as well as a
hallucal granule, sometimes occur in abnormal
specimens. For example, WAM Ml 854 S. murina
has an enlarged hallucal, post-hallucal and slightly
enlarged posterior metatarsal granule. Some speci-
mens (e.g. Macleay Museum M 1 183) of 5. rufigenis
from the Herbert River in Queensland have a
slightly enlarged oval posterior metatarsal granule
and an enlarged hallucal granule. These species of
Sminthopsis , however, are otherwise unlike species
of Ningaui.
In considering the diagnostic dental characters
of Ningaui, Sminthopsis ooldea (= Sminthopsis
murina ooldea Troughton, 1965) demonstrates
some of the dental characters of species of Ningaui .
For example WAM M8077 has slightly reduced
paracones and talonids, although the degree of
reduction is not as great as that in species of
Ningaui. In all other respects S. ooldea possesses
Sminthopsis characters and is easily distinguished
from specimens of species of Ningaui.
Because of the blurring of these diagnostic
characters between Ningaui and Sminthopsis it is
tempting to regard Ningaui as a possible derivative
or ancestor of that genus. Many of the characters of
Ningaui occur in Sminthopsis as well, where they
occur as arid-adaptations of the structurally more
generalized form. For example, a detailed study of
Sminthopsis (in preparation) reveals that there are
several species-groups which probably have ach-
ieved arid-adaptation independently. Characters
involved in such arid-adaptation include small
body size, relatively short premolar rows, well-
evacuated palates, and high-crowned teeth. Some
of these characters can possibly be interpreted as
mechanical rather than physiological specializ-
ations demanded by thick cuticles of prey species of
that environment. In the case of these very small
mammals, prey species are likely to be insects with
hard cuticles and small vertebrates such as
skinks.To masticate such foods it is necessary to
have teeth mainly adapted to shear; this may be the
reason for the relatively enlarged metacones and
trigonids. However, some characters cannot, at
present, be understood as mechanical adaptations,
such as the lack of contact between the squamosal
and frontal. The presence of a squamosal-frontal
contact has a peculiar distribution among mar-
supials. It is present in some Antechinus macdon-
nellensis, Neophascogale and Phascolosorex spp.,
some Phascolarctos cinereus (Ride 1957), Vom-
batids, all Sminthopsis spp., Thylacinus spp., at
least some borhyaenids (those whose condition
can be determined from figures of Sinclair 1906),
hypsiprimnodontine and potoroine macropodids
(Pearson 1950), peramelids (ibid, and all living
genera checked by the present author), zygomat-
urine, palorchestine and nototheriine diprotodon-
tids (Stirton 1967, Woodburne 1967). It is not
present in other dasyurids, macropodine macro-
podids (Pearson 1950), didelphoids (ibid and
absence confirmed in this study in Didelphis,
Marmosa, Monodelphis, Metachirus, and Philan-
der ), caenolestoids (ibid and confirmed in this study
in Caenolestes), and all other phalangeroids. The
absence of this character from all didelphoids and
caenolestoids does not suggest that, among mar-
supials in general, the character is primitive.
Moreover, among Australian marsupials it is
absent from generalized phalangeroids. If the
presence of a squamoso-frontal contact is primitive
it would seem that a number of marsupial phyla
have achieved it independently from more primi-
tive forms.
I would suggest that the characters of Smin-
thopsis are a derivation from a non-arid-adapted
Ningaui-\ike ancestor.
Origin of Generic Name
The generic name Ningaui is here given mascu-
line gender. It is an Aboriginal name given to tiny
mythological beings that are hairy, have short feet,
and only come out at night to hunt for food all of
which is eaten raw (Roberts and Mountford 1969).
The allusion to these dasyurids involves their very
tiny size, hairy and (compared with the related
dasyurid Sminthopsis spp.) short feet, and noctur-
nal habits.
Species
The genus Ningaui contains two species: N.
timealeyi and N. ridei.
Ningaui timealeyi sp. nov.
(Figs. 1-3; Plates 28A-C, 29A)
Planigale tenuirostris: Ride, 1970, pp. 120, 200 (in part)
(nec Troughton 1928).
Holotype: Western Australian Museum specimen
WAM M6181, young adult female, skull, dentaries, and
carcase in spirit, collected by Mr A. Snell, 7 July 1963,
32-2 km southeast of Mt. Robinson, northwestern
Western Australia. Specimen ‘caught while escaping
burning Spinifex’.
Paratypes: WAM M5076, male collected 1957 (E. H.
M. Ealey, in litt., 19 September 1972) by Dr E. H. M.
Ealey from aerodrome of Abydos Station, W.A.
(21 °25'S 118°54'E). WAM M8041, male, collected July
1969 by Mr T. Fletcher at Kangan Station, W.A.
(21 °09'S 118 G 30'E). WAM M8042, male, collected June
1969 by Mr T. Fletcher from Pilbara Townsite, W.A.
(21 °15'S 118°18'E).
244
MEMOIRS OF THE QUEENSLAND MUSEUM
Referred Specimen: WAM M8729, female, found by
Messrs A. Baynes and M. K. Youngson 15 December
1968, in small cave in breakaway about 30 m above sea
level, on North West Cape near lighthouse (21°48'S
114°6'E). Found freshly killed with head missing,
presumably removed by predator. Specimen referred to
N. timealeyi on basis of foot structure and general
appearance.
Diagnosis
This species differs from N. ridei as follows: Inner
and outer posterior metatarsal granules of hind feet
elongate; distal end of hallux extends to level of or
beyond posterior edge of interdigital pads; hind
foot relatively short (HF/NV ratio between 0T7
and 020); supratragus of ear relatively long antero-
posteriorly; tail exceeds nose-vent length; ali-
sphenoid tympanic wing relatively less well-
developed (M 1-3 /outside bullar distance-inside
bullar distance value 0-69); I 2 length approximately
equals I 4 ; nasals may be slightly expanded pos-
teriorly; uninflated ventral portion of periotic
larger than periotic tympanic wing; paracrista M 4
considerably larger than that of M 3 .
Description
Pelage: Ridgeway (1912) colours for holotype
(spirit specimen) are as follows: side of face Salmon
Colour to Bister; mid-back near Blackish Brown
(3) to Chaetura Black; belly near Massicot Yellow
to Pale Chalcedony Yellow.
Tail: All specimens have thin tails which exceed
nose-vent length (TV/NV ration between 1-13 and
1 36).
Hind Feet: Pads somewhat variable. In holo-
type, posterior metatarsal granule shorter than
post-hallucal granule; in WAM M5076 posterior
metatarsal granule present. Hallucal granule also
varies in similar manner. WAM M5076, left foot,
post-hallucal granule entire; right foot with divided
but small separate granule at anterior end. Gran-
ules barely striate (when held in incident light some
striae reflect light demonstrating very slight surface
expression; in others striae are visible, but without
surface expression, e.g. as in holotype).
Ear: Supratragus of ear relatively long antero-
posteriorly (St/E ratio between 0-25 and 0-29).
Nipple Number: Six (holotype and referred
specimen WAM M8729).
Skull and Dentition: Table 1 gives absolute
measurements and ratio values. Dorsal, ventral
and lateral views of holotype shown in Fig! 3.
Characters of teeth shown in Plate 28 where they
are accurately illustrated by use of scanning
electron microscope. In Fig. 3 damaged areas
illustrated unrestored. For example, portion of left
jugal, both hamular processes of pterygoid, both
alisphenoid tympanic wings and right ectotym-
panic missing or broken. Also portion of right
tympanic wing displaced towards basicranial mid-
line and minor depressed fracture of right ali-
sphenoid. Mild asymmetry of basicranium ac-
curately illustrated. Some basisphenoid foramina
of holotype not prepared sufficiently for illus-
tration and as result condition in WAM M8042
shown in Plate 29. In Fig. 3C dentary slightly tilted
such that tip of ascending ramus rotated
buccally. This shortens apparent distance between
tip of angular process and articular condyle. Nasals
as shown in Fig. 3A appear to expand posteriorly
slightly more than they actually do in holotype. In
Figs. 3B and 3C teeth only approximations and
their characters should be examined in Plate 28.
Figs. 2A and 2B have been made using a camera
lucida.
Habitat
Burbidge (1959) and Ealey (1967) give habitat
information for Abydos Woodstock area which
includes locality of paratype WAM M5076. Bur-
bidge (1959) describes flat plain areas as spinifex
(Triodia) steppe with Acacia pyrifolia, Grevillea
pyramidalis and Eucalyptus dichromophloia. Al-
though Triodia dominates the grasses, other small
plants and bushes occur sporadically. Ealey (1967)
describes area as one of erratic summer rainfall
(between ten and twelve inches per annum) in
which driest months are September to November.
He notes that since 1915 number of native grasses
in area severely reduced as result of stocking with
sheep. Paratype WAM M5076 collected by Ealey
amongst Triodia pungens and T. lanigera. Ealey (in
litt. 19 September 1972) adds that habitat was
shallow sand overlying hard pan with very sparse
corkbark trees. Collection area about half mile
from water and granite outcrops.
Reproduction
Holotype has relatively undeveloped nipples.
WAM M8041 and WAM M8042 juvenile males
collected June and July. Referred specimen WAM
M8729 collected in middle December and probably
lactating. Has very well-developed pouch with six
large nipples.
Origin of Specific Name
The specific name is in honour of Dr E. H. M.
(‘Tim’) Ealey, of Monash University, who besides
collecting the first known specimen of a species of
Ningaui while an officer of the Wildlife Survey
Section of CSIRO, was also responsible for the
collection of specimens of rare and little-known
species such as Antechinus rosamondae from
ARCHER: N INGA UI, A NEW GENUS OF DASYUR1DS
245
Fig. 3: Holotype Ningaui timealeyi sp. nov. (WAM M6181). A, dorsal; B, ventral; C, lateral skull and dentary. Nasals,
left jugal and parts of basicranium damaged. Right ectotympanic and most of left and right alisphenoid tympanic
wings missing. See text for comments. Line represents one mm.
246 MEMOIRS OF THE QUEENSLAND MUSEUM
northwestern Western Australia from 1953 to
1962.
Ningaui ridei sp. nov.
(Fig. 4; Plates 27, 28D, 29B)
Plcmigale ingrami: Ride, 1970, pp. 120, 238, pi. 35 (as
Planigale, close to P. ingrami), (in part) (nee Thomas
1906).
Holotype: Western Australian Museum WAM
M8080, very young adult female with P4 erupting, skull,
dentaries and spirit carcase, caught by Dr and Mrs E.
Pianka, February 1967, 38-6 km along White Cliffs Road
East-northeast of Laverton, Western Australia (28°30'S
I22°47'E).
Paratype: WAM M8081, juvenile male with dP4 in
place, same collection locality as holotype WAM M8080.
Diagnosis
N. ridei differs from N. timealeyi as follows: Post-
hallucal and metatarsal granules generally more
oval than elongate; distal end of hallux does not
reach base of interdigital pads; supratragus outer
edge of ear relatively short; tail- vent length just less
than nose-vent length; alisphenoid tympanic wing
well-developed; periotic tympanic wing relatively
large; I 2 just larger than I 4 and somewhat recurved;
nasals do not appear to expand posteriorly;
paracrista M 4 subequal to paracrista M 3 .
Description
Pelage: Ridgway (1912) colours for holotype
(spirit specimen) as follows: side of face near
Salmon Colour to Buffy Brown; mid-back near
Fiscous-Black to Chaetura Black; belly near Ivory
Yellow to Pale Russian Blue.
Tail: All specimens have thin, relatively short
tails (TV/NV ratio 0-96 and 0-97).
Hind Feet: All hallucal and metatarsal granules
on hind feet of holotype oval. However, hallucal
and posterior metatarsal granules of paratype
more elongate than oval; interdigital pad granules
either long, partially fused series of small median
granules, or shorter somewhat oval apical granule;
left foot of holotype shows questionably fused
median granules on 2nd and 4th interdigital pads
and smaller but isolated apical granule on 3rd
interdigital pad; in paratype, situation similar with
3rd interdigital pad having unfused but enlarged
oval apical granule; 4th interdigital pad appears to
have fused elongated apical granule; 2nd in-
terdigital pad has elongate apical granule but
margins indented halfway along to suggest incom-
plete fusion of median granules; some suggestion of
slightly enlarged granule between hallucal and
apical granule of 2nd interdigital pad of left foot of
paratype; as result, three conspicuous but smaller
inner post-interdigital pad granules present; in
holotype only two, a hallucal granule and one
between it and apical granule of 2nd interdigital
pad.
Ear: All specimens have relatively short outer
edge of supratargus (St/E ratio 0-20 and 0-22).
Nipple Number: The holotype is a young female
with seven barely distinguishable nipples. The
referred specimen is a juvenile male.
Skull and Dentition: Holotype just barely
adult (P4 practically fully erupted); paratype
TABLE 1 : External Measurements (mm) of Ningaui timealeyi and N. ridei gen. et spp. nov.
Species
TV
NV
HF
E
St
TV/NV
HF/NV
St/E
N. timealeyi
WAM M6181
62-0
55-0
10-0
10-6
3-0
1.13
0-18
0-28
WAM M8041
680
?
10-0
11-7
3-4
1 36
0-20
0-29
WAM M8042
79-0
?
10-0
—
—
1-32
0-17
—
WAM M5076
650
57-0
11-0
11-5
2-9
1-14
0-19
0-25
WAM M8729
650
—
9-0
—
—
—
—
—
X
67-8
56-0
100
11-3
3-1
s
6-61
1-41
0-71
0-59
0-27
V
9-75
2-52
7-10
5-22
8-71
N. ridei
WAM M8080*
50-6
53-0
11-6
10-9
2-2
0-96
0-21
0-20
WAM M8081*
48-0
49-7
11-3
11-3
2-5
0-97
0-23
0-22
X
49-3
51 4
11-5
111
2-4
^Indicates juvenile or very young adult specimens whose absolute body measurements are probably not
indicative of adult body measurements. Abbreviations: TV = tail tip to cloacal vent; NV = nose tip to
cloacal vent; HF = hind foot from heel to toe tips not including claws; E = ear from base of notch to tip of
pinna; St = supratragus maximum length.
ARCHER: N INGA VI, A NEW GENUS OF DASYURIDS
Fig. 4: Holotype Ningaui ridei sp. nov. (WAM M8080).
Damage described in text. Line represents one mm.
juvenile (dP4 in place). Absolute and relative ratio
values given in Table 1. As noted above, the
periotic and alisphenoid tympanic wings of N. ridei
are more enlarged than those of N. timealeyi (Plate
29). It is possible that in older individuals of N. ridei
this difference may become even greater.
Habitat
The only two specimens were caught by Dr and
Mrs Pianka in February 1967. The holotype was
caught in a pit trap dug for lizards. The paratype
was collected in spinifex (Triodia). Pianka and
Pianka (1970) describe the area within which the
holotype and paratype were collected, as open
savannah woodland with gently rolling red sand
plains, and low lying Triodia meadows. The
woodland consists in part of mallee eucalypt and
Eucalyptus gongylocarpa, some of which grow to 1 5
m in height, as well as Acacia sp. and a ground
cover of Triodia.
247
^Indicates measurements made on juvenile or very young adult specimens which are probably smaller than corresponding measurements made on adult specimens. For
example, antero-posterior growth of rostrum and dentary increases A, E, K and L. Similarly, growth of cranium in lateral direction increases B, C, D, H, and I.
248
MEMOIRS OF THE QUEENSLAND MUSEUM
Ningaui ridei in Captivity
The holotype was kept in captivity until 1 3 April
1967, during which time it was photographed. It
was also drawn by Mrs E. Fry and forms the basis
of plate 35 in Ride (1970). The photographs (Plate
27) reveal that in some respects the living animal
does not clearly resemble species of either Smin-
thopsis or Planigale. The body is covered with
guard hairs which give the animal a bristly
appearance, not unlike that of Antechinus apicalis ,
but markedly unlike that of any Sminthopsis I have
seen. It does resemble the pelage condition of
Planigale gilesi as suggested by photographs in
Aitken (1972). The photographs show that the foot
width is broader than that of Sminthopsis but
distinctly narrower than the feet of Planigale. The
animal has been photographed while eating a large
grasshopper (Austracris guttulosa, in litt. J. H.
Calaby, 1973). Although the initial seizing of the
grasshopper’s head is made with the front of the
mouth, appendages and projecting body parts are
subsequently shifted to the molar region of the
dentition where they are sheared off by the large
metacristae and paracristids. Dr W. D. L. Ride
(pers. comm.) noted that the jumping legs of the
grasshopper were quickly severed, perhaps as a
way of immobilizing the animal.
Discussion
The holotype of N. ridei is a young adult animal
and the paratype a juvenile. All of the specimens
representing N. timealeyi are adult. Because of this
age difference, many of the cranial ratios given in
Table 1 are not directly comparable between the
two species of the genus.
There are other characters of the canine and
premolar region which, when more specimens
referable to N. ridei are available and the degree of
variation is better understood, may prove to be
diagnostic. For example, in the only specimen of N.
ridei in which P 4 is visible, it considerably exceeds in
length the crown of P, . In N. timealeyi on the other
hand, the P 4 crown is approximately the same
length as that of Pj.
There also appears to be a difference in the shape
of the premolars. In N. ridei P 1 appears to be
broader and more massive in appearance than P 1 of
N . timealeyi. Similarly, and P! of N. timealeyi
appear relatively more elongate and narrower than
those teeth in N. ridei.
The difference in nipple number may, when more
specimens of both species are know, prove to be
diagnostic. At the moment these characters are not
listed in the diagnosis because it is not clear how
much variation will be demonstrated by larger
samples.
Origin of Specific Name
The specific name is in honour of Dr W. D. L.
Ride, who recognized the complexity and possibly
polyphyletic nature of the small marsupials earlier
referred to Planigale, including the forms described
here as species of Ningaui.
RE-DIAGNOSIS OF PLANIGALE
The recognition of the difference between species
of Ningaui and those of Planigale enables the genus
Planigale to be re-diagnosed as follows:
Dasyurids smaller than Antechinus and differing
from these in having an extremely reduced maxil-
lary vacuity; very small paracone; and small talonid
on M i _ 3 . Generally smaller in size than Smin-
thopsis and Antechinomys and differing from these
in possessing a straight external edge on the
supratragus of the ear; short broad pentadactyl
hind feet; enlarged metatarsal granules; broadened
nasals; lack of squamosal-frontal contact on the
outside of the skull; posterior cingula on M 1-3 ;
reduced paracone, protocone, and talonid on
Ml-3; single-rooted or absent P 4 ; and lack of a
palatine vacuity.
Under this concept, Planigale contains P. in-
grami. P. macu/ata, P. suhtilissima, P. tenuirostris.
P. novaeguineae and P. gilesi (discussed as part of a
revision of Planigale, in preparation).
ACKNOWLEDGMENTS
While this study was made, the author alter-
nately held Fulbright Scholarships, a grant in aid
from the American Explorers’ Club, and a Re-
search Assistantship to Dr W. D. L. Ride who was
in receipt of a Research Grant from the Australian
Research Grants Committee. Dr W. D. L. Ride,
Director of the Western Australian Museum and
Dr M. O. Woodburne of the University of
California at Riverside kindly read and crtiticised
drafts. Mr J. Hardy of the University of Queens-
land helped take the scanning electron microscope
photographs. Mr A. Easton, Queensland Museum,
helped produce photographs not taken by Mr
Hardy or Dr Ride. Mrs P. Johnson, formerly of the
Western Australian Museum, produced the draw-
ings for Figs. 3-4. Dr E. H. M. Ealey, Monash
University, kindly provided information about the
capture of specimens of N. timealeyi. Mrs C.
Farlow and Miss P. Rainbird of the Queensland
Museum, typed drafts of the manuscript.
ARCHER: NINGAUI , A NEW GENUS OF DASYURIDS
249
LITERATURE CITED
Aitken, P. F., 1972. Planigale gilesi (Marsupialia,
Dasyuridae), a new species from the interior of south
eastern Australia. Rec. S. Aust. Mus. 16 : 114.
Archer, M., 1974. The development of the cheek-teeth in
Antechinus flavipes (Marsupialia, Dasyuridae). J. R.
Soc. West. Aust. 57 : 54-63.
Gould, J., 1863. ‘The mammals of Australia’. Vol. 1,
Introduction, 40 pp. (The Author: London).
Gray, J. E., 1843. ‘List of the specimens of mammalia in
the collection of the British Museum’, 216 pp.
(British Museum: London).
Macleay, W. S., 1841. Notice of a new genus of
Mammalia discovered by J. Stuart, Esq., in New
South Wales. Am. Mag. nat. Hist. 8: 242,
Pearson, J., 1949. The relationships of the Potoroidae to
the Macropodidae (Marsupialia). Pap. Roy. Soc.
Tas 1949 : 211-29.
Pocock, R. I., 1926. The external characters of Thyla-
cinus, Sarcophilus, and some related marsupials.
Proc. zool. Soc. Lond. 68: 1037-84.
Ride, W. D, L., 1957. Protemnodon parma (Waterhouse)
and the classification of related wallabies ( Pro-
temnodon, Thylogale, and Setonix). Proc. zool. Soc .
Lond. 128 : 327-46.
1970. ‘A guide to the Native Mammals of Australia’,
xiv + 249 pp. (Oxford University Press: Melbourne).
Roberts, A., Mountford, C. P., 1969. The dawn of
time’, 79 pp. (Rigby Limited: Adelaide).
Sinclair, W. J., 1906. Mammalia of the Santa Cruz beds.
Marsupialia. Rep. Princeton Univ. Exped. Patagonia,
Princeton , N.Y. 1896 1899 4 : 333-460.
Stirton, R. A., 1967. The Diprotodontidae from the
Ngapakaldi Fauna, South Australia. Bur. Min.
Resour. Aust. Bull. 85: 1 44.
Tate, G. H. H., 1947. On the anatomy and classification
of the Dasyuridae (Marsupialia). Bull. Amer. Mus.
nat. Hist. 88: 101-5.
Thomas, O., 1888. ‘Catalogue of the Marsupialia and
Monotremata in the collection of the British
Museum (Natural History)’, 401 pp. (British
Museum: London).
1906. Proc. zool. Soc. Lond., Abstract 1906 (32): 6.
Troughton, E. Le G., 1928. A new genus, species, and
subspecies of marsupial mice (Family Dasyuridae).
Rec. Aust. Mus. 16 : 281-8.
1965. A review of the marsupial genus Sminthopsis
(Phascogalinae) and diagnoses of new forms. Proc.
Linn. Soc. N.S.W. 89: 307-21.
Waterhouse, G. P., 1846. ‘A natural history of the
Mammalia. Vol. I. Marsupiata or pouched animals’,
553 pp. (Hippolyte Bailliere: London).
Westwood, J. O., 1840 ‘An introduction to the modern
classification of insects. Vol. I, Synopsis’, 158 pp.
(Longman, Orme, Brown, Green, and Longmans:
London).
Woodburne, M. O., 1967. The Alcoota Fauna, Central
Australia. Bur. Min. Resour. Aust. Bull. 87: 1-187.
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 27
Holotype (WAM M8080) N. ridei photographed live shortly
after capture. A and B show bristly appearance, relatively broad
hind foot, fold line of ear, curled supratragus, and use of
anterior molar region (presumably large metacones) to cut
locust’s leg (photographs, W. D. L. Ride).
ARCHER: N1NGAUI, A NEW GENUS OF DASYURIDS
Plate 27
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 28
A, B. Scanning electron microscope photographs (composite
overlays) of teeth of N. timealeyi (WAM M8041). Line is 1 mm
in length. A, RIj-M*. B, RP l -M 4 .
C. D, Left hind foot of Ningaui spp. C, Holotype (WAM
M6181) N. timealeyi. D. Holotype (WAM M8080) N. ridei.
Ruled lines are 1 mm apart.
ARCHER: NINGAUI, A NEW GENUS OF DASYURIDS
Plate 28
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 29
A. B. Ventral view of skulls of Ningaui gen. nov. A, N.
timealeyi sp. nov. (WAM M8042). B, N. ridei sp. nov. (WAM
M8080). Outlines show differences in basicranial region of two
species, and in particular, relative development of alisphenoid
tympanic wings. Line represents one cm.
ARCHER: N1NGAUI, A NEW GENUS OF DASYURIDS
Plate 29
Mem. QdMus. 17(2): 251 -65, pis. 30--5. [1975]
ABNORMAL DENTAL DEVELOPMENT AND ITS SIGNIFICANCE IN DASYURIDS
AND OTHER MARSUPIALS
Michael Archer
Queensland Museum
ABSTRACT
Abnormal dental developments in dasyurids and other marsupials are described, including
supernumerary and lost teeth, divided and fused crowns, abnormal molar crown morphology,
and accidents of development including abnormal tooth positions or postures and malocclusion.
Instances of ephemeral teeth' are also given although these are not regarded as abnormal dental
developments. The literature of abnormal dental development in non-marsupials as well as
marsupials is briefly reviewed. Premolar number variation is not considered a valid means for
determining the position of a possibly suppressed fourth premolar in marsupials. P5 and M5
occurrences are regarded as indications of the proliferative potential of the posterior end of the
dental lamina. Some supernumerary teeth are regarded as atavisms. Evidence is given that
supernumerary teeth may originate as divided tooth crowns. Fused crowns are probably the
result of damage to crowded, developing tooth germs. Some abnormal molar crown variations
are also probably the result of antero-posterior compression of the developing tooth germ.
Examples are given of abnormal tooth shape and number which may be the result of disease.
Some abnormal developments in tooth crown shape and occlusion are more frequent among
inbred individuals. Many examples of ephemeral teeth noted are regarded as normal, representing
rarely noted vestigial milk-teeth or canine teeth in the process of phylogenetic suppression.
Although most abnormal dental developments occur in one tooth only, some occur with occlusal
and/or bilateral counterparts. These facts suggest that most single-tooth abnormalities are not
genetically determined, in contrast to bilateral and occlusal pair abnormalities which may develop
under the influence of a dental morphogenetic field.
Skulls and dentitions of 2990 individuals of
dasyurids have been examined, representing many
stages of dental development. Observations on
non-dasyurids have been more limited. Abnormal
developments (i.e. those outside limits given with
taxonomic descriptions, as noted for example by
Archer 1975) in dasyurids and other marsupials
may be placed in three categories: (1) super-
numerary or lost teeth, loss not obviously being the
result of accident or physical disturbance; (2)
morphological abnormalities including misshapen
tooth crowns, extra cusps, and composite odon-
tomas; (3) developmental accidents including teeth
erupting in unusual positions or postures, tooth
loss resulting from physical disturbance or disease,
and malocclusion. These categories are not mutu-
ally exclusive. For example, a supernumerary
molar may also have a morphologically abnormal
crown.
This review is based (a) on personal observations
of specimens housed in collections throughout
Australia, in the British Museum, and in the
American Museum of Natural History, and (b) on
literature references. For each abnormality, re-
cords from these two sources are listed in separate
paragraphs. Literature citations are by author and
date except that references to Bateson (1894) are so
numerous that they are given only as the case
number listed by Bateson (e.g. case no. 366).
Catalogue numbers are prefixed by letters as
follows: Australian Museum, e.g. AM M4343;
American Museum of Natural History, e.g.
AMNH 109524; British Museum (Natural His-
tory), e.g. BM No. 2.9. 8.7; Butler collection housed
in the Western Australian Museum, e.g. B1944;
National Museum of Victoria, e.g. C1009; Fish-
eries and Wildlife Department of Victoria, e.g.
D966; Queensland Museum fossil collection, e.g.
F4713; Queensland Museum modern mammal
collection, e.g. J23087 or JM169; South Australian
Museum modern mammal collection, e.g. SAM
M7536; Northern Territory Museum, e.g.
252
MEMOIRS OF THE QUEENSLAND MUSEUM
NTM274; Western Australian Museum modern
mammal collection, e.g. WAM M2477; Western
Australian Museum fossil vertebrate collection,
e.g. WAM 68.3.43; Queen Victoria Museum and
Art Gallery modern mammal collection, e.g. QVM
1964.1.33.
Literature references to non-erupting teeth or
teeth so small they have usually escaped detection
in description are mentioned here and although
these are not abnormal developments, some further
examples are given.
The purposes of this study are twofold. Firstly,
to attempt to discover whether ‘atavisms’ occur at
any regular positions suggesting sites of earlier
evolutionary losses from a toothrow. Secondly, to
describe abnormal structural developments in
various populations so that abnormal individuals
(e.g. unique fossils like the Fromm’s Landing
Thylacine, Archer 1971) can be more easily eval-
uated.
Molar cusp terminology is that used by Archer
(1974, 1975). Tooth nomenclature is that used by
Thomas (1888). Australian marsupial names are
those employed by Ride (1970) and Archer (1975).
Sminthopsis sp. (a) and (b) refer to two species
which will be described elsewhere. New Guinean
marsupial names are those employed by Laurie and
Hill (1954).
DASYURIDAE
Material Examined and Number of
Abnormal Developments: Incidence (in brackets)
of abnormalities, other than ephemeral teeth,
follows the number of individuals examined in the
samples listed below.
Sarcophilus harrisii 17 (3); Dasyurus hallucatus
64 (2); D. maculatus 33 (1); modern D. geoffroii 45
(2); fossil D. geoffroii 17 (0); D. viverinnus 11(1); D.
albopunctatus 3 (0); D. dunmalli 4 (0); Murexia
longicaudata 2 (0); Myoictis melas 3 (0); Neophas-
cogale lorentzii 3 (1); Phascolosorex dorsalis 5 (0);
P. doriae 4 (0); modern Dasycercus cristicauda 24
(2); fossil D. cristicauda 574 (4); Dasyuroides byrnei
46 (6); modern Antechinus flavipes 41 (5); fossil A.
flavipes 252 (0); modern A. apicalis 5 (1); fossil A.
apicalis 77 (2); A . stuartii 1 0 (0); A . swainsonii 1 0 (0);
A. bellus 14 (2); A. macdonnellensis 38 (0); A.
rosamondae 10 (0); A. godmani 3 (0); Phascogale
tapoatafa , (in access of) 50 (5); modern P. calura 19
(I) ; fossil P. calura 31 (0); Planigale maculata 43
(II) ; P. ingrami 33 (1); P. subtilissima 7 (0); P.
tenuirostris 14 (1); P. gilesi 5 (0); modern Smin-
thopsis murina 140 (7); fossil S. murina 615 (0); S.
ooldea 13 (1); S. leucopus 69 (9); S. crassicaudata
143 (7); modern S. granulipes 6 (1); fossil S.
granulipes 207 (0); S. psammophila 4 (0); S.
longicaudata (modern and fossil) 8 (0); S. hirtipes 1
(0); S. virginiae 37 (1); S. macroura 100 (6); S. sp. (a)
7 (1); S. sp. (b) 3 (2); modern Antechinomys spenceri
30 (1); fossil A. spencerae 1 70 (1); ^4. laniger 12 (0);
Ningaui ridei 2 (0); and N. timealeyi 4 (0).
The number of abormal dental developments
recorded above is certainly an underestimate of the
true number because many specimens examined
have worn teeth and wear obliterates some morph-
ological abnormalities. Also because fossil speci-
mens examined are generally incomplete, like-
lihood of detecting abnormal dental developments
in individuals is markedly reduced.
Tooth Number: Adult tooth formula of-JH'l
is regarded as structurally ancestral for marsupials
in general. In living dasyurids the maximum adult
tooth formula is -f rli- I n most juvenile
dasyurids there is also a molariform tooth, dP4,
displaced from the tooth row by the erupting P4.
Normally non-erupting tooth rudiments (of milk
teeth) also develop ontogenetically, lingual to the
incisors and canine in dasyurids (Archer 1974).
In Mrymecobius (a dasyuroid considered below
with dasyurids) there are eight cheek-teeth in the
adult dentition. Tate (1947) suggests that one of
these is dP4 which persists b ’gether with P4.
Supernumerary Teeth
Incisors: Mrymecobius fasciatus: Two teeth in
position of LI 3 (case no. 366); extra incisor on
lower right side (case no. 367 and noted by Bensley
1903). Dasyurus spp.: Swellings interpreted as
tooth germs between I 2 And I 3 , in front of I : , and
between l 1 and I 2 in sectioned specimens of D.
maculatus and D. viverinnus (Woodward 1896).
Antechinus sp.: Supernumerary tooth positions in
sectioned specimens between I 2 and I 3 , between Ij
and I 2 , and in front of Ij and these positions were
distinguished from deciduous tooth germs (Wood-
ward 1896). Deciduous teeth associated with all
incisors (Archer 1974).
Canines: Antechinus flavipes W AM M71 1 1 small
accessory LC 1 ; A. minimus D966 small accessory
LC 1 ; Sminthopsis crassicaudata J 14376 two teeth at
LC 1 position. Supernumerary canines observed in
this study may be abnormally enlarged deciduous
canine tooth rudiments, such as in Antechinus
(Woodward 1896, Archer 1974).
Premolars: Dasyurus geoffroii WAM M4464
tooth between RP 1 and RP 3 ; Phascogale calura
WAM M8069 tooth either anterior to LP 1 or
between LP 1 and LP 3 ; Antechinus flavipes WAM
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
253
M7107 tooth posterior to L and RP 4 ; WAM
M6785 extra RP x , homology uncertain, appears to
be anterior to RP 3 ; A. bellus NTM274 two teeth in
LP 3 and RP 3 positions; A. apicalis J1741 tooth
antero-lingual to LP 1 ; WAM 64.10.47 extra P,
homology uncertain; Sminthopsis leucopus C891
tooth posterior to RP 4 .
Dasyurus spp.: Swelling of dental lamina in
sectioned specimens of D. maculatus and D.
viverinnus, between PI and P3, interpreted as
rudiments of P2 (Woodward 1896). Phascolosorex
dorsalis : Premolar between LP 1 and LP 3 (case no.
386). Antechinus sp.: Swelling of dental lamina in
sectioned specimen between PI and P3 interpreted
as rudiments of P2 (Woodward 1896).
DP4: Sminthopsis leucopus D524 spicule between
RdP 4 and RM 1 .
Molars: Sarcophilus harrisii: Possible occur-
rences of M 5 discussed below as divided teeth. Fifth
molar (Green 1967). Dasyurus maculatus: Five
upper left molars and five molars on both sides of
lower jaw (case no. 385). Bateson (1894) interprets
these as LM 5 and L and RM S . I agree with
Bateson’s interpretation. Thomas (1888) describ-
ing same specimen, notes that besides being very
small, specimen has asymmetrical squamosal bone.
Missing Teeth
Premolars: Dasyurus hallucatus J16753 missing
RP 3 ; Dasycercus cristicauda J23101 missing RP 3 ;
Antechinus minimus D968, D967 missing RP 4 ;
Planigale maculata J 10989 missing L and RP 3 ;
Sminthopsis sp. (a) B 1939 missing LP 1 and possibly
RP 3 .
Antechinus flavipes: In sample of seven speci-
mens, one lacked P 3 (case no. 387).
Molars: Sarcophilus harrisii WAM 71.10.209
missing all LM X .
Divided Crowns
Incisors: Sminthopsis crassicaudata BM No.
2.9. 8. 7 L and RI 4 crowns bicuspid. Roots partly
divided.
Premolars: Dasyurus geoffroii WAM M4464
RP 3 partly divided. LP 3 has transverse groove;
Dasycercus cristicauda WAM 68.9.91 LP 3 divided
and tooth has three roots; J23098 RP 3 crown tip
divided; Phascogale tapoatafa WAM M7453 LP 3
crown divided and tooth has three roots; WAM
Ml 338 RP 3 crown divided; Sminthopsis
crassicaudata J11388 RP 3 tip of crown divided; S.
murina WAM M 1 642 RP 4 crown partially divided.
S. leucopus AM M4343 LP 3 partially divided; D793
LP 4 partially divided and has three roots; D741,
D458, Cl 00 19 RP 3 has partially divided crown.
Dasyurus geoffroii RP 4 partly divided along
plane transverse to long axis of jaw (case no. 383).
DP4: Sminthopsis leucopus D524 spicule between
RdP 4 and RM 1 may represent split portion of
RdP 4 ; S. rufigenis AM M6562 LdP 4 crown tip
divided.
Molars: Sarcophilus harrisii QVM 1964.1.201
LM 4 may be completely divided producing two
small teeth.
Fused Crowns
Premolars: Dasycercus cristicauda WAM
69.6.269 P 1 and P 3 crowns fused (an isolated
maxilla); Dasyuroides byrnei J 1 1510 RP 1 and RP 3
crowns fused, sharing one root; Antechinus bellus
CSIRO (Canberra) CM1 141 LP 1 and LP 3 fused on
point of contact; Planigale maculata J 10989 RP 4
and RP 3 almost completely fused; Planigale sp.
J 14089 LP 1 and P 3 fused at base of cfowns;
Sminthopsis murina WAM M2046 LPj and P 3
fused.
Abnormal Crown and Root Morphology
Incisors: Sminthopsis crassicaudata BM No.
2.9. 8.7 bilobed L and RI 4 .
Premolars: Sarcophilus harrisii WAM 71.10.209
grossly abnormal tooth in position of
LP’’, Phascogale tapoatafa WAM M7951, WAM
M1338 RP 4 has one root; Dasycercus cristicauda
WAM 68.9.91 LP 3 has three roots; Antechinus
flavipes B 1 8 14 RP 1 caniniform and also tallest
premolar; Planigale tenuirostris AM M5438 P 4
two-rooted; P. ingrami J7656 LP1*3 have buccal
notches or imperfections in cingula; P. maculata
WAM M420 RP 4 has extra large cusp posterior to
paraconid; Sminthopsis leucopus D793 LP 4 re-
sembles enlarged and divided dP 4 ; S. macroura AM
M4403 RP 4 has very large antero-lingual cingular
shelf; S. sp. (a) B1939 tooth posterior to LP X
(topographic homologue of P 3 ) morphologically
resembles molarized protoconid such as occurs on
M 4 ; S. murina WAM M6998 L and RP 3 _ 4 possess
small postero-lingual cuspids.
DP4: Sminthopsis leucopus D524 RdP 4 mildly
deformed, possibly result of split-off corner.
Molars .Sacrophilus harrisii QVM 1964.1.134
M 4 very small; Dasyurus geoffroii WAM M4464
talonids very reduced, trigonids enlarged, parti-
cularly M 3 ; D. maculatus J 1 6744 LM 4 either part of
254
MEMOIRS OF THE QUEENSLAND MUSEUM
composite dental odontoma or grossly abnormal;
Dasyuroides byrnei J 1 1 509 L and RM 3 ectolophs
compressed antero-posteriorly; Planigale maculata
J 16721 small buccal cusp on talonid near base of
protoconid; Antechinus apicalis unregistered LM 1
paracone slightly displaced, paracrista absent, and
extra cusp present anterior to displaced paracone;
A. flavipes WAM M8092 M 2 has prominent
antero-lingual cusp on flank of protocone; Smin-
thopsis macroura WAM M6903 L and RM 1 ” 3
ectolophs compressed antero-posteriorly; J7407 L
and RM 1-2 have conspicuous protoconules; WAM
M5701 very large entoconid M 4 ; WAM M541 1
M 4 talonid has lingual cuspules and M x has tiny
cusp between paraconid and metaconid; J23555 R
and LM 3 ectolophs compressed antero-posteriorly
with buccally displaced parastyles; S. ooldea WAM
M5888 LM X has cusp between paraconid and
metaconid and tiny cusp in position of entoconid
RM x _ 3 ; S. murina WAM M2477 L and RM^j
have large entoconids; SAM M7536 LM X has cusp
anterior-buccal to paraconid; S. leucopus C6343
M 1 ” 3 ectolophs compressed antero-posteriorly;
Cl 009 M 1 _ 4 have tiny metastylids; AM M4343
Mj_ 3 have tiny entoconids; C9566 L and RM 4
abnormally shaped; S. crassicaudata WAM M373
M 4 talonid tricuspid; S. sp. (b) J5459 RM 3
entoconid split transversely; Antechinomys spen-
cerae WAM 68.2.265 RM 2 with small entoconid;
WAM M2860 M 3 has two tiny cusps in topog-
raphic position of entoconid.
Dasyurus spp.: RM 4 of D. viverinnus larger than
normal sized LM 4 (case no. 384). Specimen of D.
maculatus with supernumerary molars (case no.
384, see above). Bateson interprets (I believe
correctly) supernumerary molars as LM 5 , LM 5 and
RM S . LM 4 and L and RM 4 enlarged and mor-
phologically abnormal. Tooth in position of LM 4
more closely resembles normal M 3 , and teeth in
positions of L and RM 4 b< >th resemble normal M 3 .
Abnormal Molar Stylar Cusps: Dasyurus hal-
lucatus WAM M8085 stylar cusp B distinct on L
and RM 1 ; D. viverinnus J20413 stylar cusp B
distinct on L and RM 1 ; Phascogale tapoatafa
WAM M2855 stylar cusp C occurs between B and
D on M 1 ; WAM Ml 338 stylar cusp C large on
anterior flank of D on L and RM 1 ; WAM M6390
stylar cusp anterior to D on L and RM 1 ; Neophas-
cogale sp. AMNH 109524 stylar cusp D divided
transversely on RM 1 ; Planigale maculata WAM
M420 prominent stylar cusp A on RM 1 ; J19668
stylar cusp C on L and RM 1 ” 3 ; J16722 tiny stylar
cusp E on M 1 ” 3 (uncommon but occurs also in
three other specimens); Sminthopsis sp. (b) J5173
extra stylar cusps between B and D on L and RM 1
and LM 2 ; S. granulipes WAM M6062 stylar cusp D
divided transversely on L and RM 1 ” 2 ; S. murina
WAM M8652 stylar cusp D unusually enlarged on
L and RM 1 , projecting buccally; S. leucopus C6343
stylar cusp A on M 1 , extra stylar cusps between B
and D on RM 3 , and extra stylar cusp on M 4 ; C9566
L and RM 4 have two stylar cusps each.
Eruption and Occlusion
Sminthopsis crassicaudata: WAM M4503 has
what appears to be reversed (antero-posteriorly)
LP 4 . Tooth slightly smaller than RP 4 , WAM
M4497 shows abnormal occlusion resulting from
very short lower jaw. RC 2 bites behind RC 1 and
RC 1 occludes with RI 3 . LC 3 just passes anterior to
LC 1 . Upper incisors do not occlude. Planigale
maculata: J8070 has lower canines, premolars and
molars heavily worn, while upper cheek-teeth
almost unworn. Reasons unknown. Possible that
specimen includes wrongly associated dentary.
Phascogale calura: WAM M8069 maloccluded.
RM 1 rotated out of position. Result is hypoconid
of RMj opposes, rather than shears anterior to,
metacone of RM 1 and protoconid of RM X bites
into protocone basin of RM 1 , rather than anterior
to protocone. Specimen also possesses super-
numerary left premolar (see above). Dasyuroides
byrnei: J 1 1433 missing LM 1 while spaces set RM 1
and RM 4 off from RM 2-3 . Left and right lower
molar rows crowded antero-posteriorly so that
lower molars on both sides distorted out of
position. Teeth maloccluded. J10935 has maloc-
cluded RI 2 which bites lingual to lower incisors.
J 1 1 509 maloccluded. LM 1 deflected postero-
lingually and hypoconid of LM X passes over tip of
metacone producing abnormal wear facet. LM2
also distorted. LM 2 protoconid occludes with LM 2
protocone basin, rather than shearing past it with
very large wear facet across LM 2 protocone and
paracone. Series of thirty Dasyuroides byrnei
specimens (including all noted above) in Queens-
land Museum were bred in captivity. Comparison
of measures of brachycephaly (maximum skull
width/maximum skull length) in wild-caught and
laboratory-bred individuals indicates higher in-
cidence of brachycephaly in latter. Concurrently,
much higher incidence of dental abnormalities such
as malocclusion, tooth loss, supernumerary pre-
molars, and antero-posteriorly compressed molars
evidenced in laboratory-bred samples (Archer and
Vernon in preparation).
Disease or Trauma
Dasyurus maculatus: J 16744 has complex LM 4
which may be either composite odontoma or
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
255
teratoma, possibly result of disease. Dasycercus
cristidauda: WAM 69.6.165 dentaries fused at
symphysis. Antechinomys spencerae: J23103 isol-
ated right dentary has RIi_ 3 , C x and M 2 _ 4 with
alveoli for M l . No trace of any premolar. Dentary
in area where premolars missing has roughened
surface. Large abscesses occur beneath and
below posterior root of M 4 . Abscess below M x has
perforated buccal surface of dentary below al-
veolar margin of Case may be example of
partial anodontia following disease. Sminthopsis
crassicaudata: WAM M8082 missing LI 3 , Cj and
P, . These possibly lost during life, with alveoli then
overgrown by bone.
Normally non-erupting or Ephemeral Teeth
Dasyurus geoffroii: WAM M6370 rudimentary
spicule-like tooth in position of normally absent
RP 4 ; Dasyuroides byrnei J 11435 tiny calcified
rudimentary tooth adhering to postero-lingual
corner LP 3 . Dasyurus, Myrmecobius, Antechinus:
Calcified rudiments or positions for milk-teeth
associated with incisors of Dasyurus spp., Myrme-
cobius fasciatus, and Antechinus (species not given)
in sectioned material (Woodward 1896). Milk-
tooth rudiments associated with Cl in Dasyurus
maculatus, D. viverinnus, Antechinus sp., and Myr-
mecobius fasciatus in sectioned material (Wood-
ward 1896). Dasyurus hallucatus: DP 4 (Tate 1947).
DP4 and P4 normally absent Dasyurus (except D.
dunmalli). Antechinus flavipes: Milk-canine tooth
rudiments and non-erupting milk-tooth rudiments
in association with incisors in sectioned material
(Archer 1974). Examples noted above of super-
numerary canines may represent abnormal de-
velopment of normally non-erupting milk-canines.
THYLACINIDAE
Abnormal molar cusps, supernumerary stylar
cusps, enlarged basal cingula and proximation of
paracone and metacone of upper molar in Thyla-
cinus spp. (Archer 1971). Thylacinus cynocephalus
with four lower premolars (Rose 1892). Calcified
but rudimentary tooth in T. cynocephalus in-
terpreted as dP 4 (Flower 1868).
PERAMEL1DAE
Suggestion of undeveloped incisor tooth pos-
ition between I 3 and I 4 (Woodward 1896). Similar
tooth rudiments in Perameles (Wilson and Hill
1897). Echymipera: In 22 per cent of skulls,
supernumerary upper incisor present (Ziegler
1971). Ziegler interprets this as I s which normally
present in most other peramelids. In series of six
skulls of Echymipera rufescens (J 123063-8), no
examples of supernumerary incisors or other dental
abnormalities. Isoodon obesulus: J23082 shows
morphologically abnormal L and RM,. RM X has
normal talonid. Anterior to this are five principal
cusps. Anterior three may represent trigonid. Two
large accessory cusps also present on tooth, one
(normal on some specimens of I. obesulus) anterior
to hypoconid on buccal edge of crown and other
posterior to possible homologue of protoconid.
LMj identical except that last mentioned accessory
cusp less well-developed. LdP 4 has enlarged
antero-buccal cusp, usually homologous with
protoconid of molars. This cusp less well-
developed in RdP 4 . Isoodon macrourus: J 13743
with RP 3 missing. Position marked by pebbly
knobs of calcified tissue not extending above oral
epithelium. Much more complex pebbly knobs
occur near base of P 4 , surrounding P 3 and
surrounding posterior margin of P 1 . L and RP 3
partially resorbed at points around crown. Eroded
pit in anterior root of LP 4 . LI 1 missing although
reason not clear. J8765 has four upper right
premolars. Extra tooth apparently between RP 1
and RP 3 . J21908 has no teeth posterior to R and
LC 1 , no right upper incisors, rudiments only of two
left upper incisors, and no lower teeth. Gum lines
irregular and all post-canine alveoli (if they existed)
filled with bone.
PHALANGERIDAE
Trichosurus vulpecula: Variation in occurrence of
small teeth between C 1 and P 4 and between I t and
P 4 (Kingsmill 1 962). P 1 variably present (case no.
378). ‘Premilk’ teeth in association with I 3 in
sectioned material (Woodward 1896). J23083 isol-
ated left maxilla without normal P 1 . J23070
isolated right maxilla with abnormal tooth in RdP 4
position. Tooth much larger than normal dP 4 and
has unusual lingual cusp. Tooth with unclear
morphology projecting buccally. J23080 has RP 4
erupting out of alignment antero-buccal to M 1 .
Phalanger spp.: P, orientalis with LI 3 imperfectly
bifid, crown almost completely divided, but root
single (case no. 368). Individuals of P. orientalis:
four upper premolars on one side; one premolar
absent; two teeth occur in place of P 1 ; and tooth
present between normal P 4 and P 3 (case no.
372-275). P. macula tus lacking L and RI 3 (case no.
369). ‘Premilk’ teeth in incisor and canine region
sectioned specimens of Phalanger sp. (Woodward
1896).
PETAURIDAE
Pseudocheirus spp.: P.forbesi has no I 3 and no P 1
(case no. 37 1 ). In present study, P. peregrinus skulls
256
MEMOIRS OF THE QUEENSLAND MUSEUM
(including 23 dentaries) from caves in Western
Australia show following abnormalities and vari-
ations: J23076 has incipiently two-rooted RP 1 ;
J23078, left maxillary fragment, shows same P 1
condition; J23075 has four left upper premolars,
extra premolar either first or second in row, both
being simple peg-like teeth. Teeth posterior to I x
and anterior to P 4 lost but alveoli vary in number
from one (J23072) to two (J23074) to three
(J23073). J 1 1427 has small calcified tooth adher-
ring to postero-buccal corner of RP 4 . Long pos-
terior root present on this tooth and appears that
anterior root broken off. Tooth probably dP 4 , not
previously recorded in this group because either
lost very early in development or not normally
developed to stage of calcified crown. Schoinobates
volans: J22083 has tiny calcified tooth adherring to
antero-lingual tip of RMi . No roots apparent. This
rudiment may represent dP 4 , previously unre-
corded in genus. Petaurus sp.: In 25 skulls, two
show variations in number of post-^-pre^ teeth
(case no. 380). One has four on left side. Hemi-
belideus leadbeateri: J9294 has slightly abnormal L
and RM 4 . Entoconids and postero-lingual corners
of teeth reduced.
PHASCOLARCTIDAE
Phascolarctos cinereus: In sectioned specimen,
small calcified incisor present in front of I x and
another uncalcified incisor present behind
Toothgermsfor lower canine, two lower premolars,
and additional upper premolar present; none attain
functional maturity (Woodward 1896). Rudimen-
tary dP4 (Thomas 1887b). Tooth rudiments noted
by Woodward and Thomas probably best regarded
as uncommon observations of normally occurring
structures. J 1 0023 has two incisors in place of RI 3 .
Both resemble normal LI 1 . J8811 has very tiny
hypocones on L and RM 1-4 . Size of protoconule
varies in this species, being absent to miniscule in R
and LM 1-3 (e.g. J8811) to conspicuous in same
teeth (e.g. J 10023). J 13278 shows two tiny calcified
teeth in shallow sockets on right lower side between
erupting RI : and RP 4 . First of these immediately
posterior to . Second adpressed to antero-lingual
corner of P 4 crown. On left side, same two teeth
present but anterior one lost, perhaps during
preparation of specimen. J7209 has no L or RM 4
but has L and RM 4 . J5749 has mildly twisted L and
R dentary, resulting in malocclusion and abnormal
resting position of incisors.
MACROPODIDAE
Supernumerary Teeth
Incisors: Macropus giganteus J23087 small
tooth anterior to LI 3 .
Premolars: Macropus giganteus J23089 RP 5
erupting beneath RP 4 ; J23108 LP x present anterior
to P 4 ; J23105 RP X present anterior to P 4 .
Molars: Macropus giganteus J23110, J23151,
J23135, J23150, J23137, J23120, J23125, J23083,
J23085 L and RM 5 ; J23107 L and RM 5 and LM 5 ;
J23128 L and RM 5 and ?LM 6 ; J23109, J23129,
J23130 LM 5 ; J23140, J23134 RM 5 ; Macropus
robustus J23122 L and RM 5 ; J23117 RM 5 . Meg-
aleia rufa J23091 supernumerary between RM 3 and
RM 4 ; J23086, J23084, J23152 LM 5 ; M. sp. J231 15,
J23147 RM 5 ; J23136 LM 5 ; Potorous tridactylus
WAM 70.7.242 LM S ; Bettongia lesueur WAM
68.10.2 M s .
Supernumerary molars in Peradorcas concinnus
common (Tate 1948, Troughton 1967). Macropus
spp.: Supernumerary molars, premolars, presence
of rudimentary canines, and absence of P4 (Kirk-
patrick 1965). Kirkpatrick (1965) notes frequency
of some abnormalities (e.g. presence of paired M5
on maxillae) to be as high as seven per cent in one
species. Bettongia spp.: M5 in B. penicillata (case
no. 390) and B. lesueur (Waterhouse 1846, Thomas
1888). L and RM 5 in two other specimens B.
lesueur and L and RM 5 and L and RM 5 in B.
gaimardi (as B. cuniculus) (Thomas 1888).
Missing Teeth
Macropus giganteus J23 1 14 L and RM 4 missing;
J23119 L and RM 4 missing; J23133 L and RM 4
missing; J23126 all lower molars missing R side
only; J23134 RM 4 missing; M. robustus J23117 L
and RM 4 missing; M. sp. J23123 RM 4 missing;
J23144RM* missing; J23145RM 4 missing; J23149
RM 4 missing.
Bettongia spp.: B. penicillata molar formula was
M 1 7 5 (Bateson 1894). LM 4 missing (case no. 389).
M 4 \ . . is itself often aborted in Bettongia, there
being then only three molars’ (Thomas 1888, p.
105).
Abnormal Tooth Morphology
Incisors: tCfacropus giganteus J23092 L and RL
show two pronounced lingual grooves and crowns
deflected buccally; J23087 RI 3 has large buccal
projection from walls of crown adjacent to incisive
groove; J23090, J23153, J23111 RI 3 distorted
shape; J231 12 L and RI 3 shape abnormal.
Molars: Macropus giganteus J23083 LM 5
horseshoe-shaped, RM 5 similar but more complex;
J23085 LM 5 horseshore-shaped but RM 5 mol-
ariform; J23139 RM 4 peg-shaped, LM 4 mildly
abnormal; J23109 LM 5 submolariform with one
loph and one cusp; J23131 possible odontoma in
crypt buccal to LM ; M. robustus J231 17 prominent
257
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
mesostyles on L and RM 3 ; Megaleia rufa J23088
RM 2 with one major transverse loph, RM 4 may
also be peg-shaped (tooth gone); J23091 abnormal
supernumerary tooth between RM 3 and RM 4 ,
three-rooted and tricuspid; J23084 LM 5 metaloph
appears complexly folded; J23068 LM 5 simple
conical cusp surrounded by cingulum; Macro-
podid, indet., F4713 LM 4 lacks clear homologue of
hypolophid.
In macropodids, abnormally shaped teeth re-
latively more common than in other marsupial
families. Abnormalities in shape also different from
those found in other groups in that divided cusps
and crowns, fused crowns, and buccally com-
pressed crowns extremely rare.
Normal morphological variation in premolars of
several modern and fossil species of large macro-
podids described (Bartholomai 1973, 1974), with
view to interpreting variation in fossil species.
Variations in Eruption and Occlusion
Incisors: Macropus giganteus J23087 RI 3 on
premaxilla-maxilla boundary; J23092 RI 3 pos-
itioned near maxilla boundary; J23 104 skull distor-
ted and RI, occludes with LI 3 .
Premolars: Macropus fuliginosus WAM M6956
R and LP 4 erupting near buccal wall of maxilla.
Molars: Macropus giganteus ill 132 L and RM 4
erupted abnormally high in tooth row; J23113
RM 3 erupted abnormally high; J23126 all upper R
molars erupted abnormally, lower molars missing.
Normally non-erupting or Ephemeral Teeth
Incisors and possibly Canines: Macropus irma
WAM M8127 (basicranial length 44-3 mm) two
tooth rudiments, one antero-buccal to unerupted
I 1 , other on premaxilla-maxilla boundary; M.
robustus WAM M6976 (51-4 mm) one tooth
rudiment on premaxilla-maxilla boundary; WAM
M6137 (62-9 mm) no rudimentary teeth, but socket
only just resorbed; M. fuliginosus WAM M6588
(50-4 mm) shallow socket present on premaxilla-
maxilla boundary; M. giganteus J23087 (adult)
spicule-like tooth immediately anterior to LI 3 .
These teeth undoubtedly normal in developing
dentitions of many species and no examination of
sectioned macropodid material fails to mention
them, although not commonly observed in gross
skeletal preparations.
Many cases cited of normally non-erupting teeth
in macropodids (e.g. Flower 1868, Woodward
1893, Hopewell-Smith and Tims 1911, Tate 1947a,
Johnson 1964, Berkovitz 1968c, Bartholomai
1973). Incisors of macropodids normally have
deciduous predecessors that resorb without erupt-
ing (Kirkpatrick 1969). Normally non-erupting
canine develops ontogenetically (Kirkpatrick
1969).
DIDELPHIDAE
Bensley (1906) presented comprehensive anal-
ysis of normal and abnormal variations in stylar
cusp morphology in species of Monodelphis (as
Peramys), Marmosa, Caluromys, Metachirus,
Chironectes, and Didelphis. Didelphis marsupialis:
Six right upper incisors (case no. 363). Four right
lower incisors (case no. 364). Upper incisor missing
from left and right sides (case no. 365). One out of
79 had no R or LM 4 (case no. 388). RM 4 larger
than LM 4 (also case no. 388).
STAGODONTIDAE
Didelphodon padanicus: Type specimen (dentary
fragment) of this Cretaceous didelphid may have
had four premolars (Clemens 1966).
CAENOLESTIDAE
Garzonia patagonica: Specimen of Tertiary Gar-
zonia may have supernumerary number of ante-
molar teeth (Sinclair 1906, Ride 1962, Ziegler
1971).
VOMBATIDAE
Extra calcified (but rudimentary) teeth reported
in vombatids (Owen 1840-45, Rose 1893).
EUTHERIANS
Abnormal dental developments in human teeth
have received considerable attention. The most
important general treatise on the subject is Stones,
Farmer and Lawton (1966). Several other papers
(not noted in Stones et. al.) dealing with general
dental abnormalities are: Black (1902), Kraus,
Jordon and Pruzansky (1966); to papers dealing
with specific abnormalities such as shovel-shaped
incisors, Carbonell (1963); double-rooted lower
canines, Alexandersen (1963); carabelli’s tubercle,
Meredith and Hixon (1954); abnormal cusp de-
velopment in addition to carabelli’s tubercle,
Kallay (1966); dens in dente, Swanson and
McCarthy (1947); and to papers dealing with the
genetic nature of dental abnormalities such as
Brothwell, Carbonell and Goose (1963), and
Hopewell-Smith (1913).
Minor dental abnormalities in some placentals
are documented, for example by Allow (1971),
Bateson (1894), Berkovitz (1968), Berkovitz and
258
MEMOIRS OF THE QUEENSLAND MUSEUM
Musgrave (1971), Chasson (1955), Churcher
(1959), Fish and Whitaker (1971), Forsten (1973),
Frisch (1963), Garn and Lewis (1963), Hooper
(1955), Jones (1960), Hooper (1955), Jones (1960),
Mech, Franzel, Karns and Kuehn (1970), Miller
and Tessier (1971), Peterson and Fenton (1970),
Schitoskey (1971), Spinage (1971), Van Valen
(1966, 1964), Wolfe and Layne (1968), Zakrzewski
(1969), Ziegler (1971).
Several studies have been made using dental
abnormalities in eutherians to interpret factors
controlling tooth development. These include Ber-
kovitz (1969), Butler (1963), Gaunt (1967), Grune-
burg (1951, 1965), Hitchin (1966), Johnson (1952),
Kurten (1955, 1957, 1963, 1967), Stockard et. al.
( 1 94 1 ), Van Valen ( 1 962, 1 970) and Wallace (1968).
DISCUSSION
Supernumerary Teeth
Premolar Number: Diversity of position in
abnormally occurring teeth noted in the present
study indicates that some interpretations attempt-
ing to clarify the maruspial premolar dental
formula are probably unjustified. For example,
Thomas (1887) concludes that appearance of a
premolar between the first and third premolars
represents an atavistic reappearance of a lost
marsupial second premolar. This view is not held
by Owen (1840—5) or by Ziegler (1971). Ziegler
concludes (p. 240) that ‘The premolar position
vacant in all marsupials is . . . most logically
homologized with that of the retained first milk
premolar of placentals . . . accordingly, the first
three metatherian post canines are . . . designated
the second, third and fourth premolars.’ Owen (op.
cit.) believes that premolar teeth are normally lost
from the front of the premolar row. However,
Bateson (1894 p. 249) after an examination of
dental variations in marsupials in general con-
cludes that \ . . the system elaborated by Thomas
breaks down; not because there is any other system
which can claim to supersede it, but because the
phenomena are not capable of this kind of
treatment’. Considering cases noted in the present
study, it seems that extra premolars may occur at
almost any position in the tooth row, as well as
anterior and posterior to the first premolar. In
addition, recent ontogenetic studies by Archer
(1974), Berkovitz (1968) and others have not
supported the suggestion of Woodward (1896) that
there is clear evidence for a suppressed premolar
position between PI and P3 in marsupials, nor the
view held by Owen ( 1 840-45) or Ziegler (1971) that
a similar premolar position has been lost in
marsupials anterior to PI.
P5: Development of P5 in dasyurids and macro-
podids invariably results in a premolariform tooth,
clearly indicating that potential for premolar
production exists posterior to P4. Production of P5
presumably occurs later than P4 since it erupts later
than and posterior to P4. Archer (1974) de-
monstrates that even before P4 is calcified in
Antechinus , the dental lamina connections between
it and adjacent teeth are already breaking down.
Prior to normal breakdown of dental lamina, this
tissue is continuous posterior to the canine. For this
reason, the extra premolar probably could not
result simply from an abnormal posterior extension
of the premolar part of the dental lamina. It
appears that the P5 tooth family position is
established well before the tooth actually develops,
posterior to the P4 position and anterior to the dP4
position on the continuous band of dental lamina.
M5: Molars sometimes occur posterior to M4 in
macropodids but rarely in other marsupials. How-
ever, one case noted above of a Dasyurus maculatus
specimen with L and RM 5 and LM S is of this kind.
It differs in that L and RM 4 resemble a normal L
and RM 3 . This specimen was, in part, the basis for
Bateson’s (1894) formulation of the concept of
homoeosis in meristic series. This concept is that
particular teeth in a series may vary in shape, and
come to resemble aspects of adjacent teeth. In
macropodids with more than four molars, homo-
eosis is not clearly evident because the normal
macropodid M4 closely resembles the normal M3,
and homoeotic variation in M4 would not be as
obvious as in dasyurids.
M5 probably develops as an extra tooth family
position at the posterior end of the dental lamina
(Kirkpatrick 1969). In Peradorcas , production of
additional molars is the normal condition (see Tate
1948). Obstructions to addition of teeth at the
posterior end of the tooth row result from lack of
space. This crowding may be, in part, responsible
for abnormal shape of many supernumerary teeth.
Butler (1956), Gruneberg (1937), and Lefkowitz,
Bodecker and Mardfin (1953) have suggested that
distortion of tooth germs can produce abnormally
shaped teeth. Sofair, Bailit and MacLean (1971)
and Stein (1943) note that this is most likely to be
the case with posterior teeth of a particular tooth
series. ‘Normality’ of supernumerary molars in
macropodids may be due to forward progression of
molars and consequent relative freedom from
crowding.
Probable Atavisms
In some taxa with a dental number reduced
below that possessed by close relatives, teeth
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
259
occasionally occur in the position of the ‘missing’
tooth family. Such occurrences appear to be rather
regular and should be regarded most probably as
atavisms. These would include appearances of
teeth in some dasyurids (e.g. Dasyurus ) at the P4
position, and in some peramelids (e.g. Echymipera)
at the I 5 position.
Eutherians
Berkovitz(1969), Johnson (1969), Kurten (1963)
and others have presented interpretations regard-
ing supernumerary teeth in various eutherians.
Berkovitz (1969) demonstrates the existence of dl 4
in a eutherian carnivore. He suggests this is
evidence for the presence of four incisors in
primitive eutherians. Johnson (1969) notes the
appearance of M 4 in a murid rodent and considers
that this may be the homologue of the normal
eutherian M 3 and that in murid rodents the so-
called M 1 is actually a persistant dP 4 . Kurten
(1963) believes that in one lineage of felids, so-
called supernumerary molars at the rear of the
dentition represent an atavisitic reappearance of a
molar previously lost through evolution.
Although many Mesozoic mammals had more
than seven molariform teeth (e.g. Peramus and
Amphitherium ), there is no particular reason for
believing that the abnormal marsupial M5 noted in
the present study is an atavistic reappearance of a
lost tooth. More probably, these teeth are simply
the result of abnormal activation of a potential
tooth-producing structure, the posterior end of the
dental lamina.
Missing Teeth
Missing teeth, partial anodontia, or hypodontia
noted in the present study occur mostly in the
premolar region and only rarely in the molar
region. Some instances are presumably due to
trauma, others perhaps to disease, while others
seem likely to be caused by genetic defects. Stones,
Desmond and Lawton (1966) note that anodontia
in humans is frequently due to a gene mutation in
the X chromosome. Brekhus, Oliver and Montelius
(1944) note that there are often clear correlations
between tooth absences. For example, loss of Ml is
usually correlated with absence of other molars. In
the only instances of non-traumatic molar loss
noted in the present study, there were other teeth
missing including premolars as well as molars.
However, the great majority of cases of premolar
loss do not involve molar loss.
Abnormal Tooth Morphology
Divided Teeth: Divided teeth were found in the
present study only among ante-molar teeth. Bate-
son (1894) notes that when teeth are divided, the
plane of division is usually transverse to the long
axis of the tooth row. This was invariable in
examples considered here. A tendency for the
divided tooth to be a RP 3 among dasyurids is
present. There is also evidence for correlation of
divided teeth and supernumerary premolars among
dasyurids. In Dasyurus geoffroii (WAM M4464) an
extra premolar occurs on the upper right side and
the RP 3 has a divided crown. Division of single
tooth germs in various stages of development may
be one way in which supernumerary teeth are
produced.
Concepts of tooth development, as discussed by
Butler (1956), do not provide a mechanism for
actually dividing established cusps. Rather, a
divided crown tip could develop from two centres
of suppressed mitosis in the developing tooth germ.
Fused Teeth: Fused, geminated or connated
teeth were found in the present study only among
premolars. Fusion may involve only roots or
anything up to and including the whole crown.
Hitchin and Morris (1966) suggest that actual teeth
do not become fused by any other means than by
cementum after tooth formation is complete. They
present reasons (p. 575) why fusion of tooth germs
is unlikely to occur after epithelial contact between
teeth is broken and follicles develop around each
germ. They suggest (p. 583) that \ . . primary
developmental abnormality in connation is per-
sistence of the dental lamina between the teeth
germs.’ However, it seems equally plausible that
physical trauma could result in ruptured follicles
and subsequent fusion of previously separated
portions of epithelium. Some fused teeth noted in
the present study are also abnormally shaped.
Combinations of abnormal features might result
from fusion of previously ruptured and adjacent
tooth germs. These developmental accidents cer-
tainly have no evolutionary significance.
Other Abnormal Morphology: Mis-shapen
molars noted in the present study are also often
supernumerary teeth. In some cases they may be
sub-molariform with part of the tooth, such as a
protoloph, resembling a serially homologous struc-
ture in an anterior molar. M5 is commonly a tooth
of this sort. Horseshoe-shaped and peg-like M5s
are also known. Supernumerary molars occurring
between other molar teeth are generally not
molariform.
Some grossly abnormal molars suggest struc-
turally ancestral cusp patterns. Examples of this
may be cited among abnormal macropodid molars
which show isolated cusps rather than crests.
However, an equivalent number of abnormal
260
MEMOIRS OF THE QUEENSLAND MUSEUM
crowns do not suggest structurally ancestral
shapes, such as the horseshoe-shaped or peg-like
supernumerary molars. Stein (1934, p. 1817) notes
that \ . . from a reasonably large collection of
human third molars, different specimens could be
selected and arranged in such order as to prove
almost any theory of the evolution of the human
dentition’. Abnormally shaped non-
supernumerary molars are also noted above.
Compression is a common abnormality of this
kind. This was also noted by Archer (1971) in
thylacine teeth. Probably this results from com-
pression of the tooth germ follicle. Gruneberg
(1937) and Lefkowitz, Bodecker and Mardfin
(1953) have stressed the importance of the follicle
in production of normally-shaped teeth.
Osborn (1902) notes that increasing brachycephaly
of rhinoceratids is correlated with antero-pos-
teriorly shorter but wider molars. Butler (1956)
suggests the possibility that this is the result of
antero-posterior compression of the tooth germ
follicles. In most of the examples given in the
present study, antero-posterior molar compression
is correlated with brachycephaly, shortening of the
molar row, and sometimes malocclusion.
Number of stylar cusps present in dasyurid
molars is clearly variable within species, although
these variants are generally uncommon and their
significance is unclear. Except in Neophascogale
and Phascolosorex, there are normally only two
conspicuous stylar cusps, st.B and st.D. St. A is not
usually distinguishable from st.B. Sometimes, a
stylar cusp (or pair) is developed between st.B and
st.D, and it has been called here st.C. Stylar cusps
may appear posterior to st.D. Some didelphids
(considered structurally ancestral to dasyurids)
have five stylar cusps. Other didelphids have
practically no stylar cusps. Bensley (1906) has
demonstrated considerable variation in didelphid
stylar cusps. He concludes (pp. 12-13) that al-
though these \ . . relatively small and subsidiary
structures in the molar crown are certain to exhibit
signs of variation, they are surprisingly constant in
their relations . . . they show throughout the family
indications of a general type . . .’. The same can be
concluded for the stylar cusp area of dasyurids.
Despite intra-specific variation and even inter-
specific modification, the dasyurid basic pattern
described above is clear. However, more infor-
mation about the Tertiary record of dasyurid stylar
cusp development is required before the struc-
turally ancestral dasyurid condition can be defined.
Eruption and Occlusion Abnormalities
Malocclusion is rare among marsupials but
several instances have been described. An example
of exaggerated overbite noted above is similar to
examples presented by Stockard et al. (1941)
among dogs. They suggest that upper and lower
jaw development is under separate genetic control
because, for example, in a cross between a Saluki
Dog and a Bassethound, the hybrid had a skull of a
Saluki-type in length and a dentary of a
Bassethound-type in length, resulting in gross
malocclusion with the C x biting behind the C 1 .
Another case of malocclusion noted in the
present study, involves Dasyuroides byrnei. A large
series of these animals were trapped and lodged in
the Queensland Museum’s collection (e.g. J 10226).
Some bred in captivity (Mack 1961) through
several generations. Comparison of 30 skulls of
animals bred in captivity and the 9 animals caught
in the wild indicates that problems of malocclusion
and missing teeth occurred only among animals
bred in captivity. In addition there was occurrence
of abnormally compressed molars and brachy-
cephaly in some animals bred in captivity. Reasons
for this are not clear but factors such as diet and
inbreeding are likely to have been involved.
Abnormalities and Diseases
Examples of morphological variation, tooth
loss, and bony accretion around the tooth roots
noted in the present study may be the result of
disease. In other instances, some teeth show
abnormal dentine accretions around roots, and one
specimen may possibly represent a composite
odontoma. The difference between composite
odontomas and congenital teratomas such as
ovarian dermoid cysts (Stones, Farmer and Law-
ton 1966) seems to be one largely of position. The
case noted in the present study involves an
abnormal M 4 with apparently several surrounding
and related calcified structures. There is, however,
no evidence that these calcified structures and M 4
were fused.
Ephemeral Teeth
Ephemeral teeth observed in sectioned material
or carefully prepared juvenile specimens, do not
really represent abnormal dental developments in
the sense outlined in the introduction. However,
they have been considered here either because they
are not commonly observed or because they
represent teeth previously unobserved.
Observations of the occurrence of small calcified
incisors are reported in this paper and by others
(e.g. Berkovitz 1967, Woodward 1896), parti-
cularly among macropodids. They rarely persist in
adults. These teeth probably represent true de-
ciduous teeth related to the incisors (as suggested
by Kirkpatrick 1969). Clear embryological evid-
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
261
ence for them is known (e.g. Archer 1974, Ber-
ko vitz 1968c, Rose 1892, Woodward 1896) for
many marsupial groups and they are generally
assumed to be rudimentary deciduous teeth. This
may not be true for rudimentary canines in
macropodids. These may represent rudimentary
non-deciduous teeth, which in other macropodids
(e.g. potoroines) are parts of the functional adult
dentition.
It has been noted that some ephemeral teeth
occurring in the position of a ‘missing’ tooth family
in some closely related forms should be interpreted
as atavisms. Tate (1947) notes a case of a dental
rudiment in the P 4 region of a specimen of Dasyurus
hallucatus and considers this a dP 4 . However, its
actual identity is doubtful, since it could be either a
rudimentary dP 4 or a rudimentary P 4 . Reduction of
size in these teeth is not necessarily correlated. For
example, it has been noted above that while the
Phascolarctos P4 is large, dP4 is a mere rudiment
(Thomas 1887). In macropodids, while dP4 is as
large as a molar, P4 may be (in some species) a
rudiment. This is clearly so in thylacinids (Flower
1868), adding support to the idea (Archer 1974,
Berko vitz 1966) that the two teeth do not belong to
the same tooth family and hence development of
one may not depend on development of the other.
Bateson (1894) concludes that meristic variation
in teeth is discontinuous, and that a structure was
either a tooth or not a tooth. However, ephemeral
teeth in the incisor and canine regions clearly
present exceptions to this concept. These have been
found in all stages of development from mere
lingual growths of dental lamina to calcified and
even erupted teeth. Commonly, even after
calcification, the teeth are resorbed. Sometimes
they persist into the adult dentition as do the small
canines in some species of macropodine and most
potoroine macropodids.
Dental Abnormalities as Indicators
of Pattern in Development
Value of dental abnormalities in general in
understanding factors controlling development of
teeth is doubtful. There have been many attempts
to interpret the nature of controlling factors by
analysis of the kinds and incidence of dental
abnormalities.
Butler (1967) concludes, after noting studies of
abnormal conditions of human jaws, that distur-
bances in migration of mesenchyme from the
neural crest may account for abnormal develop-
ments of teeth and their supporting bones. In cases
where the tongue is doubled, a median series of
teeth may develop between the two tongues.
Similarly in unilateral hypertrophy of the face,
teeth are enlarged on the affected side. It is
therefore interesting to consider the possible
significance of unilateral and bilateral dental
abnormalities. Bateson (1894) finds that dental
abnormalities sometimes occur simultaneously on
both sides of the head but rarely occur simul-
taneously in upper and lower dentitions. Neverthe-
less, he cites examples of extra molars on both
upper sides only (e.g. case nos. 178, 179 and 194),
on both lower sides only (e.g. case nos. 171 and
251), on upper and lower right sides only (e.g. case
nos. 190 and 196), on both lower sides and only one
upper side (e.g. case nos. 182 and 385), on both
upper sides and only one lower side (e.g. case nos.
166 and 167), and on both upper and both lower
sides (e.g. case 1 89). Similar examples of correlated
variations are given for other tooth series and for
missing teeth. Apparently all combinations of this
sort are possible. In specimens noted in the present
study, particularly among macropodids, L and
RM 5 may be grossly different (e.g. J23085) or
mirror images of one another (e.g. J23083). The
case given of a grossly abnormal L and RMj in
Isoodon obesulus shows that complex bilateral
abnormalities may occur, without abnormal oc-
clusal counterparts.
Butler (1961, p. 122) considers that ‘If the
mutation of teeth (upper, lower and adjacent) were
fortuitous, variability of pattern would result so
frequently in malocclusion that natural selection
would operate to reduce that variability to a
minimum. Yet molar teeth show a high degree of
individual variation.’ Accordingly he proposes that
genes controlling tooth shape (and presumably
number) may have a dual or pleiotropic effect,
producing mirror image structures on surfaces of
teeth which contact. This idea relates to the concept
of dental morphogenetic fields (Butler 1937, and
discussed by Butler 1961, Van Valen 1962, 1970,
Wallace 1968, et. al.). The concept is based on the
belief of the equipotential nature of all tooth germs
of a particular species (suggested by Bolk 1922).
This potential is modified by other factors such as
heredity, position in the tooth row, chemical
imbalances, disease, trauma, and available room.
Many of these factors have been analyzed (e.g.
Stones, Farmer and Lawton 1966). However, the
way in which position in the tooth row controls
tooth shape is not clearly understood. Butler
(1967), notes that Remane (1926) figures a speci-
men of Colobus whose first and second permanent
molars are dwarfed and premolariform. Stein
(1934) notes a specimen of a human M 3 which
corresponds cusp for cusp, ridge for ridge, and
groove for groove with a normal P 2 . These
examples suggest that position in the tooth row is
262
MEMOIRS OF THE QUEENSLAND MUSEUM
not always the most important factor in develop-
ment. Butler (1963) regards such variation in dP 3 as
evidence for slight shifts in dental morphogenetic
field at the molar-premolar boundary, and this may
also apply to Remane’s (1926) specimen but not to
Stein’s (1934).
The present study offers no additional examples
among marsupials which bear on the question of
developmental importance of position in the tooth
row. It does provide examples (e.g. bilaterally
symmetric abnormal L and RMQ of abnormal
molars that have not been correlated with abnor-
mal occlusal counterparts. It may be that com-
position of dental morphogenetic fields, if they
exist, is much more complex than has been
visualized (e.g. by Wallace 1968 and Van Valen
1970).
CONCLUSIONS
Abnormal dental developments do not appear to
provide a basis for interpreting position of a
supposedly phylogenetic lost premolar tooth in
marsupials other than P4 of some dasyurids. It has
been concluded elsewhere (Archer 1974) that
ontogeny in some dasyurids similarly fails to reveal
a ‘lost’ premolar position in structurally ancestral
marsupials. There does not appear to be any sound
reason for believing that marsupials phylogeneti-
cally suppressed a particular premolar position.
In some dasyurids with only two premolars on
each side, there is evidence from dental variations
that the P4 position has been suppressed. Such
dental variations should be referred to as atavisms.
Abnormal production of P5 and M5 in mar-
supials suggests that the dental lamina may remain
proliferative at its posterior end even after the
normal complement of tooth families has been
established. This is particularly evident in the
macropodid Peradorcas where continuous pro-
duction of supernumerary molars may occur.
Abnormal crown morphology occurs most com-
monly among premolars. These often exhibit fused
or divided crowns. Fused crowns may result from
damage to adjacent developing teeth. Divided
crowns, because they are sometimes related oc-
clusally to supernumerary teeth, may be one stage
in a process in which supernumerary teeth are
produced. Abnormal variations are also common
on the stylar shelf of molars. Grossly abnormal
molar crowns are uncommon, in marsupials in
general but are more common in supernumerary
macropodid molars, distorted molar crowns may
result from crowded tooth germs. Among dasy-
urids, antero-posteriorly compressed molars are
among the most commonly encountered molar
abnormalities.
Malocclusion and abnormal molar crown mor-
phology appear, among some dasyurids, more
common among inbred than among trapped
individuals, suggesting some abnormalities may
have a genetic basis.
From ontogenetic studies, many ephemeral teeth
developing in canine and incisor position's appear
to represent short-lived milk-teeth (e.g. macro-
podid incisors) or small vestiges of teeth in the
process of phylogenetic reduction (e.g. macro-
podine canines). These are sometimes found in dry
skulls representing very juvenile individuals, if the
skulls are carefully prepared.
Most abnormal dental developments appear to
be of little or no significance in interpreting
phylogeny, while others do appear significant and
suggest that mechanisms determining tooth shape
are complex. Dental morphogenetic fields do not
always ensure that abnormalities occur with oc-
clusal counterparts and the majority of dental
abnormalities occur in one tooth row only. Other
examples do involve occlusal or bilateral counter-
parts, and add support to the concept of dental
morphogenetic fields. The apparent inconsistency
may be resolved if the majority of dental abnormal-
ities lack a genetic basis.
ACKNOWLEDGMENTS
Dr W. D. L. Ride, as my supervisor, con-
structively criticised a draft of this work. Dr A.
Bartholomai and Mr B. Campbell of the Queens-
land Museum also provided helpful criticism.
Various other people helped by bringing my
attention to interesting dental abnormalities in-
cluding Dr D. Merrilees, Mr A. Baynes, and Ms J.
Porter, of the Western Australian Museum, and Dr
T. K. Kirkpatrick of the Queensland Department
of Primary Industries who also allowed me to study
the interesting series of abnormal teeth in his
collection of macropodid skulls. Mr K. Thompson,
University of Western Australia, kindly donated
J23082, the abnormal Isoodon obesulus. Dr H. Van
Deusen, American Museum of Natural History,
Ms J. Covacevich, Queensland Museum, Ms J.
Dixon, National Museum of Victoria, Mr B. J.
Marlow, Australian Museum, Mr R. Warneke,
Fisheries and Wildlife Department, Victoria, Mr P.
Aitken, South Australian Museum, Mr S. Parker,
formerly of the Arid Zone Research Centre, Mr R.
Green, Queen Victoria Museum and Art Gallery,
and Dr E. Hill, British Museum, Natural History,
allowed me to examine specimens in museum
collections. Mr D. Vernon, Queensland Museum,
kindly gave me access to records he has kept of the
breeding population of Dasyuroides byrnei in the
Queensland Museum. Mr A. Easton, Queensland
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYUR1DS
263
Museum, helped with photography.
While this research was carried out, I received a
Fulbright Scholarship, a grant in aid from the
American Explorers’ Club, and a Research Assis-
tantship to Dr W. D. L. Ride, who was in receipt of
a Research Grant from the Australian Research
Grants Committee.
LITERATURE CITED
Alexandersen, V., 1963. Double-rooted human lower
canine teeth. In D. R. Brothwell, (Ed.) ‘Dental
Anthropology’, pp. 235-44, (Pergamon Press: Ox-
ford).
Allo, J., 1971. The dentition of the Maori Dog of New
Zealand. Rec. Auckland (N.Z.) Inst. 8: 29-45.
Archer, M., 1971. A re-evaluation of the Fromm’s
Landing thylacine tooth. Proc. R. Soc. Viet. 84:
229-34.
1974. The development of the cheek-teeth in Antechinus
flavipes (Marsupialia, Dasyuridae). J. R. Soc. W.
Aust. 57: 54-63.
1975. Ningaui, a new genus of tiny dasyurids (Mar-
supialia) and two new species from arid Western
Australia, N. timealeyi and N. ridei. Mem. Qd. Mus.
Bader, R. S., 1954. Variability and evolutionary rate in
the oreodonts. Evolution 9: 1 19-40.
Bartholomai, A., 1971a. Morphology and variation of
the cheek teeth in Macropus giganteus Shaw and
Macropus agilis (Gould). Mem. Qd Mus. 16: 1-18.
1971b. Dasyurus dunmalli , a new species of fossil
marsupial (Dasyuridae) in the upper Cainozoic
deposits of Queensland. Mem. Qd Mus. 16: 19 26.
1973. The genus Protemnodon Owen (Marsupialia:
Macropodidae) in the upper Cainozoic deposits of
Queensland. Mem. Qd Mus. 16: 309-63.
Bateson, W., 1894. ‘Materials for the study of variation
treated with special regard to discontinuity in the
origin of species.’ pp. 1-598 (Macmillan and Co.:
London).
Bensley,, B. A., 1903. On the evolution of the Australian
Marsupialia: with remarks on the relationships of
marsupials in general. Trans. Linn. Soc. Lond. 9:
83-217.
1906. The homologies of the stylar cusps of the
upper molars of the Didelphyidae. Stud. Univ.
Toronto, Biot. Ser. 5: 1-13.
Berkovitz, B. K. B., 1966. Homology of the premolar
teeth in Setonix brachyurus (Macropodidae: Mar-
supialia). Archs oral Biol. 11: 1371-84.
1967. The dentition of a 25-day pouch young specimen
of Didelphis virginiana (Didelphidae: Marsupialia).
Arch oral Biol. 12: 1211-2.
1968a. Some stages in the early development of the
post-incisor dentition of Trichosurus vulpecula (Phal-
angeroidea: Marsupialia). J. Zool., Lond. 154:
403-14.
1968b. Supernumerary deciduous incisors and the
order of eruption of the incisor teeth in the albino
ferret. J. Zool.. Lond. 155: 445-9.
1968c. The early development of the incisor teeth of
Setonix brachyurus (Macropodidae: Marsupialia)
with special reference to the prelacteal teeth. Archs
oral Biol. 13: 171-90,
1969. Supernumerary deciduous incisors in the polecat.
Archs oral Biol. 14: 863.
Berkovitz, B. K. B. and Musgrave, J. H., 1971. A rare
dental abnormality in an adult male orang-utan
(Pongo pygmaeus); bilateral supernumerary maxil-
lary premolars. J. Zool., Lond. 164: 266-8.
Black, G. V., 1902. ‘Descriptive anatomy of the human
teeth’. (5th Ed.). Pp. 1-169. (S. S. White Dental
Manufacturing Company: Philadelphia).
Bolk, L., 1922. Odontological essays. 4. On the relation
between reptilian and mammalian teeth. J. Anat.,
Lond. 56: 136.
Brekhus, P. J., Oliver, C. P., and Montelius, G., 1944.
A study of the pattern and combinations of con-
genitally missing teeth in man. J. dent. Res. 23:
117-31.
Brothwell, D. R., Carbonell, V. M., and Goose, D.
H., 1963. Congenital absence of teeth in human
populations. In D. R. Brothwell (Ed.), ‘Dental
Anthropology’, pp. 179-90. (Pergamon Press: Ox-
ford).
Butler, P. M., 1937. Studies of the mammalian den-
tition. The teeth of Centetes ecaudatus and its allies.
Proc. zool. Soc. Lond. (Series B) 109: 1-36.
1956. The ontogency of molar pattern. Biol. Rev. 31 :
30-70.
1961. Relationships between upper and lower molar
patterns. Int. Colloq. Evol. Lower and Non-
specialised Mammals, Brussels 1: 1 17 26.
1963. Tooth morphology and primate evolution. In D.
R. Brothwell, (Ed.) ‘Dental Anthropology’, pp.
1-13. (Pergamon Press: Oxford).
1967. Dental merism and tooth development. J. dent.
Res. 46, supplement Sept. -Oct. 1967: 845-50.
Carbonell, V. M., 1963. Variations in the frequency of
shovel-shaped incisors in different populations.//! D.
R. Brothwell, (Ed.) ‘Dental Anthropology’, pp.
211-34. (Pergamon Press: Oxford).
Chasson, R. B., 1955. Dental abnormalities of the
Alaskan Fur Seal. J. Mammal. 36: 562-64.
Churcher, C. S., 1959. The specific status of the new
world red fox. J. Mammal. 40: 513-20.
Clemens, W. A. Jr., 1966. Fossil mammals of the type
Lance Formation, Wyoming. Part 2. Marsupialia.
Bull. Dep. Geol. Univ. Calif. 62: 1-122.
Daniel, M. J. and Kershaw, M., 1964. Recent con-
genital anomalies in rdd deer in New Zealand. J.
Mammal, 45: 480-83.
Darwin, C., 1882. ‘The variation of animals and plants
under domestication’. (2nd Ed.) Vol. 1: 1-473. (John
Murray: Lond.).
Fish, P. G. and Whitaker J. O. Jr., 1971. Microtus
pinetorum with grooved incisors. J. Mammal. 52:
827.
Flower, W. H., 1868. On the development and suc-
cession of the teeth in the Marsupialia. Phil. Trans.
157: 631-41.
Forsten, A., 1973. Abnormal enamel morphology in
fossil equid teeth. J. Mammal. 54: 255-8.
Frisch, J. E., 1963. Dental variability in a population of
Gibbons. In D. R. Brothwell, (Ed.) ‘Dental
264
MEMOIRS OF THE QUEENSLAND MUSEUM
Anthropology’, pp. 15-28. (Pergamon Press: Ox-
ford).
Garn, S. M. and Lewis, A. B,, 1963. Phylogenetic and
intraspecific variations in tooth sequence poly-
morphism. In D. R. Brothwell, (Ed.) ‘Dental
Anthropology’, pp. 53-73 (Pergamon Press: Ox-
ford).
Green, R. H., 1967. Notes on the Devil (Sarcophilus
harrisi) and the Quoll (Dasyurus viverrinus) in north-
eastern Tasmania. Rec. Q. Viet. Mus. 27: 1-13.
Gruneberg, H., 1951. The genetics of a tooth defect in
the mouse. Proc. R. Soc. (Series B) 138: 437-51.
1965. Genes and genotypes affecting the teeth of the
mouse. J. Embryol. exp. Morph. 14: 137-59.
Hitchin, A. D., and Morris, L, 1966. Geminated
odontome-connation of the incisors in the dog-its
etiology and ontogeny. J. dent. Res., supplement to
no. 3 45: 137-59.
Hooper, E. T., 1955. Extra teeth in the pygmy mouse
Baiomys musculus. J. Mammal. 36: 298-9.
Hopewell-Smith, A., 1913. ‘An introduction to dental
anatomy and physiology descriptive and applied’,
pp. 1 372. (Lea and Febiger: Philadelphia).
Hopewell-Smith, A. and Tims, H. W. M., 1911. Tooth-
germs in the wallaby Macropus billardieri. Proc. zool.
Soc. Lond. 1911: 926-42.
Johnson, D. H., 1952. The occurrence and significance of
extra molar teeth in rodents. J. Mammal. 33: 70-2.
1964. Mammals of the Arnhem Land Expedition. Pp.
427-517 In. Records of the American- Australian
Scientific Expedition to Arnhem Land, Vol. 4,
(University Press: Melbourne).
Jones, J. K., 1960. Absence of third upper premolar in
Eutamias. J. Mammal. 41: 269.
Kallay, J., 1966. Extra cusp formation in the human
dentition. J. dent. Res. 45: 1381-94.
Kingsmill, E., 1962. An investigation of criteria for
estimating age in the marsupials Trichosurus vul-
pecula Kerr and Perameles nasuta Geolfroy. Aust. J.
Zool. 10: 597-616.
Kirkpatrick, T. H., 1965. Studies of macropodidae in
Queensland. 2. Age estimation in the grey kangaroo,
the red kangaroo, the eastern wallaroo and the red-
necked wallaby, with notes on dental abnormalities.
QdJ. agric. Sci. 22: 301-17.
1969. ‘The dentition of the marsupial family Macropo-
didae with particular reference to tooth development
in the Grey Kangaroo Macropus giganteus Shaw’.
Unpublished thesis submitted to the Department of
Zoology, University of Queensland: 1-128.
Kraus, B, S., Jordan, R. E. and Prunzansky, S., 1966.
Dental abnormalities in the deciduous and per-
manent dentitions of individuals with cleft lip and
palate. J. dent. Res. 45: 1736-46.
Kurten, B., 1955. Contribution to the history of a
mutation during 1,000,000 years. Evolution 9:
107-18.
1957. A case of Darwinian selection in bears. Evolution
11: 412 16.
1963. Return of a lost structure in the evolution of the
felid dentition. Comment, biol. Helsingf. 26: 3-11.
1967. Some quantitative approaches to dental mic-
roevolution. J. dent. Res. supplement to 46: 817 28.
Macintosh, N. W. G., and Mahoney, J. A., 1964. ‘A
4,000 years old thylacine tooth (Dasyuridae) from
Shelter 2’ Appendix 3 (pp. 507-16) In D. J.
Mulvaney, et. al. Archaeological excavation of
Rock Shelter no. 6 Fromm’s Landing, South
Australia. Proc. R. Soc. Viet. 77: 479-516.
Mack, G., 1961. Mammals from south-western
Queensland. Mem. Qd Mus. 13: 213-29.
Mech, D. L., Frenzel, L. D. Jr., Karns, P. D., and
Kuehn, D. W., 1970. Mandibular dental anomalies
in White Tailed deer from Minnesota. J. Mammal.
51: 804-6.
Meredith, H. V., and Hixon, E. H., 1954. Frequency,
size and bilateralism of Carabelli’s tubercle. J. dent.
Res. 33: 435-40.
Miller, F. L., and Tessier, G. D., 1971. Dental
anomalies in Caribou, Rangifer tarandus. J.
Mammal. 52: 164-73.
Mills, J. R. E., 1963. Occlusion and malocclusion of the
teeth of primates. In D, R. Brothwell, (Ed.)
‘Dental Anthropology’, pp. 29 5 1 . (Pergamon Press:
Oxford).
Oringer, M. J., 1948. Anomalies of human dentition.
Oral Surgery Oral Medicine Oral Pathology 1:
1119-30.
Owen, R., 1840-5. Odontography. Ixxiv and 665 pp.
(Hippolyte Balliere: London).
Remane, A., 1926. Eine seltsame Gebissanomalie bei
einem StummelaflFen, zugleich ein Beitrag zur Frage
der Selektionswirkung bei der Gebiss-
differenzierung, Z. Saugetierk. 1: 114-20.
Ride, W. D. L., 1962. On the evolution of Australian
marsupials. Pp. 28 1-306 in G. W. Leeper, (Ed.) The
evolution of living organisms’. (Melb. Univ. Pr.:
Melbourne).
1964. Antechinus rosamondae, a new species of dasyurid
marsupial from the Pilbara District of Western
Australia; with remarks on the classification of
Antechinus. W. Aust. Nat. 9: 58-65.
1970. ‘A guide to the native mammals of Australia.’ xiv
and 249 pp. (Oxford Univ. Pr.: Melbourne).
Rose, C., 1892. Uber die Zahnentwicklung der Beutel-
tiere. Anat. Anz. 7: 639-50 and 693-707.
1893. Uber die Zahnentwicklung von Phascolomys
Wombat. S.B. preuss. Akad. Wiss. 38: 749-55.
Ruston, M. A., 1936. Some dilated composite
odontomes. Dent. Rec. 56: 766-74.
Schitoskey, F. Jr., 1971. Anomalies and pathological
conditions in the skulls of Nutria from southern
Louisiana. Mammalia 35: 31 1-14.
Sinclair, W. J., 1906. Mammalia of the Santa Cruz beds,
Marsupialia. Rep. Princeton Exped. to Patagonia 4:
333-408.
Sofair,J. A., Bailit, H. L., and MacLean, C. J., 1971. A
developmental basis for differential tooth reduction
during hominid evolution. Evolution 25: 509-17.
Spinage, C. A., 1971. Two records of pathological
conditions in the impala (Aepyceros melampus). J.
Zool., Lond. 164: 269-70,
Stein, M. R., 1934. Some variations of the upper third
molar. J. Amer. dent. Ass. 21: 1815-9.
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
265
Stockard, C. R., et. al, 1941. The genetic and
endocrine basis for differences in form and
behaviour. Amer. anat. Mem. 19 : 367-72.
Stones, H. H., Farmer, E. D. and Lawton, F. E., 1966.
‘Stone’s oral and dental diseases.’ (5th Ed.) Pp.
1-1 1 12. (E. and S. Livingstone Ltd.: London).
Sutton, J. B., 1890. ‘Evolution and disease.’ Pp. 1-285.
(Walter Scott: London).
Swanson, W. F., and McCarthy, F. M., 1947. Bilateral
dens in dente. J. dent. Res. 26: 167-71.
Thomas, O., 1887a. On the homologies and succession of
the teeth in the Dasyuridae with an attempt to trace
the history of the evolution of mammalian teeth in
general. Phil. Trans. 178 : 443-62.
1887b. On the milk dentition of the koala. Proc. zool.
Soc. Lond. 128 : 338-9.
1888. ‘Catalogue of the Marsupialia and Monotremata
in the collection of the British Museum (Natural
History).’ xiii and 401 pp. (British Museum (Natural
History): London).
Tate, G. H. H„ 1947. Results of the Archbold
Expeditions. No. 56. On the anatomy and
classification of the Dasyuridae (Marsupialia). Bull.
Am. Mus. nat. Hist. 88: 101-55.
1947a. An example of ‘prelacteal incisors’ in advanced
pouch young of Macropus. J. Mammal. 28 : 399-400.
1948. Results of the Archbold Expeditions. No. 59.
Studies on the anatomy and phylogeny of the
Macropodidae (Marsupialia). Bull. Am. Mus. nat.
Hist. 91 : 233-351.
1951. The banded anteater Myrmecobius Waterhouse
(Marsupialia). Amer. Mus. Novit. 1521 : 18.
Tratman, E. K., 1949. An unrecorded form of the
simplest type of the dilated composite odontome.
Brit. dent. J. 86: 271-5.
Troughton, E. Le G., 1967. Furred animals of
Australia. (9th Ed.) xxxii and 384 pp. (Angus &
Robertson Ltd.: Sydney).
Van Valen, L., 1962. Growth fields in the dentition of
Peromyscus. Evolution 16 : 272-7.
1964. Nature of supernumerary molars of Otocyon. J.
Mammal. 45 : 284-6.
1966. Nearly rooted incisors in an abnormal rat, and
control of tooth growth. Evolution 20: 428-30.
1970. An analysis of developmental fields. Devi. Biol.
23: 456-77.
Waddington, C. H., 1966. Fields and gradients. Symp.
Soc. Develop. Biol. 25 : 105 24.
Wallace, J. T., 1968. Analysis of dental variation in
wild-caught California house-mice. Amer. Midi.
Nat. 80 : 360-80.
Waterhouse, G. R., 1846. ‘A natural history of the
Mammalia’. Vol. 1. Marsupiata, or pouched
animals. Pp. 1-553. (Hippolyte Bailliere: London).
Wilson, J. T., and Hill, J. P., 1896. Observations upon
the development and succession of the teeth in
Perameles ; together with a contribution to the
discussion of the homologies of the teeth in
marsupial families. Quart. J. micr. Sci. 39 : 427-588.
Wolfe, J. L., and Layne, J. N., 1968. Variations in
dental structures of the Florida mouse, Peromyscus
floridanus. Amer. Mus. Novit. 2351 : 1-7.
Woodward, M. F., 1893. Contributions to the study of
mammalian dentition. Pt. 1. On the development of
the teeth of the Macropodidae. Proc. zool. Soc.,
Lond. 1893 : 450-73.
1896. On the teeth of the Marsupialia with special
reference to the premilk-dentition, Anat. Anz. 12 :
281-91.
Zakrzewski, R. J., 1969. Dental abnormality in the
genus Castor. J. Mammal. 50 : 652-3.
Ziegler, A. C., 1971 . A theory of the evolution of therian
dental formulas and replacement patterns. Quart.
Rev. Biol. 46 : 226-49.
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 30
A. Antechinus flavipes (WAM M6785) with extra right lower
premolar anterior to .
B. Sarcophilus harrisii (WAM 71.10.209) with abnormal tooth
posterior to LP 1 and otherwise missing posterior left upper
teeth. Specimen from cave deposit.
C. Phascogale tapoatafa (WAM M7453) with three-rooted
partially divided LP 3 .
D. Dasyurus maculatus (J 16744) with abnormal LM 4 , possibly
part of composite dental odontoma.
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
Plate 30
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 31
A. Sminthopsis sp. (b) (J5459) with divided entoconid on RM 3 .
B. Sminthopsis sp. (b) (J 5173) with bifid stylar cusp C on RM 1 .
C. Sminthopsis granulipes (WAM M6062) with stylar cusp C on
RM 1 or split stylar cusp D.
D. Neophascogale sp. (AMNH 109524) with divided stylar cusp
C and possibly also divided stylar cusp D on RM 1 .
E. Dasyuroides byrnei (specimens in Queensland Museum). El,
captured live, presents normal skull form. E2, inbred in
captivity, shows extreme brachycephaly and related
distortions of ‘wild’ condition.
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
Plate 31
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 32
A. Isoodon obesulus (J 23082) with grossly abnormal L and
RMj.
B. Isoodon macrourus (J 21908) showing almost total anodontia.
C. Trichosurus vulpecula (1, J 23071; 2, J 23070). Cl shows
normal RdP 4 . C2 shows abnormal ?RdP 4 erupting above this
tooth through side of maxilla.
Plate 32
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 33
A. Pseudocheirus peregrinus (J 11427) with calcified ?RdP 4
adhering to buccal side of RP 4 .
B. Phascolarctos cinereus (1, J 8811; 2, J 10023). B1 shows
reduced hypocones on RM 1-4 . B2 shows extra right incisor
and unreduced hypocones on RM 1 ^,
C. Macropus irma (WAM M8127) showing calcified 7LC 1
with maxillary-premaxillary crypt and calcified ?L and Rdl 1
anterior to unerupted L and RI 1 .
ARCHER: ABNORMAL DENTAL DEVELOPMENT IN DASYURIDS
Plate 33
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 34
A. Macropus robustus (1, WAM M6976; 2, WAM M6137). A1
shows 7LC 1 in maxillary-premaxillary crypt formed by
anterior extension of maxillary bone. A2 shows older
individual in which anterior projection of maxillary bone has
been resorbed (former position indicated by arrow) and 7RC 1
lost.
B. Macropus giganteus (J 23092) with abnormal L and RI 2 and
RP erupting in abnormal position.
C. Macropus giganteus (J 23087) with RP abnormal in structure
and position, and supernumerary incisiform tooth anterior to
LP.
MEMOIRS OF THE QUEENSLAND MUSEUM
Plate 35
A. Macropus giganteus (J 23083) with abnormal L and RM 5 .
B. Megaleia rufa (J 23091) with abnormal supernumerary tooth
between RM 3 and RM 4 .
C. Megaleia rufa (J 23088) with supernumerary abnormal tooth
posterior to RM 1 and alveolus for another posterior to ?RM 3 .
D. Megaleia rufa (J 23084) showing abnormal LM 5 .
E. Megaleia rufa (J 23086) showing abnormal conical LM 5 .
F. Macropus giganteus (J 23085) showing abnormal but
bilaterally symmetric L and RM 5 .
G. Macropus giganteus (J 23089) showing RP 5 replacing RP 4
anterior to RM 1 .
H. Macropodid (F 4713) from Pliocene Chinchilla Sands
Formation, showing abnormally shaped LM 4 or M 5 .
ARCHER: ABNORMAL DENTAL DEVELOPMENT TN DASYURIDS
Plate 35
Mem. QdMus . 17(2): 267-92. [1975]
THE GENUS DIG ASTER (MEGASCOLECIDAE: OLIGOCHAETA) IN QUEENSLAND
B. G. M. Jamieson
Department of Zoology, University of Queensland
ABSTRACT
Digaster is here emended to exclude Perissogaster excavata and P. nemoralis but to retain P.
queenslandica. The redefined genus is limited, as previously, to the montane coastal province of
the Eastern Subregion of Australia. It consists of 17 species: 13 in southern Queensland, 2 in New
South Wales and 2 species shared between the two states. Evidence given suggests that the genus
has originated locally from holonephric species with a pair of combined male and prostatic pores
on XVIII, or from meronephric monogastric descendants of these, and that origin, or at least
major speciation, of the genus has been relatively recent. It is deduced that the digastric or
trigastric condition of the genus is derived from the monogastric condition and that this
multiplication of gizzards has occurred in Australia independently of other regions. A new generic
definition and a key to all species are given together with descriptions of 5 new species, 2 new
subspecies and of new material of 5 previously known species.
The genus Digaster was erected by Perrier (1872)
for a single species, D. lumbricoides , from the vicin-
ity of Port Macquarie in northern New South
Wales. This remained the only known Australian
oligochaete with two gizzards (the digastric or
digiceriate condition) until Fletcher (1887) erected
the genus Didymogaster for Didymogaster sylvatica
from further south in New South Wales. The latter
genus remained monotypic until Jamieson and
Bradbury (1972) added the geographic replace-
ment species Didymogaster prothecata and val-
idated generic distinction from Digaster. The first
additions to Digaster were D. armifera Fletcher,
1887 and D. perrieri Fletcher, 1889, also from
New South Wales. Subsequently, Spencer (1900)
added three species and Michaelsen (1916) and
Boardman (1932) each added one species, all from
Queensland. Three further Queensland species
were added by Jamieson (1970, 1972). The present
paper is devoted to description of 5 new species and
2 new subspecies of Digaster recorded in intensive
collecting for oligochaetes in Queensland, from its
southern border, at approximately 29 °S., north-
wards along its eastern seaboard to Cooktown, at
approximately 15°30'S., on Cape York Peninsula.
A single, brief collecting foray on the Peninsula
from its east coast at 15°30'S. to Normanton, near
its west coast, at approximately 18°S. was unpro-
ductive, presumably because conditions were dry.
The oligochaete fauna of the northern half of the
Peninsula and of its Carpentaria catchment thus
remains unknown. The map of localities sampled
will be limited to southern Queensland as the most
northerly record for Digaster is 25 S.
The new additions bring the generic complement
to sixteen digastric species and evidence will be
given for the inclusion of a seventeenth species, the
trigastric Perissogaster queenslandica Fletcher,
1889.
Perissogaster was first placed in the synonymy of
Digaster by Beddard (1895) who included its three
species and also Didymogaster. Michaelsen (1900,
1907) separated the three genera but Sweet (1900)
and Jamieson (1963) again included the three
species of Perissogaster , but excluded Didymogas-
ter. Jamieson (1970) presented evidence in support
of exclusion of the type-species of Perissogaster, P.
excavata Fletcher, 1888 and of P. nemoralis
Fletcher, 1889 and retention in Digaster of P.
queenslandica but these changes were deferred
pending examination of Digaster lumbricoides. The
latter species was redescribed in Jamieson (1971b),
in which a lumbricoides species-group was defined,
and it will be appropriate in the present work to
redefine the genus.
Genus Digaster Perrier, 1872
Digaster Perrier, 1872, pp, 94-6; Michaelsen, 1900, p.
196; 1907, p. 162; Stephenson, 1930, p. 839.
268
MEMOIRS OF THE QUEENSLAND MUSEUM
Digaster (part.); Beddard, 1895, p. 484; Jamieson,
1963, pp. 85-6; Jamieson, 1971a, pp. 74-5;
Jamieson, 1971b, pp. 1307-8.
Diagnosis: Setae lumbricine. Gizzards 2, in V
and VI or VI and VII, or 3, in V, VI and VII;
extramural calciferous glands absent though
poorly differentiated oesophageal pouches may be
present. Intestine commencing in or behind XVII;
typhlosole absent. Meronephric; the medianmost
nephridium in caudal segments exonephric with
preseptal funnel. Combined pores of a pair of
racemose or tubuloracemose prostates and the
vasa deferentia in XVIII or rarely in XVII. Testis-
sacs absent. Spermathecal pores intersegmental or
at the anterior margins of their segments.
Detailed Definition: Small to large terrestrial
worms (28 mm to more than a metre long).
Prostomium zygolobous to tanylobous. Circular in
cross section throughout, with terminal anus. First
dorsal pore in 3/4^11/12 (rarely far posterior?).
Setae 8 per segment, in straight or irregular
longitudinal rows; fairly closely paired; cd
significantly wider than ab (minimally 1-2 ab); dd:u
0-2-08 in the forebody. Nephropores scattered,
only sporadically visible. Clitellum annular, oc-
cupying 4-8 segments, the first segment occupied
usually being XIII or XIV. Combined male and
prostatic pores a pair on XVIII varying from
median to a to b lines. Accessory genital markings
present or absent. Female pores paired or, rarely,
single; anteromedian to setae a of XIV. Spermathe-
cal pores 2 or, rarely, 3 pairs, intersegmental or at
the anterior borders of their segments, the last in or
immediately behind 8/9, varying from median to a
to b lines.
Dorsal blood vessel single, continuous onto the
pharynx; last hearts in XII or XIII; supra-
oesophageal vessel present but often poorly differ-
entiated; the last hearts in and posterior to X
latero-oesophageal; dorsoventral commissurals
present anteriorly to the hearts. Subneural vessel
absent. Gizzards 2, in V and VI or VI and VII or
{queenslandica and rare individual variation) 3, in
V, VI and VII; contiguous or separated by
unmodified oesophagus. Oesophagus vascularized
and internally folded and often dilated, in a few to
most segments occupied, but extramural calci-
ferous glands absent; intestine commencing in
XVII to XIX; typhlosole, muscular thickening and
caeca absent.
Wholly meronephric; micromeronephridia in
anteriormost segments aggregated in bands or
usually forming bucco-pharyngeal or exonephric
tufts; succeeding nephridia exonephric astomate
micromeronephridia; in the hindbody the median-
most nephridium on each side possessing a pre-
septal funnel which usually (and typically) is
enlarged as an exonephric megameronephridium.
Flolandric (testes and funnels a pair in each of X
and XI) or metandric (these in XI only); testis sacs
absent; seminal vesicles in IX and XII, XI and XII
or XII only. Prostates 1 pair, racemose or {queens-
landica) tending to tubuloracemose; unipartite or
bipartite; the vasa deferentia (always?) joining the
junction of gland and duct. Penial setae present or,
more commonly, absent. Ovaries 1 pair, in XIII;
ovisacs absent. Spermathecae 2 or 3 pairs, with 1 or
2 often multiloculate diverticula. All structures
sometimes ( D . lumbricoides and D. anomala ) dis-
placed one segment anteriorly by suppression of an
anterior metamere but gizzards never anterior to
V).
Distribution: New South Wales (Port Mac-
quarie and Kyogle areas and the Sydney Basin);
Queensland (South of 25 °S., coastal to as far west
as 151 °E.).
Type Species: Digaster lumbricoides Perrier,
1872.
Species: (NSW = New South Wales; Q =
Queensland).
1 .
D. anomala Jamieson, 1970
Q
2.
D. armifera Fletcher, 1887
NSW
3.
D. binnaburra sp. nov.
Q
4.
D. bradburyi Jamieson, 1970
Q
5.
D. brunneus Spencer, 1900
Q
6.
D. gayndahensis Spencer, 1900
Q
7.
D. gwongorellae Jamieson, 1972
Q
8.
D. lamingtonensis Michaelsen,
1916
Q
9.
D. longmani Boardman, 1932
NSW, Q
10.
D. lumbricoides Perrier, 1872
NSW, Q
11.
D. minima sp. nov.
Q
12.
D. minor Spencer, 1 900
Q
13.
D. nothofagi sp. nov.
Q
14.
D. perrieri Fletcher, 1889
NSW
15.
D. pseudoperichaeta sp. nov.
Q
16
Perissogaster queenslandica Flet-
cher, 1889
Q
17.
D. sexpunctata sp. nov.
Q
Remarks: Jamieson (1970) showed that the type-
species of Perissogaster, P. excavata (see also Bage,
1910), and P. nemoralis had only astomate micro-
meronephridia posteriorly. The third known spec-
ies, P. queenslandica , was shown to have the
dichogastrin condition, with the medianmost neph-
ridium enlarged and stomate. Demonstration
(Jamieson, 1971b) that the dichogastrin condition
was typical of Digaster supported the contention in
the previous paper that P. queenslandica could be
accommodated in Digaster while Perissogaster
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
269
should be revived for P. excavata and R. nemoralis.
Perissogaster is here formally re-instated although
it must be stated that existing material of the genus
is not in sufficiently good condition to allow full
characterization of the genus.
With the re-instatement of Perissogaster as an
independent genus, the number of di- or tri-gastric
genera in the Megascolecinae is raised to thirteen of
which only Digaster, Didymogaster and Per-
issogaster are Australian. The fact that all thirteen
genera have the advanced (meronephric) condition
of the nephridia, all holonephric megascolecines
being monogastric or agastric, suggests that mul-
tiplication of gizzards is a secondary condition, at
least in the Megascolecinae. Ten of the genera with
multiplied gizzards ( Benhamia , Dichogaster, Eudi-
chogaster, Eutrigaster, Lennogaster, Millsonia,
Omodeona, Pellogaster, Rillogaster and Trigaster)
differ from the three Australian genera in lacking
the megascolecin of the male terminalia (pores of a
pair of prostates and of the vasa deferentia
combined on XVIII). This difference, in addition to
its diagnostic value, is of interest in suggesting that
multiplication of the gizzards has occurred in
Australia independently of replication in groups in
other regions. This inference is supported by the
fact that the primitive (that is, holonephric,
monogastric) Australian genera have the same,
megascolecin condition of the male pores, a fact
which suggests that they are ancestral to the di- or
tri-gastric Australian genera and contraindicating
descent of the latter from allopatric genera with
non-megascolecin male pores whether with one or
more gizzards.
The five new species of Digaster and two new
subspecies are all assignable to the very homo-
geneous lumbricoides species-group defined by
Jamieson (1971b). Inclusion of the remaining
species, D. armifera, D. perrieri and D. queenslan-
dica, in the genus confers greater heterogeneity on
the genus but their exclusion cannot on present
evidence be justified. The displacement of the
gizzards in the lumbricoides group from VI and VII
to V and VI in D. lumbricoides and D. anomala by
deletion or supression of an anterior metamere,
together with occurrence of gizzards in V, VI and
VII in the morphologically close D. queenslandica
and (Jamieson, 1963) a variant in D. perrieri, do not
allow convincing subdivision on the basis of
distribution of the gizzards.
Perissogaster s. strict., now limited to P. ex-
cavata and P. nemoralis , from New South Wales, is
diagnosed from all other megascolecids by the
combination of three gizzards, megascolecin male
pores and non-dichogastrin (megascolecin) neph-
ridia. Didymogaster, also from New South Wales,
is distinguished by the combination of two giz-
zards, in VI and VII, megascolecin male pores, the
segmental (not intersegmental) location of the
spermathecal pores and the suboesophageal testis-
sacs.
While the tribes Perionychini and Dichogastrini,
characterized by holonephric and dichogastrin
nephridia respectively, are acceptable groupings by
virtue of their homogeneity, the tribe Megasco-
lecini, diagnosed by non-dichogastrin mer-
onephridia, which includes Perissogaster , is less
satisfactory, being more heterogeneous in mor-
phology and probably in origin. Relegation of
Perissogaster s, strict, to the Megascolecini still
leaves the possibility of close relationship of this
genus with Digaster.
The distribution of Digaster (like that of Did-
ymogaster and Perissogaster ) conforms with the
montane coastal province of the Eastern Subregion
of Australia recognized for the avifauna by Kik-
kawa and Pearse (1969; see Jamieson, 1974). In
view of the large number of very similar species in
the genus, this limited geographical distribution
does not appear to be evidence, as might have been
thought, that Digaster is a relict genus with a
former wide distribution in Australia. If it were
taken to be a relict with a much-contracted
distribution it would be necessary to explain why
the similarly digastric Didymogaster and the tri-
gastric Perissogaster also survived only in this
province. Nor can the limited distribution be
explained in terms of recent introduction from
outside Australia for this would require either
concomitant introduction of all its species and,
presumably, of Didymogaster and Perissogaster, or
massive speciation since introduction. It seems
more acceptable that Digaster (and the clearly
related Didymogaster and the less certainly related
Perissogaster) originated locally from monogastric
holonephric worms, or their meronephric de-
scendants, sufficiently long ago for considerable
speciation but not long enough ago for col-
onization of other parts of Australia. Part of the
restriction in range must be due to ecological
preferences and not to lack of time for dispersion
but it must be reiterated that the digastric condition
suggests relatively recent, and local, origin of the
genus for reasons given above.
Key to the Species and Subspecies of Digaster
1. 3 oesophageal gizzards present . .
D. queenslandica *
2 oesophageal gizzards present . . . . 2
2. Male pores in XVII 3
Male pores in XVIII 4
3. Accessory genital markings a pair of oval
270
MEMOIRS OF THE QUEENSLAND MUSEUM
glandular areas in front of and behind the
male pores, posteriorly in XVI and ante-
riorly in XVIII. Other accessory genital
markings absent
. .D. lumbricoides lumbricoides, Fig. 9F
Accessory genital markings midventral, un-
paired transverse pads in 19/20 and 20/21;
sometimes also in varying numbers of in-
tersegments 18/19, 21/22 24/25, and in
8/9-10/11 . . . . D. anomala. Fig. 9B
4. Gizzards in V and VI 5
Gizzards in VI and VII . . . . . . 6
5. A square glandular pad midventral in each of
XI and XII. At maturity, 2 elliptical genital
markings, one behind the other, lateral of the
male pores. A pair of ellipses often present in
each of XVII and XIX
D. armifera*, Fig. 9C
Genital markings absent from XI and XII. 3
transverse ventral ridges typically present, in
17/18, XVIII and 18/19 . . . D . perrieri*
6. Dorsal setal couples ( cd) in caudal segments
displaced far dorsally, the 4 setae in 4
equispaced lines . . 7
Dorsal setal couples not displaced far dorsally;
significantly further apart than the setae of
each couple 8
7. Accessory genital markings a midventral pad
in X, XVI and XVII, and a pair of white
tumescences presetally in XIX. .
D. pseudoperichaeta , Fig. 7
Accessory genital markings a midventral cir-
cular papilla in XIII, XX, XXI and in
varying numbers of segments XII, XV, XVI,
XIX, and XXII— XXIV. A paired marking
sometimes present in X
D. minima, Fig. 6A
8. One or more midventral unpaired genital
markings (papillae or transverse pads) in or
within a segment or two of IX . . . . 9
Midventral unpaired genital markings absent
or, if present, not in the vicinity of IX . 13
9. 3 pairs of small disc-like markings located
between the male porophores on XVII
D. sexpunctata, Fig. 8
No genital markings between the male poro-
phores . . . . . . 10
10. Metandric (testes and funnels in XI, seminal
vesicles 1 pair, in XII only). First dorsal pore
in 5/6 or 6/7 . . D. longmani (part.)
Holandric (testes and funnels in X and XI,
seminal vesicles 2 pairs, in IX and XII or XI
and XII). First dorsal pore in 4/5 . . 11
11. Seminal vesicles in XI and XII. First dorsal
pore in 5/6 . . . . D. longmani (part.)
Seminal vesicles in IX and XII. First dorsal
pore in 4/5 12
12. Midventral unpaired papillae transversely
elongate on segments following the sper-
mathecal pores; intersegmental in the vic-
inity of the male pores where there may be
paired papillae also D. nothofagi, Fig. 6B
Midventral unpaired papillae circular or longi-
tudinally extended on segments following
and often including the spermathecal pores;
not present in the vicinity of the male pores
where there are usually paired segmental
markings . . . . D. binnaburra, Fig. 2
13. A midventral circular, oval or almost bifid pad
on XVIII between or including the male
pores 14
No midventral pad in XVIII. Genital markings
usually present elsewhere 15
14. Female pore unpaired. No genital markings
present in addition to that on XVI II . .
. . D. gwongorel/ae, Fig. 9E
Female pore paired. Typically with an ad-
ditional genital marking on XIX..
D. minor*. Fig. 9D
15. Metandric (testes and funnels in XI, seminal
vesicles in XII only) 16
Holandric (testes and funnels in X and XI,
seminal vesicles 2 pairs, in IX and XII or XI
and XII) 18
16. Unpaired transverse midventral pads in-
tersegmental on the clitellum. First dorsal
pore in 9/1 0-1 2/13
D, brunneus (part.), Fig. 4B
Clitellar pads absent. First dorsal pore in
5/6-12/13 17
17. First dorsal pore in 9/10 or further posteriorly
D. brunneus (part.), Fig. 3B
First dorsal pore in 5/6 or 6/7
D. longmani (part.)
18. A pair of glandular patches present in the vic-
inity of 17/18 in front of the male pores.
Other genital markings present or absent
19
No paired glandular patches anterior to the
male pores. Male pores in ab of XVIII on
small papillae within a common lip-like
ridge. Midventral unpaired transverse pads
extending to ab in 19/20, 20/21 and 21/22
D. gayndahensis* , Fig. 9G
19. A pair of genital markings in or just anterior to
17/18 and a similar, unilateral (left) marking
in or just anterior to 18/19. First dorsal pore
in 5/6, Spermathecal ducts not dilated . .
D. lamingtonensis*. Fig. 9H
Paired markings in or near 17/18; markings if
present near 18/19 paired. First dorsal pore
in 4/5 or 8/9-10/11. Spermathecal ducts not
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
271
dilated 20
20. First dorsal pore in 4/5. Male pores preceded at
17/18 and succeeded at 18/19 by a pair of
oval glandular areas
D. lumbricoides kondalilla, Fig. 5
First dorsal pore in or behind 8/9. Male pores
preceded at 17/18 by a pair of glandular
areas but paired markings behind the pores
indistinct or absent 21
21. Body approximately 90-140 mm long and 4-5
mm wide . D. bradburyi bradburyi , Fig. 9A
Body approximately 185 265 mm long and
8-10 mm wide
D. bradburyi bunyaensis. Fig. 3A
* Asterisked species have not been recorded, and are therefore not described, in this study. Published illustrations of
their genital fields are, however, reproduced. The fields of D. perrieri (see Jamieson, 1963) and D. queenslandica (see
Fletcher, 1889) on available specimens do not merit illustration.
Digaster anomala Jamieson, 1970
Figs. 1, 9B; Table 1
Digaster anomala Jamieson, 1970, pp. 40-3, figs. 1 B, C;
2D, E.
Material Examined: Or 3, 152°50 / E. 27°15'S.,
Kobble, rocky hillside covered by Lantana, E. Bradbury,
10 Jul 1970 and 10 Aug 1970, BM(NH) 1973.10.1-5,6-15;
rocky hillside by road, E. Bradbury, 8 Oct 1970, BM(NH)
1 973. 1 0. 1 6-21 . Or 5, 1 52 °45'E. 27 2 US., 6 miles from M t.
Nebo on Mt. Glorious road, on side of eucalypt-covered
hill by ferns and nettles, E. Bradbury, 12 Jun 1970,
BM(NH) 1973.10.22-23; Mt. Glorious, rainforest, E.
Bradbury, 12 Jun 1970, BM(NH) 1973.10.24; 8 Oct 1970
BM(NH) 1973. 10.25; I. Naumann, 1 3 Sep 1971 , BM(NH)
1973.10.26. Or 7, 152°47'E. 27 23'S„ Mt. Nebo road Mt.
Nebo, in loamy soil in gully in eucalypt area, E. Bradbury,
12 Jun 1970, 3, Jamieson collection; 1 mile from Mt. Nebo
on road, E. Bradbury, 10 Aug 1970, BM(NH) 1973.10.27-
28. Or 11, 152°54'E. 27°28'S., Gold Creek Road,
Brookfield, in loose damp soil in leaf mould near creek, E.
Bradbury, 24 Apr 1970, BM(NH) 1973.10.29-44, 45. Or
12, 152 54.5'E. 27°30'S., Willunga Street, Brookfield, on
dirt road surface after heavy rain, B. Jamieson, 9 Dec
1970, BM(NH) 1973.10.46-55; Brookfield, E. R. John-
son, no date, BM(NH) 1973.10.56. Or 13,
152°5TE. 27 C ‘32'S., banks of Kholo Creek, Mt. Crosby
Road, near Brisbane, B. Jamieson and/or E. Bradbury, 1 3
Mar 1970, 24 Apr 1970, 2 Feb 1970, 4 Mar 1970, 2 Jun
1966, BM(NH) 1973.10.57, 58-59, 60-65, 66-67, 68-70. Or
14, 152°53'E. 27°35 / S., Moggill, near Brisbane, E. R.
Johnson, no date, BM(NH) 1973.10.71-72. Or 16,
152°58'E. 27°28'S., Eastern part of Mt. Coot-tha,
Brisbane, in sandy Lantana soil, L. Lyndon, 5 Sep 1971,
BM(NH) 1973.10.73-74. Or 18, 152°58'E. 27 C 33'S., Fig
Tree Pocket, Brisbane, cleared land near river bank, Mrs
C. C. Wallace, 27 Oct 1969, QM G83 16-24. Or 20,
152°49'E. 27°49'S., Mt. Flinders, H. Mayne, no date,
BM(NH) 1973.10.75-77, 78.
The following account is abstracted from Jamie-
son (1970). New material is listed under ‘Material
Examined’ and is discussed under ‘Genital Field
Variation’, below.
1 - 69-105 mm, w (midclitellar) = 3-4 mm, s
103-146. Unpigmented excepting the brick red
clitellum. Prostomium epilobous -. First perforate
dorsal pore 4/5. Setae in 8 longitudinal rows
throughout; commencing on segment I;
aa:ab:bc:cd:dd in XII averaging 2-9: 1:4- 3:3-0: 19-6;
dd:u = 047-0-55. Clitellum annular, XIII-XVII.
Male pores on XVII in a. Accessory genital
markings midventral unpaired transverse pads in
19/20 and 20/21 and sometimes also in 18/19,
21/22, 23/24 and 24/25; sometimes with slight
ventral epidermal elevations in some or all of
intersegments 8/9-10/11. Female pores antero-
median of seta a in XIII. Spermathecal pores 2
pairs, well median of a lines, in 6/7 and 7/8.
Last hearts in XII. Gizzards in V and VI;
calciferous glands absent; intestinal origin XVII.
Nephridia meronephridia with exonephric tufts in
III and, caudally, an exonephric stomate mega-
meronephridium median to astomate micromero-
nephridia in each segment. Holandric; testes in IX
and X; seminal vesicles racemose in VIII or VIII
and XI. Ovaries in XII; ovisacs absent. Prostates
bipartite, in XVII. Spermathecae 2 pairs, in VII and
VIII; diverticulum large, ovoid-subspherical, in-
ternally multiloculate, almost sessile on the ectal
end of the duct; length of a spermatheca = 3-0 mm;
ratio of length: length duct = 3-5; ratio of length:
length diverticulum = 4-7.
Genital Field Variation: Specimens from the
type-localities (including the new material from Or
12, Willunga Street, and Or 13, Kholo Creek)
differ from material from all other recorded
localities in having the spermathecal pores median
to setal lines a, not in or lateral of a. These two
alternative conditions may, for convenience in
discussion, be said to characterize a typical and an
atypical population, respectively. Table 1 lists the
distribution of transverse genital markings in 31
typical specimens, including the holotype and 1 1
paratypes as described by Jamieson (1970), and in
30 atypical specimens from all localities sampled. It
is seen that in the typical population transverse
genital markings are present in a few to most of
intersegments 6/7-11/12, 15/16, 16/1 7—24/25. The
distribution in the atypical population agrees with
this except for absence in 11/12, 16/17 and
22/23-24/25. These absences in atypical specimens
272
MEMOIRS OF THE QUEENSLAND MUSEUM
Fig. 1: Queensland records of Digaster. • = D. anomala. Other species shown by labelled symbols. Unlabelled blank
circles indicate sampled localities which did not yield Digaster. Localities are referred to in the text by co-ordinates
and number (e.g. Or 3). The inset shows the Australian distribution of this endemic genus.
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
273
may not represent a significant difference as the
frequency of markings at these sites is low in the
typical specimens. In both populations the genital
markings are most frequent in 19/20-20/21, being
virtually constant in 19/20 and very frequent in
20/21. Good agreement between the two pop-
ulations is seen in the frequency of 23-26% in the
occurrence of a marking in 21/22 but a major
difference is seen in the frequency in 17/18 and
18/19, for in the typical population the marking is
more frequent in 17/18 (55%) than in 18/19 (19%)
whereas in the atypical population the frequency is
reversed, 13% in 17/18 against 83% in 18/19. A
further difference is that the marking in 18/19 when
present is always simple in the typical specimens
while it is usually composed of a pair of more or less
conjoined ellipses in the atypical specimens. Penial
setae have not been observed in typical specimens
whereas in the atypical population in 19 specimens
examined for penial setae they were present in 13
specimens (BM(NH) 1973.10.1,6, 7, 16, 17, 25,27,
30, 45, 74. Jamieson collection, Mt. Nebo), though
absent in 6 specimens (BM(NH) 1973.10.22, 23, 56,
71, 75, 78). All specimens show the anomala
condition, apparent suppression of the first met-
amere so that segment I is setigerous and structures
including the ovaries and prostates are displaced
one segment anteriorly relative to their position in
most Megascolecinae.
TABLE 1: Distribution of Genital Markings in the
Digaster anomala complex.
Intersegment
occupied
Number of specimens
°/
/O
Typical
population %
(of 31)
Atypical
population
(of 30)
6/7
17
54.8
11
36-6
7/8
17
54-8
11
36-6
8/9
10
32-3
1
3-3
9/10
10
32-3
1
3-3
10/11
6
19-4
1
3-3
11/12
3
9-7
0
0
12/13
0
0
0
0
13/14
0
0
1
3-3
14/15
0
0
1
3-3
15/16
1
3-2
2
6-7
16/17
6
194
7
0
17/18
17
54-8
4
13-3
18/19
6
19 4
25
83-3
19/20
31
100
29
96-7
20/21
28
90-3
. 24
800
21/22
8
25-8
7
23-3
22/23
1
3-2
0
0
23/24
3
9-7
0
0
24/25
2
6-5
0
0
Type-Locality: Brisbane (Gap Creek Road and
Willunga Street, Or 12; Kholo Creek, Or 13).
Remarks: The anomalous segmentation, with
male pores on XVII, together with the presence of a
transverse genital marking in 19/20 and usually in
20/21 , distinguish D. anomala from all other species
of Digaster , though the same segmental anomaly
occurs in the nominate subspecies of D. lumb-
ricoides. The term ‘D. anomala complex’ is here
proposed for the combined typical and atypical
populations as defined above. That the two
populations are conspecific remains to be
confirmed but they are clearly very closely related
and if specifically distinct would have to be
considered sibling species.
Digaster binnaburra sp. nov.
Figs. 1; 2; 10A, B; 11 A; Table 2, 3
Material Examined: Lamington National Park,
South Queensland: Ps 4, 153°11 / E. 28°12'S: 3 miles
from Binna Burra, in rainforest with Casuarinas, B.
Jamieson and E. Bradbury, 25 Mar 1970, Holotype(H)
QM G7382, Paratypes (P) 1-6 (clitellate), P7-11 (with
genital markings but aclitellate), QM G7383-93; Binna
Burra, in Tristania-Casuarina and mixed broadleaf
forest, with tree ferns and palms, B. Jamieson and E.
Bradbury, 25 Mar 1971, P14-17, 29, BM(NH)
1973.10.79-82, 83; Binna Burra, rainforest, B. Jamieson
and E. Bradbury, 24 Mar 1971, PI 9, 20 (clitellate), 21 25
(with genital markings but aclitellate), BM(NH)
1973.10.84-85, 86-90. Ps 3, 153°09'E. 28°14'S„ in park
near O’Reilly’s Guest House: at Elbana Falls, B.
Jamieson, 3 May 1970, P18, BM(NH) 1973.10.91; in
rainforest, B. Jamieson and T. Walker, 18 Mar 1972, P12,
13, BM(NH) 1973.10.92-93; B. Jamieson, 19 Mar 1972,
P26^28, BM(NH) 1973.10.94-96.
1 = 62-65 mm. w (midclitellar) = 3 mm, s
188,185 (H,P1). Form circular in cross section
throughout; moderately slender; lacking strong
secondary annulation. Pigmentless, excepting the
brick-red clitellum, in alcohol. Prostomium pro-
epilobous; no canaliculi present dorsally or vent-
rally on it or the segments. First dorsal pore 4/5;
pores inconspicuous. Setae in 8 regular longitu-
dinal rows, commencing on II; setae a and b absent
in XVIII. Nephropores not externally visible.
Clitellum annular, XIV-XVII(H), 1 XVIII(Pl),
with weak extension to ^ XIII; intersegmental
furrows and dorsal pores obscured excepting at
13/14 and 17/18; setae retained. Male pores minute
in ab of XVIII, each at the centre of a small, low,
circular porophore which at maximum develop-
ment (PI) fills the segment longitudinally. Ac-
cessory genital markings small inconspicuous
transversely oval pads paired postsetally in XVII,
XIX (H,P1) and XX(H), in ab; the markings and
274
MEMOIRS OF THE QUEENSLAND MUSEUM
the male porophores contained within a ventral
glandular area. Unpaired low boss-like accessory
genital markings present anteriorly, a postsetal
marking in each of VIII(H), IX and X(H,P1) and a
presetal marking in IX and XI (H) or X (PI).
Female pores small but distinctly visible ante-
romedian of a of XIV, about \ aa apart. Sper-
mathecal pores 2 pairs, in 7/8 and 8/9, in a lines on
small papillae (H), or in a and ab respectively (PI).
Fig. 2. Digaster binnaburra sp. nov. Genital field of
holotype. For key to abbreviations, see Fig. 9.
Some preclitellar septa thickened; 10/11 the
thickest, moderately strongly thickened. Dorsal
blood vessel single, continuous onto the pharynx.
Last hearts in XII, those in X XII latero-
oesophageal, each with a connective from the
dorsal and the supra-oesophageal vessel. Supra-
oesophageal vessel present but extent indetermin-
able as it is indistinctly delimited from the roof of
the oesophagus. Subneural absent. Oesophagus in
V thinwalled but fusiform and glossy and therefore
gizzardlike in appearance, hidden by septal glands.
Two strong slightly elongate spherical gizzards,
each with anterior rim, in VI and VII, separated by
a short region of unmodified oesophagus. Oeso-
phagus almost suppressed, by backward extension
of the gizzards, in VIII and IX; short in X-XII;
swollen and vascularized in XIII-XVII, especially
in XIV(Pl), XV and XVI(H,P1) in each of which it
has a reniform but not separate dilatation on each
side which is slightly bilobed owing to a circumfer-
ential vessel which arises from the closely adherent
dorsal vessel. Parallel sinuous rugae present on the
internal wall of the oesophagus in XV and XVI but
no true extramural calciferous glands present.
Intestinal origin XIX; typhlosole, caeca and mus-
cular thickening absent. Nephridia meronephridia:
small tufts in II, III and IV sending a common
composite duct on each side to the peristomium
near the mouth; large tufts in V sending a
composite (exonephric) duct to the region of
segment II(H,P1) where (PI) it appears to continue
to the wall of the pharynx. Thereafter with lateral
bands of numerous astomate, exonephric micro-
meronephridia. In the anterior intestinal region
with 8 such nephridia on each side; caudally with
the medianmost nephridium enlarged as an exo-
nephric megameronephrium with preseptal funnel.
Holandric; gymnorchous (sperm funnels iridescent
in X and XI); seminal vesicles racemose, in IX and
XII. Prostates racemose, tongue-shaped, that on
the left in the holotype consisting of adpressed
dorsal and ventral halves; vas deferens joining the
short, muscular medianly directed duct at its
junction with the gland. Penial setae absent.
Metagynous; ovaries, consisting of several strings
of oocytes, and funnels in XIII; ovisacs not
recognizable. Spermathecae 2 uniform pairs; am-
pulla elongate ovoid; duct cylindrical, bearing an
iridescent rounded multiloculate diverticulum at its
ectal extremity; length of right spermatheca of VIII
(H) =1-7 mm; ratio of length spermatheca: length
duct = 2-4; ratio of length spermatheca: length
diverticulum = 5-8.
Remarks: The midventral anterior genital mark-
ings which, like the paired markings in the male
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
275
TABLE 2: Genital Markings in 18 Specimens of
Digas ter binnaburra.
Specimen*
Total
Midventral markings
VI Presetal
0
Postsetal
P13, 18
2
VII
Presetal
—
0
Postsetal
P12-14, 17-20, 26, 28
9
VIII
Presetal
P3, 13 14, 17 20, 26
8
Postsetal
H, P12-15, 17-20, 26, 28
12
IX
Presetal
H, P3, 12-15, 17-20, 26-28
13
Postsetal
H, PI 5, 12-15, 18-20, 26-28
16
X
Presetal
Pl-5, 12-15, 17-20, 26-28
16
Postsetal
H, PI -5, 12-20, 26-28
18
XI
Presetal
H, PI 5, 12 15, 17 20, 26-28
17
Postsetal
P2-5, 12-20, 26-28
16
XII
Presetal
P5, 19
2
Postsetal
—
0
Paired markings in ab
XVIII Postsetal
H, Pl^l, 12-13, 16-20, 26
13
XIX
Postsetal
H, Pl-4, 12-20, 26-28
17
XX
Presetal
H (left), PI 7, 26
3
Postsetal
H, P12-13, 16, 18, 27, 28
7
XXI
Presetal
P26
1
Postsetal
P26
1
*H = Holotype, P = paratype.
field, are present in partly mature aclitellate and in
clitellate specimens, permit ready diagnosis of D.
binnaburra. The postsetal markings in X are
constant in the type sample and are almost
invariably accompanied by markings postsetally in
IX, presetally in X, and pre- and post-setally in XI.
Some fusion of presetal and postsetal markings is
common.
Digaster bradburyi Jamieson, 1970
Digaster bradburyi Jamieson, 1970, pp. 35-40, figs. 1A,
2A-C, 3.
The following account is based on Jamieson
(1970) and new material, comprising a new sub-
species, from Bunya Mountains.
1 = 88-265 mm, w (midclitellar) = 4-10 mm, s
= 116-269. Body slender, circular in cross section
throughout, pigmentless or pale brown, clitellum
pigmented purplish brown. Prostomium epilobous
(or sometimes prolobous?). First dorsal pore 8/9,
9/10 or 10/11. Setae in eight longitudinal rows
throughout, commencing on II; minute and not, or
only sporadically visible in the forebody, con-
spicuous on the clitellum owing to pale encircling
fields; aa:ab:bc\cd:dd in XII (measured for nom-
inate subspecies only) averaging 3-9: 1 :3-3:2-7: 13-5;
dd\u — 0-41-0-45; c and d not especially dorsal
posteriorly. Nephropores not externally recogniz-
able. Clitellum annular, XIII, ^XIII, XIV-XVIII,
i, ^XIX (= 5|-6^ segments) but its pigmentation
may reach XII XX. Male pores on XVIII in a or
ab. Accessory genital markings an approximately
oval glandular marking occupying the postsetal
portion of XVII and extending over intersegment
17/18 slightly median or slightly lateral of the male
pores. One or two pairs of less distinct markings
present behind the male pores, in XVIII and/or
XIX, or absent. Female pores anteromedian to
setae a of XIV. Spermathecal pores 2 pairs, in 7/8
and 8/9, in a lines.
Dorsal blood vessel continuous onto the phar-
ynx. Last hearts in XII; those in X XII latero-
oesophageal, each receiving a connective from the
dorsal vessel and from a poorly or well defined
supra-oesophageal vessel; subneural vessel absent.
Gizzards in VI and VII, not separated by
unmodified oesophagus. Extramural calciferous
glands absent. Intestinal origin \ (or anterior?)
XVIII; typhlosole, caeca and muscular thickening
absent. Nephridia meronephridia: tufts in II-VI;
those in IV VI, at least, enteronephric. Holandric;
testes in X and XI; seminal vesicles racemose, in IX
and XII. Ovaries in XIII; ovisacs absent. Prostates
276
MEMOIRS OF THE QUEENSLAND MUSEUM
bipartite, in XVIII; penial setae absent though
follicles a and b may be present in the male
porophores. Spermathecae 2 approximately equi-
sized pairs, in VIII and IX, though the ampullae
may be displaced into the preceding segment, each
with an approximately ovoid ampulla and a well
demarcated, dilated duct which is widest slightly
ectal of its midlength and bears, at or ental of its
widest part, 1 or sometimes 2 bulb-shaped, exter-
nally simple diverticula; length of a spermatheca
2- 4-4-0 mm; ratio of length spermatheca: length
duct = 1 -9—2-8; ratio of length spermatheca: length
diverticulum = 4-9-6-2.
Type-Locality: Kholo Creek, near Brisbane
(Or 13).
Wider Distribution: Bunya Mountains (Nq 2).
Digaster bradburyi bradburyi Jamieson, 1970
Figs. 1; 9 A
Digaster bradburyi Jamieson, 1970, pp. 35-40, figs. 1A,
2A-C, 3.
Material Examined. Or 13, 1 52°51 E. 27°32'S.,
Kholo Creek at Ugly Gully, E. Bradbury, 10 May 1970, 4
specimens, BM(NH) 1973.10.112-115.
For detailed account see Jamieson (1970). Char-
acters as for specific description, above, with 1 =
88-140 mm, w (midclitellar) - 4-5 mm. Body,
excepting clitellum, pigmentless. Prostomium epi-
lobous |4 Clitellum i XIII, XIV XVIII, 4 \
XIX (= 4-6 segments). Male pores in ab. Ac-
cessory genital markings a pair of approximately
oval, sunken, translucent markings occupying the
postsetal portion of XVII and intersegment 17/18
only; slightly median to the male pores.
Supra-oesophageal very well developed. In-
testinal origin \ XVIII. Nephridial tufts in IV
sending a composite duct on each side to the lateral
extremity of the mouth; those in V and VI to
pharynx behind brain. Ectal portion of spermathe-
cal duct dilated and joined entally by diverticulum;
length spermatheca = 2 4-3-6 mm; ratio length:
length duct = 1 -9-2-8; ratio length: length diverti-
culum = 4-9-7- 8.
Type-Locality: Kholo Creek, near Brisbane (Or
13).
Remarks: Of the above characters, the only
significant differences from the second subspecies
appear to be the smaller size, perhaps effluence of
the tufted nephridia of V and VI into the pharynx
and not into the buccal cavity into which those in
III- VI discharge in the Bunya specimens, and
possibly the unpigmented body.
Digaster bradburyi bunyaensis subsp. nov.
Figs. 1; 3A; 10C, D; 1 IB.
Material Examined: Nq 2, 151°35'E. 26°57 , S.,
Bunya Mountains National Park, in mixed rainforest at
c.1000 metres, B. Jamieson and E. Bradbury, 20 Feb
1971, Holotype (H), QM G8333; Paratype (P) 1,
BM(NH) 1973.10.97; P2, QM G8334: P3, Jamieson
collection.
I = 185 (PI), 265 (H) mm, w (midclitellar) = 8 0
(PI), 9-6 (H) mm, s = 231 (PI), 237 (H). Secondary
Fig. 3: A, Digaster bradburyi bunyaensis subsp. nov. Genital field of holotype. B, D. brunneus Spencer, 1900, specimen
from Wolvi, BM(NH) 1973.10.105; the clitellum is shown widened owing to dorsal incision. For key to
abbreviations, see Fig. 9.
JAMIESON: GENUS DIG ASTER IN QUEENSLAND 277
TABLE 3: Imtersetal Distances in Segment XII in Digaster
Species
Intersetal distance, mm
Standardized as
% of periphery
aa
ab
be
cd
dd
dc
cb
ba
aa
ab
be
cd
dd
dc
cb
ba
D. binnaburra
Holotype
0-9
0-3
1-3
0-7
3 4
0-7
1-2
0-3
10 1
2-8
14 2
8-5
39-7
8-5
13-4
2-8
Paratype 1
0-8
0-3
1-2
0-8
3-0
0-8
M
0-3
9-3
3-1
14 9
9-9
36-5
9-3
13 6
3-4
Mean
9-7
3-0
14-5
9-2
38-1
8-9
13 5
3-1
Interval/ab
3-3
1-0
4-9
3-1
12-8
3-0
4-5
1-1
D. lumbricoides
kondalilla
Holotype
1-1
0-3
1-1
0-9
4-2
1-1
1-1
0-4
10-9
31
11-1
9-0
40-8
10-2
11-1
3-9
Interval/ab
3-6
1-0
3-6
2-9
13 3
3-3
3-6
1-3
D. minima
Holotype
0-5
0-2
1-3
0-2
1-0
0-2
1-3
0-2
10-8
3-4
26-9
4-0
20-6
4-0
26-9
3-4
Interval/ab
3-2
1-0
7-9
1-2
6-0
1-2
7-9
10
D. nothofagi
Holotype
1-3
0-3
1-3
0-7
2-1
0-7
1-3
0-2
16-3
3-4
16-5
9-2
26-4
8-8
16-3.
3-1
Interval/ab
4-8
1-0
4-8
2-7
7-7
2-6
4-8
0-9
D. pseudoperichaeta
Holotype
0-8
0-3
1-0
0-3
2-3
0-3
1-3
0-3
12 9
4-0
15 3
4-8
34-5
4-4
19-3
4-8
Paratype 1
11
0-3
1-4
0-4
1-4
0-3
1-5
0-3
15-6
4-7
21-1
6-3
21-1
4-7
21-9
4 7
Mean
14-2
4-4
18-2
5-5
27-8
4-6
20-6
4-8
Interval/ab
3-3
1-0
4-2
1-3
6-4
1-1
4-7
1-1
D. sexpunctata
Holotype
1-2
0-4
1-3
0-7
6-8
0-8
H
0-3
9-5
3-0
10-0
5-8
54-1
6-0
9-0
2-5
Paratype
1-2
0-4
1-0
0-8
7-4
0-8
11
0-4
9-2
3-1
7-7
5-8
57-0
5-8
8-2
3-1
Mean
9-4
3-1
8-9
5-8
55-6
5-9
8-6
2-8
Interval/ab
3-1
1-0
2-9
1-9
18-1
1-9
2-8
0-9
annulation strongly developed but not obscuring
the primary segmentation. Pigmented pale brown
with the clitellum deep purplish brown. Pro-
stomium epilobous \, closed (H); or appearing
prolobous with lateral borders continued to 1 /2 but
not more conspicuous than the numerous longi-
tudinal peristomial grooves (PI). First dorsal
pore 9/10. Postclitellar setae visible with some
difficulty; setae a and b in XVIII represented by two
follicles behind each male pore (only a single, left,
follicle in PI). Clitellum annular, XIII ^XIX but
continued as rosy pigmentation throughout XII
and XX; interrupted in bb behind the setal arc of
XVII; only intersegmental furrow 13/14 complete;
dorsal pores occluded; setae visible. Male pores
transverse slits equatorially in XVIII in a , on
elliptical porophores. Accessory genital markings a
pair of oval unelevated glandular patches pos-
teriorly in XVII and apparently extending slightly
on to XVIII (intersegmental furrow 17/18 ob-
scured) in ab (all specimens), each patch (H) with a
whitish laterally papillated border; a further pair of
patches presetally in XIX centred in b but in-
distinctly delimited; in P2 a pair of glandular
depressions situated posteriorly in XVIII and
similarly placed in XIX; each slightly extending
onto the succeeding segment. Midventral and other
accessory genital markings absent. Female pores
approximately } aa apart. Spermathecal pores 2
pairs of conspicuous slits in a on small papillae.
Septa 8/9-11/12 very thick (6/7, 7/8 and 12/13
moderately strongly thickened). Latero-
oesophageal hearts, in X XII, each arising from
the dorsal vessel and receiving a slender connective
from an imperfectly differentiated supra-
oesophageal vessel. Pharynx ending in IV; oeso-
phagus narrow in V; gizzards firm but only
moderately large, tubular. Oesophagus almost
suppressed in VIII by backward extension of the
gizzards. Nephridia (H): tufts in II-VI, increasing
in size posteriorly to large in VI; those in II
apparently (but not certainly) exonephric; those in
II-VI enteronephric, sending composite ducts to
the anterior region of the buccal cavity. Dense
278
MEMOIRS OF THE QUEENSLAND MUSEUM
lateral bands of nephridia on the posterior septum
of YII are exonephric by numerous ducts which
enter the body wall anteriorly in the segment;
nephridia in succeeding segments less numerous,
exonephric micromeronephridia on the body wall
posteriorly in their segments; nephridia in anterior
intestinal segments equatorial on the body wall,
approximately 1 0 per side in each segment; caud-
ally with a slightly enlarged exonephric megamer-
onephridium, with preseptal funnel, median to
astomate, exonephric micromeronephridia on each
side. Prostate ducts bound by a muscular and
fibrous covering, on each side, to the ventral body
wall. Spermathecae each with an ovoid though
asymmetrical ampulla and a well demarcated
broadly fusiform duct which bears dorsolaterally,
at its widest point, 1 or (H, right VIII) 2 bulb-
shaped diverticula which are externally simple but
internally have complex lumina; length of right
spermatheca of VIII (H) = 4 0 mm; ratio length:
length duct = 2-0; ratio length: length diverticulum
= 6 - 2 .
Remarks: The large size of the specimens of this
taxon and general morphological similarity (in-
cluding location of the first dorsal pore in the
vicinity of 9/10) to those specimens of D. brunneus
which lack transverse genital markings at first
suggested their close relationship, if not
conspecificity with D. brunneus. However, it
differs from D. brunneus but conforms with D.
bradburyi, in dilatation of the spermatheca! ducts,
holandry, in the bipartite form of the prostates and
in the form of the genital markings in 17/18.
Though the body size is much greater than that of
D. bradburyi, the dimensions and proportions of
the spermathecae are strikingly similar, though
also similar to those of D. brunneus. Dilatation of
the spermathecal ducts and bipartite prostates are
also seen in the closely similar D. lumbricoides
kondalilla but the latter differs in location of the
first dorsal pore in 4/5. It is possible that the
Kondalilla specimens are closer to D. bradburyi
than to lumbricoides but the uncertainty in this
respect is a reflection of the especially close
relationship, within the genus, of lumbricoides and
bradburyi and, less closely, brunneus and lamington-
ensis.
The validity of placing bunyaensis in D. brad-
buryi as a subspecies in the sense of a geographical
race is questionable. The Bunya Mountains are
geographically somewhat isolated but whether the
Bunya and typical populations could interbreed if
in contact is doubtful owing to their great differ-
ence in body size. If reproductively and geographi-
cally isolated they would have to be considered
separate, though very similar (sibling?) species. If,
on the other hand, gene exchange is possible
through as yet unknown intervening populations,
with specimens intermediate in size, the grounds
for separation into distinct subspecies or species
might be considered lost. Taxonomic recognition
of the large mountain forms seems desirable,
however, on present evidence and similarities with
D. bradburyi warrant inclusion in this species as a
distinct subspecies.
Digaster brunneus Spencer, 1 900
Figs. 1; 3B; 4A, B; 10E-G; 11C.
Digaster brunneus Spencer, 1900, p. 66, pi. 12, figs. 103-
5.
Material Examined: No 5, 151 0 53'E. 24°55'S., 10
miles from Gin Gin, in very damp clay soil, Mrs C. C.
Wallace, 7 Dec 1969, QM G8325-6. Np 2, 151°35'E.
27°37'S., Burnett River, Gayndah: in loam covered by
weeds on river bank, E. Bradbury, 1 1 Mar 1970, BM(NH)
1973.10.98-99; orange orchard on river bank, E. Brad-
bury, 11 Mar 1970, BM(NH) 1973.10.100. Nq 1,
151°57'E. 26°94'S., Murgon, in shallow soil, collector
and date not known, BM(NH) 1973.10.101. Op 2,
152°0LE. 25°59'S., 5 miles N. ofTansey, on road covered
with water, E. Bradbury, 7 Feb 1971, BM(NH)
1973.10.102-104. Op 3, 152°36'E. 25°50'S„ 1 mile south
of Bauple, in dry clay soil in grassy area near creek, Mrs
C. C. Wallace, 6 Dec 1969, 2 Jamieson collection, QM
G8327-31. Oq 2, 152°50'E. 26°10 , S„ Wolvi, C. A.
Scarlett, no date, BM(NH) 1973.10.105.
The following account is based on two Gayndah
specimens, BM(NH) 1973.10.98 and 100, with
some additional data from other specimens where
indicated. Spencer’s brief account is in agreement
except where noted.
1 — 210 H — 430 mm (150 mm, Spencer); w
(midclitellar) = 10-11 mm (6 mm, Spencer); s =
161 H — 397 (Specimen 98 is a posterior amputee).
Form slender, circular in cross section throughout;
secondary annulation strongly developed, obscur-
ing the primary segmentation. Pigmentless in
alcohol excepting the pale chocolate-brown clitel-
lum. Prostomium prolobous; peristomium with
numerous longitudinal grooves. First dorsal pore
10/11 (with a rudiment at 9/10, specimen 100) or
11/12 (specimen 98); location not given by Spencer.
Setae minute, only sporadically visible; a and b
absent in XVIII. Nephropores not recognizable.
Clitellum annular, fXIII(?), XIII-XIX; interrupted
ventrally between and behind the male pores;
intersegmental furrows weakened and laterally
interrupted and other annulations well marked;
dorsal pores faint; setae not visible. Male pores
transverse slits equatorially in ab of XVIII, on
elliptical porophores. Accessory genital markings
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
279
in 7 specimens (2 from No 5, 2 from Np 2 and 3
from Op 3): unpaired, midventral transverse pads
in intersegments 12/13 (4 Gayndah specimens);
13/14 (7 specimens); 14/15, 15/16 and 16/17, with a
similar pad posteriorly in XVII and a callus-like
pad posterior to and confluent with each male
porophore and crossed near its posterior border by
intersegmental furrow 18/19 (6 specimens). Trans-
verse unpaired equatorial pads illustrated in XVII,
XIX and (partial) in XX by Spencer. Female pores
only slightly anterior of the equator of XIV, almost
contiguous medianly (specimen 98) or well sep-
arated though median to a (specimen 100). Sper-
mathecal pores 2 pairs of stellate apertures, each on
a small rounded tubercle, in 7/8 and 8/9, in ab
approximately; seen to lie nearer a than b in
specimen 1 00 where ventral setal couples are here
visible; (apparently in a lines, Spencer).
Septa 4/5 1 1/12 very thick (12/13 moderately
strongly thickened). Dorsal blood vessel single,
continuous onto the pharynx. Last hearts in XIII
(or XII according to Spencer and in a specimen
from Op 3 examined); hearts in X posteriorly
latero-oesophageal, each arising from the dorsal
vessel and receiving a connective from a thin
supraoesophageal vessel. Commissurals in V-IX
valvular, like the latero-oesophageal hearts, but
differing from the latter in being dorsoventral only
and in giving off a lateral branch shortly before
joining the ventral vessel. Subneural vessel absent.
Racemose, fatbody-like masses present in XIII and
XIV lateral of the dorsal vessel in XIII and XIV
(specimen 98 only). Pharynx ending in IV; oeso-
phagus in V dilated but thinwalled; gizzards, in VI
and VII, large, globose, thin-walled anteriorly, the
musculature thick only in the posterior half; closely
abutting; not separated by unmodified oeso-
phagus. Oesophagus wide, moniliform and very
thin-walled in VIII-XVI; narrow in XVII; lacking
calciferous glands but more vascularized in
Fig. 4: Digaster brunneus Spencer, 1900, Gayndah specimen, BM(NH) 1973.10.98. A, Spermathecal field; B, Male
genital field. For key to abbreviations, see Fig. 9.
280
MEMOIRS OF THE QUEENSLAND MUSEUM
XII-XVI than elsewhere. Intestine originating in
XVIII but not greatly widening until XIX; typh-
losole, caeca and muscular thickening absent.
Nephridia meronephridia: tufts in II, III and IV,
increasing in size posteriorly to very large in IV,
adherent dorsally and laterally to the pharynx
(specimens 98 and 100). The tufts in IV divisible
into a dorsolateral mass discharging by several
ducts directly into the pharynx and by composite
ducts directly into the pharynx in III and a dorsal
mass sending a composite duct forward to the wall
of the buccal cavity in II (specimen 100). Thick
bands of nephridia on the posterior septa of V and
VI show no close association with the gut and from
their peripheral position, near the body wall, may
be exonephric; parietes lacking nephridia in V and
VI except where a few nephridia impinge on the
body wall near the septum in VI. Scattered parietal
and septal exonephric micromeronephridia in VII.
Nephridia in VIII XI (specimen 98) or -XII
(specimen 100) on the posterior septum and
adjacent parietes with ventral aggregations;
thereafter with numerous minute exonephric mic-
romeronephridia scattered on the parietes. Caud-
ally with numerous moderately large astomate
micromeronephridia on each side all of which, in a
segment, discharge into a common transverse duct
which is traceable to the midline below the
intestine, though enteronephry, if present, is not
demonstrable; the medianmost nephridium en-
larged as a stomate megameronephridium with
preseptal funnel (specimen 100). Metandric (sperm
funnels iridescent in XI); seminal vesicles racemose,
in XII only. Metagynous, bushy ovaries, and
funnels, in XIII; ovisacs not observable. Prostates
racemose, not bipartite, restricted to XVIII the
medianly directed duct weakly developed so that,
in specimen 100 especially, the gland is almost
sessile on the body wall. Penial setae absent.
Spermathecae 2 uniform pairs, each with an
elongate ovoid ampulla reflexed on or in line with
the externally poorly differentiated slightly shorter,
thicker walled duct; the duct bearing a rounded
multiloculate diverticulum dorsolaterally near its
ectal extremity (A very small double diverticulum
reported by Spencer). Length of right spermatheca
of IX = 4- 0-6- 2 mm; ratio length spermatheca:
length duct = 2- 5-2-8; ratio length spermatheca:
length diverticulum = 5-3 5-9.
Type-Locality: Gayndah (Np 2).
Remarks: An extensive search of the Gayndah
area, the type-locality, has yielded only one species
which could be identified with D. brunneus, though,
owing to the brevity of the type-description and
loss of the types (Jenz and Smith, 1969), the
identification cannot be entirely certain. Spencer
did not mention accessory genital markings but
transverse pads are illustrated on the clitellum, on
which they occur in the new material from
Gayndah. The latter material agrees closely with
Spencer’s account in other respects including
metandry which is known elsewhere in the genus
only in the allopatric D. longmani. The distribution
of the genital marking noted by Spencer, segmental
in XVII, XIX and XX, differs from that in the new
material in which markings are not present in XIX
and XX and in which those anterior to the one in
XVII are intersegmental but the fact that Spencer
did not mention the markings in the text suggests
that they were not closely observed and might have
been overlooked anteriorly to XVII. Transverse
genital markings are absent from material from
Murgon (Nq 1), Tansey (Op2) and Wolvi (Oq 2)
which is here identified with D. brunneus on general
anatomy and this variation suggests that absence of
markings anteriorly to XVII in the types, if real,
would not necessarily exclude the new Gayndah
material from the species. The specimens lacking
transverse markings, although listed in ‘Material
Examined’ above, have been excluded from the
specific description pending further elucidation of
their status. All are metandric with dorsal pores in
9/10-10/11, as in the new Gayndah material; hearts
are in XII (Wolvi) or XIII (Tansey); and the
spermathecal ducts are narrow, not dilated.
Lengths vary from 1 52 250 mm, and widths from
6-9 mm.
The specimens from Bauple (with transverse
pads) have a suggestion of a genital marking on
each side of the transverse pad in XVII and the
male porophore has a callus-like extension into
XVII and XIX. This extension supports identity
with the Wolvi material which is illustrated in Fig.
3B.
Identity of the new material listed above with the
similarly metandric D. longmani is ruled out on the
grounds that in D. longmani the transverse genital
markings are preclitellar and the first dorsal pore is
more anterior.
Digaster gwongorellae Jamieson, 1972
Figs. 1; 9E
Digaster g\vongorellae Jamieson, 1972, pp. 261^4, figs.
1A-C.
Material Examined: Ps 4, 153°11'E. 28°2LS., Binna
Burra, Lamington National Park, in rainforest, B.
Jamieson and E. Bradbury, 24 Mar 1971. QM G8332,
BM(NH) 1973.10.106.
1 > 1 10 and 75 mm (specimens 1 and 2, posterior
amputees) w (midclitellar) = 5 mm, s = ?
JAMIESON: GENUS DIGASTER IN QUEENSLAND
281
Prostomium tanylobous. First dorsal pore 5/6 faint
but perforate; larger but inconspicuous from 6/7.
Clitellum annular, \ XIII-|XIX but circumscribed
by deep furrows at 13/14 and 18/19 (extent
XIV-X-VIII in syntypes). Setae in 8 regular longitu-
dinal rows. Setae a and b absent in XVIII. Male
genital field an almost circular though transversely
somewhat widened flat topped conspicuous pad in
XVIII extending almost to the setal arcs of XVII
and XIX and laterally almost to c lines; no median
porelike marking present on this pad. In both
specimens two minute white points are included
within the margin of the pad at a and b on each side.
That in b is the larger and has been taken
(Jamieson, 1972) to be the male pore but the two
markings are possibly the follicles of the absent
setae a and b for, as noted in the type-description,
the prostate ducts enter the body wall internally in
a lines. The muscular ducts are straight and
converge medianly so that it is possible that the
male pores are shortly median of a lines (specimen
1). In specimen 2 the genital pad is narrower
longitudinally and is depressed medianly so that an
impression of a pair of medianly conjoined papillae
is produced. Female pore moderately conspicuous,
mid ventral and presetal in XIV. Spermathecal
pores 2 inconspicuous pairs, in 7/8 and 8/9 very
slightly lateral of a lines (in a lines in the types).
Last hearts in XII. Gizzards in VI and VII.
Calciferous glands absent. Intestine commencing
in XVIII but pushing septum 17/18 forwards so as
to appear to commence in XVII. Meronephric.
Holandric (sperm funnels iridescent in X and XI);
gymnorchous; seminal vesicles racemose, in IX and
XII. Prostates restricted to but expanding XVIII,
rounded tongue shaped with a deep lateral incision
but (unlike the type-specimens) not bipartite.
Penial setae absent. Metagynous; ovisacs absent.
Spermathecae 2 pairs, in VIII and IX; each with
elongate-ovoid ampulla, a slightly shorter well
demarcated duct and ectally a multiloculate diver-
ticulum as in the types, (specimen 1).
Type-Locality: Gwongorella National Park (Ps
5).
Remarks: The above brief description confirms
identity of this material with the type-series,
agreeing with the type-description in almost all
respects. The greater extent of the clitellum is
attributable to the very mature condition of the
new material and inclusion of the points (male
pores?) at b in XVIII on the protuberant genital
marking and not, as in the holotype, on each side of
it with an encircling rim, is ascribable to greater
elevation of the marking. The very slightly more
lateral position of the spermathecal pores and the
form of the prostate glands in the new material are
very minor differences.
Digaster longmani Boardman, 1932
Digaster longmani Boardman, 1932, pp. 125-7, fig. 1.
Jamieson, 1963, pp. 101-5, fig. 5.
The following account is abstracted from the
type description and that of Jamieson, 1963. Some
features of a specimen from Stradbroke Island,
Queensland, which is tentatively identified with D.
longmani, are appended to the account.
1 -■ 520-1025 mm, w (preclitellar) — 20-25 mm,
s = 309-382. Body (pigmented?) brown; clitellum
darkbrown . Prostomium zygolobous, prolobous or
proepilobous. First dorsal pore 5/6, or sometimes
6/7. Setae 8 per segment, difficult to discern or only
sporadically visible; typically, in the forebody,
aa:ab:bc:cd = 4:1:3: 1-7; dd:u > 0-5; means in the
Kyogle specimens, in IX, cd:ab = 1-6 (L3-1-8);
bc.aa = 0-4 (0-4— 0-5); dd:u = 0-7 (0-7-0-8).
Clitellum annular, } XIII, a XIII, XIV to XVIII , i
XIX-^XIX (= 5-6 segments). Male pores
equatorial or just presetal in XVIII, in ab, from a to
b. Accessory genital markings if present, a post-
setal, unpaired midventral transverse strip extend-
ing laterally to y be in each of segments VIII-XII.
Female pores presetal in XIV, transversely or
diagonally placed. Spermathecal pores 2 pairs, at
anterior borders of VIII and IX or 2 or 3 pairs in 6/7
to 8/9, in or just dorsal of a.
Last hearts in XII or, typically, XIII. Gizzards in
VI and VII; calciferous glands absent (an unpaired,
dorsal oesophageal structure reported as a gland in
the holotype was probably a fixation artefact).
Intestinal origin XVIII. Nephridia meronephridia;
septal bands present but no definite tufts observed;
caudally with a median (stomate?) megamero-
nephridium on each side. Metandric; gymnor-
chous; seminal vesicles in XII only. Prostates
racemose, tongue-shaped or discoidal, incised but
unipartite in XVIII. penial setae absent. Met-
agynous; ovisacs unknown. Spermatheca 2 or 3
pairs, tubular to tapering sacciform with the ectal
portion forming a porrly demarcated duct; diverti-
culum typically single, elongate-ovoid, obliquely
placed near the pore, extending almost across the
flattened side of the duct, containing several sperm
masses; duct also bearing 1 or 2 conspicuous
obliquely placed sacculations at midlength; if 2, on
opposite sides and convergent ectally; the pair of
sacculations the only diverticula in the Kyogle
specimens.
Type-Locality: Tamborine Mountain, South
Queensland (Pr. 9).
282
MEMOIRS OF THE QUEENSLAND MUSEUM
Wider Distribution: near Oaky Creek, Rich-
mond Range State Forest, and near Kyogle, New
South Wales.
Stradbroke Island Specimen
Material Examined: Pr5, 153°25'E. 27°30 / S., Strad-
broke Island, in deep sand, V. Pattemore, 13 Aug 1971,
BM(NH) 1973.10.107.
Postclitellar amputee with w (midclitellar) = 8
mm. Pigmentless in alcohol excepting light brown
clitellum. Prostomium prolobous but peristomium
much grooved. First dorsal pore 5/6. Setae 8 per
segment; dorsal couples varying significantly in
width and ventral couples slightly irregular, in the
forebody; both couples very irregular in width
behind the clitellum. Clitellum annular, dorsally
XIII XX. Male pores transverse slits in a, po-
rophores not developed. Accessory genital mark-
ings very faint transverse pads filling the posterior
annulus, and including the setal arc, in IX, X, XI
and XII. Female pores not certainly recognisable,
possibly unpaired. Spermathecal pores 2 pairs, in
7/8 and 8/9, in a.
Last hearts in XII. Gizzards in VI and VII;
calciferous glands absent. Intestinal origin(?)
Meronephric. Holandric; gymnorchous; seminal
vesicles in XI and XII. Prostates racemose, tongue-
shaped, unipartite. penial setae absent. Met-
agynous. Spermathecae 2 pairs, duct tapering;
?diverticulum single, small and caplike on the ectal
end of the duct.
Remarks: Typical specimens of D. longmani are
distinguished from the only other metandric spec-
ies in the genus D. brunheus, by location of the first
dorsal pore in 5/6 or 6/7 (not 9/10-11/12), location
of transverse accessory genital markings segmen-
tally in the region of VIII-XII (not in-
tersegmentally in 12/13-16/17) and in possessing
an elongate diverticulum sessile over its length or a
pair on each side of the duct (not rounded and
multiloculate or small and double). Conspecificity
of the material from the environs of Kyogle, in New
South Wales, with D. longmani is not entirely
certain as it lacks accessory genital markings and
has spermathecal diverticula restricted to the pair
of convergent sacculations reported, in addition to
a diverticulum, by Boardman but erection of a
separate species for its reception is not warranted
on present evidence. The identity of the single
Stradbroke Island specimen with D. longmani is
based primarily on the large body size and presence
of segmental preclitellar bands which are so faint
that positive recognition of them as accessory
genital markings must await discovery of further
material. The holandric gonads are otherwise un-
known in D. longmani though known in D.
bradburyi bunyaensis, a subspecies of large worms
of closely similar morphology. The Stradbroke
Island specimen can be excluded from D. brad-
buryi, however, because the latter has its first dorsal
pore in 9/10, bipartite prostate glands and dilated
spermathecal ducts. The brunneus-longmani com-
plex and its relationships with D. lumbricoides
kondalilla requires further elucidation.
Digaster lumbricoides Perrier, 1872
Digas ter lumbricoides Perrier, 1872, pp. 94-96, pi. 1, fig.
24, pi. 4, figs. 64, 65. Fletcher, 1887, pp. 559-60;
1889, pp. 1531 2. Beddard, 1895, p. 485. Michaelsen,
1900, p. 197. Jamieson, 1971b, pp. 1303-6, figs. 1,
I-K.
The following account is based on the rede-
scription of the type-specimens by Jamieson, 1971b
and on new material, comprising a new subspecies,
from Kondalilla.
1 = 82 mm, w (midclitellar) = 3-3— 3-5 mm, s =
158. Form moderately stout, circular in cross
section throughout; pigmentless buff in alcohol.
Prostomium prolobous. First dorsal pore 4/5 but
may not be perforate until 5/6 or 6/7. Setae small, in
8 regular longitudinal rows, commencing in II;
setae a and b present or absent on the segment
bearing the male pores; aa:ab:bc:cd:dd in XII
averaging 2-3 3-6: 1:3 6-4 2: 1 -8-3-3: 1 2-6-13-3; dcku
= 0-40-0-48; c and d not especially dorsal pos-
teriorly. Clitellum annular, typically XII-4XVII.
Male pores on XVII or XVIII in ab or b in an
approximately rectangular tumid field which ex-
tends to the equators of the adjacent anterior and
posterior segment, an oval glandular genital mark-
ing located near each corner of the field, posteriorly
in XVI and anteriorly in XVIII (where male pores
are in XVII) or at intersegments 17/18 and 18/19
(where male pores are in XVIII). Female pores
shortly anteromedian of setae a of XIII or XIV.
Spermathecal pores 2 pairs, in 6/7 and 7/8, or 7/8
and 8/9, in a or ab.
Dorsal blood vessel single, continuous onto the
pharynx. Last hearts in XI or XII; the last 3 pairs
latero-oesophageal, each receiving a connective
from the dorsal vessel and from the roof of the
oesophagus, no discrete supra-oesophageal vessel
being recognisable. Subneural vessel absent. Giz-
zards in V and VI or VI and VII, intervening
unmodified oesophagus short or inappreciable;
calciferous glands absent though the oesophagus is
dilated and vascularized in some of the segments
XIII-XVI; intestinal origin \ XVI or XVIII;
typhlosole, muscular thickening and caeca absent.
Nephridia meronephridia, enteronephry not obser-
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
283
ved; caudally with a stomate megameronephridium
on each side median to astomate micromero-
nephridia. Holandric; gymnorchous; seminal ves-
icles racemose in X and XI or XI and XII,
accompanied below the oesophagus in XI or in XI
and XII by an unpaired seminal vesicle. Prostates
racemose unipartite or bipartite. Penial setae
absent. Ovaries in XIII (or XII?); ovisacs unknown,
Spermathecae 2 uniform pairs; duct cylindrical
with small subspherical ectal diverticulum or duct
pyriform, ectally widening, with a small rounded
more or less bifid diverticulum on the ectal
dilatation; length of a spermatheca 2-2— 3-9 mm,
ratio of length spermatheca: length duct = 1-6-2-0;
ratio length spermatheca: length diverticulum =
5-3-7 9.
Type-Locality: Port Macquarie, New South
Wales.
Wider Distribution: Kondalilla National Park,
Queensland (Oq 8).
Digaster lumbricoides lumbricoides Perrier, 1872
Figs. 1; 9F
For detailed account see Jamieson, 1971b.
Characters as for specific description, above, with:
1 = 82 mm, w (midclitellar) = 3-3 mm, s = 158.
In XII, aa:ab:bc:cd:dd averaging 2-3:1 :4-2:l -8: 12-6;
dd:u = 0-40-0-48. Male pores in XVII, in ab; paired
genital markings posteriorly in XVI and anteriorly
in XVIII, in and lateral o( b lines. Spermathecal
pores in 6/7 and 7/8 in ab.
Septa 6/7 or 7/8 the thickest, strongly thickened.
Last hearts in XI. Gizzards in V and VI; intestinal
origin \ XVI. Seminal vesicles in X and XI,
accompanied below the oesophagus in XI by an
unpaired seminal vesicle. Prostate glands uni-
partite. Spermathecae each with a subspherical
ampulla and a wide cylindrical duct; diverticulum
ectal on the duct; length of a spermatheca 2-2 mm,
ratio of lengthdength duct = 1-6; ratio of
lengthdength diverticulum = 5-3.
Type- Locality: Port Macquarie, New South
Wales.
Digaster lumbricoides kondalilla subsp. nov.
Figs. 1; 5A, B; 10H, I; 11D
Material Examined: Oq 8, 150 o 50'E. 26°40'S.,
Kondalilla National Park, in black soil under boulder on
creek bank, in rainforest, T. Walker, 2 Jun 1972,
Holotype, QM G7396.
1 = 60+ mm (posterior amputee at 71st.
segment), w (XV) = 3-5 mm. Peristomium with
several dorsal longitudinal grooves though not
grooved ventrally. First dorsal pore 4/5. Setae
small but distinctly visible; setae a and b persisting
in XVIII in the vicinity of the male pores. Clitellum
not developed. Male pores transverse slits in XVIII
shortly anterior to setae b, each on a low oval
tumescence which bears setae a and b; the male
porophores preceded at 17/18 and succeeded at
18/19 by a pair of oval glandular areas, in ab, each
of the posterior areas accompanied by and partly
continuous with a similar glandular marking
lateral of b lines in the same intersegment. The male
porophores and these genital markings lying in an
approximately tumid quadrangular field which
extends to the setal arcs of segments XVII and
XIX and laterally almost to c lines. Female pores
shortly anteromedian of setae a of XIV on a
common transverse presetal protrusion. Sper-
mathecal pores 2 pairs of partially concealed
transverse slits at the anterior limit of small
knoblike swellings, in 7/8 and 8/9, centred in a lines.
Septa 8/9-11/12, the thickest, strongly thick-
ened. Last hearts in XII; those in XII (and X-XI?)
latero-oesophageal. Pharynx ending in IV; in V the
oesophagus is dilated and gizzard-like in ap-
pearance though not thickwalled and little more
than half the diameter of the gizzards. Gizzards
large and firm, in VI and VII; globose though with
an anterior rim which is weakly developed in the
posterior but is well developed in the anterior
gizzard; the gizzards filling their segments; and not
separated by an appreciable length of unmodified
oesophagus though the oesophagus is narrowly
constricted between them. Oesophagus longitu-
dinally constricted and lacking special modi-
fication in IX-XIII; in each of XIV, XV and
XVI dilated and vascularized, with a pair of
circumferential vessels which join the dorsal vessel,
and internally with many high clavate villi; slender
in XVII and XVIII. Intestinal origin XIX. Neph-
ridia in IV VI aggregated into loose tufts; no
enteronephry demonstrated but certain elucidation
of the excretory system not feasible as preservation
unsatisfactory; in the anterior intestinal region with
seven astomate exonephric parietal micromero-
nephridia on each side from setal line a to above d
and an aggregation of similar but smaller nephridia
median to a\ these median aggregations not present
further posteriorly. No stomate nephridia detect-
able anterior to the amputation. Sperm funnels
iridescent in X and XI; small paired seminal vesicles
racemose in XI and XII, accompanied below the
eosophagus by a large unpaired racemose seminal
vesicle in each segment. Prostate glands large but
restricted to XVIII, racemose and bipartite, the
muscular duct branching entally to the two major
lobes, then the branches in turn showing limited
284
MEMOIRS OF THE QUEENSLAND MUSEUM
Fig. 5: Digaster lumbricoides kondalilla subsp. nov. A, Spermathecal field: B, Male genital field of holotype. For key to
abbreviations, see Fig. 9.
branching; the branching visible when the prostatic
lobes are separated. Penial setae absent, though a
and b follicles are ekternally visible. Ovaries (diffuse
webs with small oocytes) and funnels in XIII.
Spermathecae each with an ovoid-sacciform am-
pulla and an elongate pearshaped, ectally widening
duct of similar length which bears a small rounded
sessile more or less distinctly bifid (iridescent)
diverticulum dorsally on the ectal dilatation; length
of right spermatheca of VIII = 3-9 mm; ratio of
length spermatheca: length duct = 2 0; ratio of
length spermatheca: length diverticulum = 7-9.
Remarks: Digaster lumbricoides, the type-species
of the genus was previously known only from
material collected at Port Macquarie in 1846
(Jamieson, 1971b). Although erection of a new
subspecies on a single specimen is undesirable, it is
preferable to total identification of this distinctive
specimen with D. lumbricoides as represented by
the earlier material. Some differences from the
nominate subspecies are: location of male pores on
XVIII; origin of the intestine in XIX (not \ XVI);
the more strongly bipartite condition of the
prostates and (significantly?) the ectal widening of
the spermathecal ducts; location of the male pores
in b and of spermathecal pores in a lines, rather
than ab, and the intersegmental, not postsetal
location of the genital markings with lateral
duplication of the posterior markings. These are
minor differences of doubtful importance.
Dilatation of the spermathecal ducts and the
bipartite form of the prostate glands are shared
with D. bradburyi with which affinities are close. On
the other hand, the quadrangular raised genital
field with genital markings on the corners, post-
testicular paired and median seminal vesicles and
anterior dorsal pores strongly indicate
conspecificity with D. lumbricoides . It is possible
that D. bradburyi consists of populations of the
earlier named D. lumbricoides but elucidation of
the lumbricoides-bradburyi complex and of re-
lationships of these with D. brunneus and D.
longmani requires further studies of morphology
and reproductive isolation of the constituent
populations.
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
285
Digaster minima sp. nov.
Figs. 6A; 10J; Table 3
Material Examined: Or 7, 152°47'E. 27°23'S., Mt.
Nebo Road, Mt. Nebo, Queensland, in loamy soil in gully
in eucalypt area, E. Bradbury, 12 Jun 1970, Holotype (H)
QM G7397, Paratypes (P) 2, 3, QM G7398 9. PI,
BM(NH) 1973.10.108. 1 mile from Mt. Nebo on road, E.
Bradbury, 10 Aug 1970, P4, BM(NH) 1973.10.109.
1 = ? (H, posterior amputee), 28 (PI), 41 (P2)
mm, w (midclitellar) = 1- 7 mm, s = 122 (P2). Form
slender, circular in cross section; pigmentless in
alcohol. Prostomium proepilobous, slightly (PI, 2)
to f peristomium (H), peristomium with or without
slight dorsal furrows. First dorsal pore 9/10 (FI, PI ,
2) with possibly some imperforate preceding this.
Setae visible with difficulty anteriorly where they lie
in 8 longitudinal rows, commencing on II; shortly
behind the clitellum they are recognisable only with
the greatest difficulty until in the last approxi-
mately 30 caudal segments where the dorsal pair
on each side is enlarged and is shifted dorsally so
that the 4 setae cd are approximately equidistant
from each other; c and d are also displaced dorsally
in anterior segments though not so markedly; setae
a and b absent in XVIII. Clitellum annular,
XIV fXVIII; (H, P2) setae visible; dorsal pores
obliterated; intersegmental pores represented only
ventrally. Male pores in longitudinal crescentic
grooves equatorially in ab of XVIII on broad, low,
medianly widely conjoined (H) or just contiguous
porophores (PI, 2) which almost reach the setal
arcs of XVII and XIX or (PI, 2) are less extensive.
Accessory genital markings midventral unpaired
circular segmental papillae with porelike centres in
each of segments XII and XIII and XX-XXIV (H).
A paired marking sometimes present in the setal arc
and bearing setae ab on its posterior aspect in X
(PI, unilateral left only; P2, 4 bilateral). Median
marking in XII (PI, 3, 4), XIII (Pl-4); XV (P2, 4);
XVI (P4); XIX (P2, 3); XX and XXI (Pl-4) and
XXII (P2, 3). Female pores a pair shortly ante-
romedian of setae a of XIV. Spermathecal pores 2
pairs, near the anterior margins of VIII and IX in
ab , nearer a (H) or in a lines, on small rounded
papillae.
Septa 6/7 and 7/8 the strongest, those to 10/11
moderately strong. Dorsal blood vessel single,
continuous ontojfie pharynx. Last hearts in XII.
Pharynx ending in IV; oesophagus wide and
thinwalled in V; gizzards well developed, globose,
not separated by unmodified oesophagus but a
narrow anterior part of the posterior gizzard
relatively unthickened. Calciferous glands absent.
Origin of intestine indefinite, apparently \ XVII in
PI; caeca and muscular thickening absent; definite
typhlosole absent but a very low dorsal ridge
present from approximately XXIV posteriorly.
Meronephric: no tufts present; nephridia few
micromeronephridia; in the anterior intestinal
region 3 on each side in longitudinal rows, one row
in ab, the second at mid be, the third in cd.
Metandric; sperm funnels strongly iridescent in XI;
gymnorchous; seminal vesicles racemose in XII (H,
PI) a spermatheca-like iridescent sac (accessory
seminal vesicle?) extends below the gut from
septum 12/13 into XIII (H only). Metagynous;
ovaries flattened laminae with numerous oocytes in
XIII; ovisacs absent. Prostates racemose, squarish
bifid lobes, occupying about four segments; vas
deferens joining the muscular anteromedianly
directed duct at its junction with the gland. Penial
setae absent. Spermathecae 2 uniform pairs, each
with ovoid ampulla and well demarcated fairly
slender duct; a single (inseminated) subspherical
diverticulum joining the duct by a narrow very
short stalk at midlength; the diverticulum exter-
nally simple but with several internal sperm masses;
length of right sperma theca of IX (H) = 1-7 mm;
ratio of length spermatheca: length duct = 1-8;
ratio of length spermatheca: length diverticulum =
5-6.
Remarks: Only D. pseudoperichaeta shares with
D. minima the extreme dorsal displacement of the
dorsal setal couples in caudal segments. Both have
the metandric condition of the male gonads seen
elsewhere in the genus only in D. brunneus and D.
longmani. It appears likely that the two species are
more closely related one to the other than to other
species. They are clearly interdistinguishable by the
male genital fields.
D. minima is the smallest known species of
Digaster.
Digaster nothofagi sp. nov.
Figs. 1; 6B; 10K; 1 IF; Table 3
Material Examined: Ps 5, 153°17'E. 28°13 , S.,
Springbrook, Best of All Lookout, under Nothofagus
(Antarctic beeches), B. Jamieson and E. Bradbury, 22
Apr 1971, Holotype, QM G7400.
1 = 81 mm, w (midclitellar) - 5-2 mm, s = 196.
Form moderately stout, circular in cross section;
pigmentless buff in alcohol. Prostomium epilobous
f open, dorsally bisected by a longitudinal groove.
Peristomium with numerous faint longitudinal
grooves but not bisected. First dorsal pore 4/5.
Setae small but visible, in 8 regular longitudinal
rows throughout, commencing on II. Setae a and b
absent.
Nephropores not externally recognisable. Clitel-
lum hardly appreciable externally but seen from the
286
MEMOIRS OF THE QUEENSLAND MUSEUM
dorsal incision to extend from XIV XVII with
some very weak development in the adjacent halves
of XIII and XVIII; apparently annular. Male pores
minute, in ab of XVIII, on small papillae. Ac-
cessory genital markings: an unpaired midventral
papilla posteriorly in IX, X and XVII and in 18/19,
19/20 and 20/21, all well within aa excepting that in
IX which extends laterally into ab ; paired papillae
postsetal in ab in XVII and lateral to b in XVIII;
segments VII and VIII with an indistinct tumid
band postsetally connecting the spermathecal
pores. Female pore unpaired, presetal in XIV.
Spermathecal pores 2 pairs in 7/8 and 8/9,
immediately lateral to a lines on minute papillae.
Septa 6/7-10/11 strongly thickened. Dorsal
blood vessel single, continuous onto the pharynx.
Last hearts in XII; those in X-XII latero-
oesophageal, taking their main origin from the
supra-oesophageal vessel but with very slender
connectives from the dorsal vessel. Supra-
Fig. 6: A, Digaster minima sp. nov. Genital field of holotype.B, D. nothofagi sp. nov. Genital field of holotype. For
key to abbreviations, see Fig. 9.
JAMIESON: GENUS DIGASTER IN QUEENSLAND
287
oesophageal ill-defined, limits indeterminable.
Subneural vessel absent. Oesophagus in V slightly
dilated, thin-walled, hidden by septal glands. Two
strong subspherical gizzards with slightly develop-
ed anterior rims, in VI and VII, the oesophagus
between them strongly constricted but very short.
Oesophagus almost suppressed, by backward
extension of the gizzards, in VIII and IX; short and
simple in X XIII; swollen and vascularized in
XIV-XVII, where a paired circumferential vessel
joins the dorsal vessel, with low, sinuous internal
rugae especially dilated in XV and XVI; oeso-
phagus narrow in XVIII. Intestinal origin XIX;
typhlosole, caeca and muscular thickening absent.
Nephridia meronephridia: masses of spiral loops in
IV, V, VI and VII, on the posterior septa of these
segments, those in V-VII apparently but not
certainly exonephric, those in IV with connections
to the pharynx which may be ducts. Sparse lateral
parietal bands of exonephric micromeronephridia
in II and III and in VIII posteriorly; in the anterior
intestinal region 8 or 9 on each side and a small
group median to these; caudally with the median-
most nephridium enlarged as an exonephric mega-
meronephridium with preseptal funnel. Holan-
dric, gymnorchous (sperm funnels iridescent in X
and XI); seminal vesicles racemose, in IX and XII.
Prostates racemose, tongue-shaped, each entally
divisible into a dorsal and a ventral lobe; vas
deferens joining the straight medianly directed
muscular duct near its junction with the gland.
Metagynous (ovaries not seen; oviducal funnels in
XIII); ovisacs absent. Spermathecae 2 uniform
pairs; ampulla elongate ovoid; duct cylindrical
bearing an iridescent multiloculate diverticulum
near its ectal extremity; length of right spermatheca
of IX = 2-6 mm; ratio of length sperma-
theca: length duct = 2-6; ratio of length sper-
matheca: length diverticulum = 41.
Remarks: D. nothofagi, from a spur on the
escarpment of the Macpherson Range, closely
resembles D. binnaburra from neighbouring parts
of the same range. The two taxa are clearly
distinguished by the genital fields which are
sufficiently different to suggest reproductive iso-
lation, one from the other.
Digaster pseudoperichaeta sp. nov.
Figs. 1; 7; 10L; 11G; Table 3
Material Examined: Or 4, 152°59'E. 27°16'S.,
Y.M.C.A. Camp Warrawee, near Petrie, in sandy loam
under Catospermum in riverine vine forest, B. Jamieson,
17 Feb 1969, Holotype (H) QM G7401, Paratypes (P) 2, 3
QM G7402-3, PI, BM(NH) 1973.10.110.
Fig. 7: Digaster pseudoperichaeta sp. nov. Genital field
of holotype. For key to abbreviations, see Fig. 9.
1 = 36 (PI )— 45 mm(H), w (midclitellar) = 2-7
(PI) 3-3(H) mm, s = ? (not countable owing to
maceration). Form circular in cross section, mod-
erately slender. Pigmentless in alcohol. Pro-
stomium epilobous t, open, faintly canaliculate.
Dorsal pores minute, commencement indetermin-
able. Setae commencing on II, in 8 regular
longitudinal rows throughout but the dorsal coup-
les (cd) caudally moving to a dorsal situation so
that by approximately 15 (PI) or 25(H) segments
from the posterior end the four setae of the two
dorsal couples are equispaced, and are large and
readily seen, whereas the more anterior setae, and
all ventral couples, are minute and inconspicuous.
Setae a and b absent in XVIII. Clitellum annular,
XIV ^XVII well developed and protuberant;
setae visible, dorsal pores occluded; intersegmental
furrows almost obscured. Male pores a pair of
transverse slits in ab on low whitish porophores
which fill the segment longitudinally. Accessory
genital markings: a midventral transverse, strongly
protuberant pad occupying the anterior two thirds
of X and extending laterally beyond b; a further
288
MEMOIRS OF THE QUEENSLAND MUSEUM
midventral pad anterior to but overlapping the
setal arc of XVI and XVII, anteriorly impinging
very slightly on the preceding segment, and extend-
ing laterally to a and ab respectively; and a pair of
white tumescences on XIX anterior to the ventral
setal couples which latter are themselves on a white
glandular transverse ridge on each side (H, PI -3).
Spermathecal pores 2 pairs, near the anterior
borders of VIII and IX, in ab on circular pro-
minences:
Septa 6/7 and 7/8 the thickest, strongly thick-
ened. Dorsal blood vessel single, continuous onto
the pharynx. Last hearts in XII (vascular system
macerated).
Oesophagus wide and very thin walled in V; two
relatively very large, strong, globular gizzards, in
VI and VII, separated by a deep constriction where
the oesophagus is unthickened though of negligible
length. Oesophagus suppressed by backward ex-
tension of the gizzards in VIII; moniliform and
vascularized but lacking calciferous glands, in
IX-XII; narrowly tubular and sinuous in
XIII-XVII; intestinal origin XV111; typhlosole,
caeca and muscular thickening absent. Mero-
nephric, with a few rows of fairly large mero-
nephridia on each side in the oesophageal and
intestinal regions of which the medianmost neph-
ridium in caudal segments has a preseptal funnel
but is not appreciably enlarged; nephridia in
pharyngeal and buccal segments more numerous
but no tufts detectable. Metandric (sperm funnels,
iridescent, XI); gymnorchous; seminal vesicles
racemose in XII only. Metagynous (ovaries with
several chains of large oocytes); ovisacs absent.
Prostates racemose, tongue-shaped but entally
bifid; restricted to XVIII; vas deferens joining the
junction of the gland with the short muscular duct.
Penial setae absent. Spermathecae 2 uniform pairs,
ampulla very slender and elongate, almost tubular,
lacking a distinct duct though the widened ectal
fourth may be considered one; diverticulum (in-
seminated) consisting of several distinct loculi,
sessile near its ectal limit; length of right sper-
matheca of IX (H) = 2-27 mm; ratio of length
spermatheca: length diverticulum = 4- 1 .
Remarks: D. pseudoperichaeta is closest to D.
minima from which it is distinguished on p. 285.
Digaster sexpunctata sp. nov.
Figs. 1; 8; 10M; 11H; Table 3.
Material Examined: Ps 6, 153°27'E. 28°05'S., 2
miles along Austinville Road, near Burleigh, South
Queensland, E. Bradbury, 10 Apr 1970, Holotype (H)
QM G7404, Paratype (P) BM(NH) 1973.10.111.
1 = 55 mm, w (midclitellar) = 5 mm, s = 142 (H;
paratype is posterior amputee). Form circular in
cross section throughout, moderately stout. Pig-
mentless. Prostomium indistinctly tanylobous,
faintly canaliculate, but almost indistinguishable
from other longitudinal furrowing of the peri-
stomium. First dorsal pore 4/5 (H) or 5/6 (P).
Setae in 8 regular longitudinal rows, commencing
on II; setae a and b absent in XVIII. Nephropores
not externally visible. Clitellum (developed in H
only, though not fully) annular, XlV-fXVIII;
intersegmental furrows and dorsal pores present
though weaker, setae present. Male pores in
Fig. 8: Digaster sexpunctata sp. nov. Genital field of
holotype. For key to abbreviations, see Fig. 9.
JAMIESON: GENUS DIGASTER IN QUEENSLAND
289
Fig. 9: Genital fields of: A, Digaster bradburyi bradburyv, B, D. anomala\ C, D. armifera; D, D. minor, E, D.
gwongorellae; F, D. lumbricoides lumbricoides; G, D. gayndahensis ; H, D. lamingtonensis. (A and B from Jamieson,
1970; C, G and H from Jamieson, 1963; D from Spencer, 1900; E from Jamieson, 1972; F from Jamieson, 1971).
ac.g.m, accessory genital marking; clit, clitellum; 9, female pore; g.m, accessory genital marking; gl. p, glandular
pad; c?, male pore; prd. sue, paired suckerlike genital marking; sp.p, spermathecal pore; sue, suckerlike genital
marking; uprd. sue, unpaired suckerlike genital marking.
290
MEMOIRS OF THE QUEENSLAND MUSEUM
XVIII in ab near b on hemispheroidal papillae
surrounded by a common prominent rim which
extends at least to the setal arcs of XVII and XIX
and laterally to mid be; the male porophores
separated by a pair of very small disc-like
equatorial genital markings which are preceded
and succeeded by a similar pair of markings which
are intersegmental in 17/18 and 18/19 (H, P);
additional genital markings on a transverse pad in
each of intersegments 6/7, 7/8, 8/9 (H, P) and 9/10
(H); those in 6/7 and 9/10 filling bb (H) or that in
6/7 median to a (P); those in 7/8 and 8/9 filling the
interval between the spermathecal pores. Female
pores anteromedian to setae a of XIV, about %aa
apart, on a common elliptical tumescence. Sper-
mathecal pores 2 pairs, at the anterior borders of
VIII and IX, on small papillae (H, P).
Septa 10/11 and 11/12 the thickest, strongly
thickened. Dorsal blood vessel single, continuous
onto the pharynx. Last hearts in XII; those in
X-XII latero-oesophageal, each receiving a con-
nective from the dorsal vessel and the weakly
Fig. 10: Spermathecae in Digaster : A, B, D. binnaburra sp. nov., holo-
type. A, right VIII, B, right IX; C, D, D. bradburyi bunyciensis sub
sp. nov., holotype, C, left VIII, D, right IX; E-G, D. brunneus Spencer,
1900, right IX, E, BM(NH) 1973.10.98; F, BM(NH) 1973.10.100, G,
Wolvi specimen, BM(NH) 1973.10.105; H, I, D. lumbricoides
kondalilla subsp. nov., holotype, right VIII, H, dorsal, I, ventral; J, D.
minima sp. nov., holotype, right IX; K, D. nothofagi sp. nov.,
holotype, right IX; L, D. pseudoperichaeta sp. nov., holotype, right
IX; M, D. sexpunctata sp. nov., holotype, right IX.
JAMIESON: GENUS DIG ASTER IN QUEENSLAND
291
developed supra-oesophageal vessel. Subneural
vessel absent. Pharynx ending in III; oesophagus in
IV and V segmentally dilated and very thin walled.
Two strong subspherical gizzards, in VI and VII,
separated by a short region of unmodified oeso-
phagus. Oesophagus almost suppressed in VIII by
backward extension of the gizzards; moniliform
and vascular in IX-XIII; in XIV, XV and XVI with
longitudinally striated lateral outpouching but not
cut off" from the oesophageal lumen and with only
low internal rugae, not forming definite calciferous
glands. Oesophagus narrow in XVII; intestinal
origin XVIII; muscular thickening, caeca and
typhlosole absent. Nephridia meronephridia (H,
P); large tufts in IV, V and VI sending ducts to a
common composite duct on each side which joins
the anterior aspect of the buccal cavity dorso-
laterally; II, III and VII posteriorly, with parietal
micromeronephridia which form dense lateral
bands in VII; approximately 10 on each side by
the anterior intestinal region. Posteriorly with
moderately enlarged exonephric megamero-
nephridium, with preseptal funnel, median to
astomate, exonephric micromeronephridia (H).
Holandric (iridescent sperm funnels in X and XI);
gymnorchous; large racemose seminal vesicles in
IX and XII. Metagynous, ovaries with numerous
united strings of large oocytes (H, P); small
ovisacs (?) in XIV in H, not observable in P.
Prostates racemose, tongue-shaped, restricted to
but enlarging XVIII; vas deferens joining the
straight medianly directed duct where this joins the
gland. Penial setae absent. Spermathecae with
widely ovoid to subspherical ampulla; the mod-
Fig. 1 1: Prostates in Digaster (L = left; R = right). A, D. binnaburra sp.
nov., holotype, R; B, D. bradburyi bunyaensis subsp. nov., holotype,
L; C, D. brunneus Spencer, 1900, BM(NH) 1973.10.98; L, D.
lumbricoides kondalilla subsp. nov., holotype, R; E, D. minima sp.
nov., holotype, L; F, D. nothofagi sp. nov., holotype, L; G, D.
pseudoperichaeta sp. nov., holotype, R; H, D. sexpunctata sp. nov.,
holotype, R.
292
MEMOIRS OF THE QUEENSLAND MUSEUM
erately narrow, well demarcated duct joined at or
shortly ectal of midlength by a subspherical
internally multiloculate (inseminated) broadly
sessile diverticulum; the ampulla packed with
nematodes (H, P); length of right spermatheca of
IX (H) = 1-5 mm; ratio of length: length duct =
1-8 ratio of length: length diverticulum = 3 0.
Remarks: The spermathecal and male genital
fields of this species are highly distinctive.
ACKNOWLEDGMENTS
I am deeply indebted to Mr R. W. Sims, the
authorities of the British Museum (Natural His-
tory), and Professor R. P. Dales of Bedford
College, for providing facilities for this study. Mr
E. Bradbury and Mr T. Walker are thanked for
their assiduous collecting and Mr Walker for aid in
mapping. The work was made possible by Royal
Society Nuffield and Australian Research Grants
Committee grants.
LITERATURE CITED
Bage, F., 1910. Contributions to our knowledge of
Australian Earthworms. The Nephridia. Proc. R.
Soc. Viet, (n.s.) 22 (2): 224-43.
Beddard, F. E., 1895. 'A monograph of the Order
Oligochaeta.’ (Clarendon Press: Oxford).
Boardman, W., 1932. Some earthworms from Queens-
land. Mem. Qd Mus. 10 (2): 125 30.
Fletcher, J. J., 1887, Notes on Australian earthworms.
Part I. Proc. Linn. Soc. N.S. W. 1: 523-74.
1 888. Notes on Australian earthworms. Part III. Proc.
Linn. Soc. N.S.W. (2) 2: 375402.
1889. Notes on Australian earthworms. Part V. Proc.
Linn. Soc. N.S.W. (2) 3: 1521 58.
Jamieson, B. G. M., 1963. A revision of the earthworm
genus Digaster (Megascolecidae, Oligochaeta). Rec.
Aust. Mus. 26 (2): 83-111.
1970. Two new sympatric species of the earthworm
genus Digaster (Megascolecidae: Oligochaeta) from
Queensland. Proc. R. Soc. Qd. 82 (3): 35-46.
1971a. A review of the megascolecoid earthworm
genera (Oligochaeta) of Australia. Part III — The
subfamily Megascolecinae. Mem. Qd Mus. 16 (1):
69-102.
1971b. Descriptions of the type-species of the
earthworm genera Plutella and Digaster (Mega-
scolecidae : Oligochaeta). Bull. Mus. natn. Hist, nat.,
Paris (2) 42 (6): 130040.
1972. A new species of Digaster (Megascolecidae:
Oligochaeta) from Queensland. Mem. Qd Mus. 16
(2): 261-4.
1974. The zoogeography and evolution of Tasmanian
Oligochaeta. In ‘Biogeography and Ecology in
Tasmania.’ Ed. W. Williams. (Junk: Den Hague).
Jamieson, B. G. M. and Bradbury, E. A., 1972.
Investigation of the Australian earthworm genus
Didymogaster (Megascolecidae: Oligochaeta) and
discrimination of populations by analysis of setal
ratios. Pedobioiogia 12: 1 1 1-22.
Jensz, R. L. and Smith, B. J., 1969. Catalogue of
Baldwin Spencer earthworm types in the National
Museum of Victoria, Australia. Mem. natn. Mus.
Viet. 29: 85-110.
Kikkawa, J. and Pearse, K., 1969. Geographical
distribution of land birds in Australia — a numerical
analysis. Aust. J. Zool. 17: 821-40.
Michaelsen, W., 1900. ‘Das Tierreich,’ Vol. 10, Vermes,
Oligochaeta. (Friedlander: Berlin).
1907. Oligochaeta In ‘Die Fauna Siidwest-Australiens.’
Bd 1. Lief. 2, pp. 117-232. (Gustav Fischer: Jena).
1916. Results of Dr E. Mjobergs Swedish Scientific
Expeditions to Australia 1910-1913. Oligochaeten.
K. svenska VetenskAkad. Handl. 52(13): 3-74.
Perrier, E., 1872. Recherches pour servir a l’histoire
des lombriciens terrestres. Nouv. Arch. Mus. Hist,
nat., Paris 8: 19-197.
Spencer, W, B., 1900. Further descriptions of Australian
earthworms, part I. Proc. R. Soc. Viet, (n.s.) 13 (1):
29-67.
Stephenson, J., 1930. ‘The Oligochaeta’. (Clarendon
Press: Oxford).
Sweet, G., 1900. On the structure of the spermiducal
glands and associated parts in Australian
earthworms. J. Linn. Soc., Zool. 28 (180): 109-39.
Mem. Qd Mus. 17(2): 293-303, pis. 36-40. [1975]
A REVIEW OF THE GENUS PHYLLURUS (LACERTILIA: GEKKONIDAE)
Jeanette Covacevich
Queensland Museum
ABSTRACT
Two new species of Phyllurus ( P. caudiannulatus and P. salebrosus) are described from mid-
eastern Queensland and the other two species of this genus, which is confined to eastern Australia,
are redescribed. A key to their identification is provided and distribution of the species is
discussed.
The genus Phyllurus Schinz, 1822, is defined in
detail by Kluge (1967, p. 1017) to include four
species — P. platurus (White, 1790), P. milii (Bory
de St. Vincent, 1825), P. cornutus (Ogilby, 1892),
and P. sphyrurus (Ogilby, 1892). Two distinct
groups are recognisable within the genus as it is
defined by Kluge. The first comprises P. milii and
P. sphyrurus. These deep-headed species, covered
in rounded tubercles, with robust moderately
depressed tails, are referred to Underwoodisaurus
by Wermuth (1965, p. 47). Species of the second
group (P. platurus and P. cornutus) have extremely
depressed heads, very broad, depressed, ‘leaf-like’
tails, and are covered in conical tubercles.
Two new species described here are referred to
Phyllurus (sensu stricto) because both have ex-
tremely depressed heads and are covered with
conical tubercles. Only one species (P. salebrosus
sp. nov.) has the characteristic ‘leaf-like’ tail of
Phyllurus species. The other (P. caudiannulatus),
has a tail that is long, slender, and round in cross-
section or slightly leaf-shaped. As P. caudiannula-
tus is very similar to P. platurus (the only obvious
differences are in tail shape and distribution) and as
U. sphyrurus and V. milii form an easily recognised
separate group, Kluge’s definition of the genus
Phyllurus must be modified to exclude P. sphyrurus
and P. milii (following Wermuth, 1965) and to
include P. caudiannulatus. This can be done by
describing the tail of Phyllurus as moderately long,
cylindrical or extremely depressed, not prehensile,
without modified terminal subcaudal lamellae.
The four species here referred to Phyllurus are
confined to eastern Australia between mid-eastern
New South Wales and northeastern Queensland.
Only one species (P. salebrosus) is found far from
the coast. Two species (P. cornutus, P. caudian-
nulatus) are apparently usually confined to closed
forests and the other two species (P. platurus and P.
salebrosus) inhabit drier rocky areas with sparser
vegetation cover.
Methods
All specimens of Phyllurus in the reference
collections of the Queensland Museum (QM), the
Australian Museum (AM), and the National Parks
Branch (Department of Forestry) Museum (RG)
have been examined. The following body measure-
ments and morphological characters have been
used in separating the species.
Snouth to vent length (SVL): From tip of snout
to anterior margin of vent.
Tail length (T); Taken in two ways (a) in species
with ‘leaf-like’ tails, because almost all tails have
been cast posterior to the vent where the leaf
begins, as shown in Fig. 1; (b) in P. caudiannulatus
which has a ‘conventional’ tail, from posterior
margin of cloaca to tip of tail.
Attenuated tip of tail (TT): As shown in Fig. 1 .
Head length (HL): Between tip of snout and
posterior margin of ear.
Head width (HW): Greatest width of head.
Snout (S): Tip of snout to anterior margin of eye.
Labials: Counted to exclude granular scales
towards angle of mouth.
Number of rows of spines across attenuated
portion of tail: Counted from first complete row
across the tail, usually corresponding with the
posterior end of the ‘leaf (see Fig. la).
Rows of tubercles on eyelid: Counted to exclude
the row forming the ‘eyelash’ and the orbital row
(see Fig. 2).
294
MEMOIRS OF THE QUEENSLAND MUSEUM
Key to Phyllurus Species
1 Throat completely covered with calcareous
deposits . . . . . P. salebrosus
Throat almost smooth, calcareous deposits
only in skin covering bones of lower jaw
P. eornutus
Throat and skin covering bones of lower jaw
smooth 2
2(1) Specimen with original tail . . . . 3
Specimen with regenerated tail or no tail 4
3(2) Tail with distinct white bands, cylindrical or
slightly leaf-shaped . . P. caudiannulatus
Tail without distinct white bands, leaf-
shaped . . . . . . . . P. platurus
4(2) Specimen from mid-eastern Queensland
. . P. caudiannulatus
Specimen from mid-eastern New South
Wales P. platurus
Phyllurus platurus (White)
(Figs. 3, 4; Plates 37B, 38C, 39C, 40D)
Lacerta platura White, 1790, p. 246, pi. 32, fig. 2.
{Australia — presumed to be near Sydney, N.S.W.;
?holotype BM xxii98a).
Agama discosura Merrem, 1820, p. 51. (Australia;
type(s) presumed lost).
\}\Phyllurus australis Swainson, 1839, p. 370.
Phyllurus inermis Gray, 1845, p. 176. (Australia;
holotype BM xxiilOOa).
Material Examined
QM Blue Mts„ J160; Buladelah J9054; AM no data,
5241, A1237, R959, R966, R992, R1550, R1575, R3588,
R3601, R3666, R5182, R11587; Bondi, R1124; Kiama,
R2306; Balmain, R2531; Hawkesbury, R3143; Woolwich,
R3182; Brooklyn, R3392; Sackville, near Windsor,
R3582; near Sydney, R3583, R8305; Callan Park, R3585;
Greenwich, Lane Cove R., R3793; Darling Point, R4396;
North Sydney, R4404, R 12209; Gosford, R4814;
Bradley’s Head, R5181; Sydney, R5520, R8087; Linfield,
R6141; Terrace Falls, Hazelbrook, R6728; Freshwater,
Manly, R7087, R7189, R8036; Pennant Hills, R7294;
Double Bay, Sydney, R7747; Woodford, Blue Mountains
R7987; Coalcliffe, R8037; Mittagong, R8125, R8126;
Hunters Hill, R8271; Mosman, near Sydney, R8277,
R8918; North Sydney, R8595; Lithgow, R8980; Crem-
ome, R9274; Northbridge, R9826; Epping, R1005,
R 10377; Lane Cove R., R 10066; Annandale, R 10068;
Watson’s Bay, R10220; Croydon Park, R10374; near
Gosford, R 10384; Harbord, R10387; Sydney, R10412,
R21047; Tuggerah Lakes, R 10429; Edgecliffe, R 10504;
Longueville, R1 1733; Northbridge, R1 1889; Northmead,
R 12907; Punchbowl, R13105; Giruan, via Stroud,
R 154 12; Jannali, Sydney, R 19084; Jenolan Caves,
R20381; Mangrove Creek, via Spencer, R25891, R25912;
Saratoga, R26208; Miranda, Sydney, R27324; Faulcon-
bridge, R27325; St. Ives, Sydney, R27330; Castle Hill,
Sydney, R27334; Hunters Hill, Sydney, R27940; Baulk-
ham Hills, Sydney, R28308; North Shore, Sydney, 4942,
All 70 1 ; Double Bay, Sydney, A9615; Wallis Lake,
Tuncurry, R8253; Buladelah, R8103. All localities are in
New South Wales.
Diagnosis
A small leaf-tailed Phyllurus very similar to P.
caudiannulatus from which it may be distinguished
readily by tail shape (depressed, broad, leaf-like
original and regenerated tail vs conventional cy-
lindrical original and regenerated tail*). Specimens
without tails virtually indistinguishable although
P. caudiannulatus is usually more spinose, es-
pecially on the head, than P. platurus and the two
species occur in widely separated localities (mid-
eastern N.S. W. vs mideastern Queensland). Distin-
guished from P. eornutus from southeastern
Queensland and northeastern New South Wales,
with which it has been confused, by size (SV 95-9
(max) vs 140 0), absence of calcareous deposits on
throat (present on skin covering bones of lower jaw
in P. eornutus ) and, less reliably, tubercles at flank
(very small vs larger, rounded or flattened and
strongly hooked in northern specimens).
Description
White’s type description of this species is very
brief and, although it is accompanied by an illustra-
tion, is hardly adequate for distinguishing the
species. J. E. Gray (1845, p. 176) examined two
specimens he referred to P. platurus from ’New
Holland’ (= Australia). G. R. Gray (1845, p. 17,
fig. 2) referred material from ’Sydney’ and ’Mac-
quarie River’ to P. platurus. His illustration shows
typical specimens of P. platurus with original and
regenerated tails (as P. inermis). It seems reasonable
to assume that White’s type specimen was included
in those examined by Gray and, in any case, the
specimen on which the type description is based
almost certainly came from near Sydney because in
1790 there were no other settlements in New South
Wales. The only other species of Phyllurus which
occurs in Sydney (and this is based on a slightly
doubtful record) is P. salebrosus sp. nov,, a very
large species which has the attenuated portion of
the tail less than ^ total length of the tail. White’s
type specimen measured 44" (114-3 mm) and his
illustration is of a specimen with the attenuated
portion of the tail equal to almost half the total
length of the tail, a feature typical of P. platurus.
Boulenger (1885, pp. 49-50) gives a detailed
description of P. platurus but this is certainly based
on several specimens (p- r, Queensland; s, Pt. Curtis
*See footnote on p. 298.
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
295
(Gladstone area, Queensland)) which are not P.
platurus if present distributions have any
significance. The measurements given are of a
specimen much larger than any examined in this
study and the description may be a composite
based on specimens of at least P. platurus , P.
salebrosus and possibly P. cornutus, highlighting
similarities in some of the external morphological
features of these species. Examination of specimens
of P. platurus makes the following elaboration of
Boulenger’s description possible. Features com-
mon to all Phy/lurus (e.g. head large, very depressed,
distinct from neck . . . covered with small granules,
intermixed with conical, spinose tubercles . . .) and
which do not serve to distinguish the species have
been omitted from this description.
Size: P. platurus is a small species, similar in size
to P. caudiannulatus. The largest specimen exam-
ined is R6728. Dimensions of this specimen are
SVL 95-9, HL 29-3, HW 22-3, (tail regenerated).
The largest specimen with an original tail (R8980)
has the following dimensions SVL 85-5, T 69 0, TT
36-5, HL 27*3, HW 22*2.
Flank Tubercles: Invariably minute if present,
and rounded; absent in some specimens (e.g.
R25912, R1550).
Original Tails: Few specimens (§f) have orig-
inal tails. The length of the attenuated tip is close to
half the total length of the tail (0-40 0-56). There
are 6-9 rows of spines across the attenuated tip of
the tail, the posterior half of which is smooth. In
most specimens the whole ‘leaf is dotted with
conical tubercles but in some the central thickened
portion of the tail is smooth. The tubercles at the
edges of the tail are long and slender or short,
almost conical. The anterior portion of the 'leaf
may be strongly heart-shaped or have almost
parallel sides.
Regenerated Tails: Smooth both ventrally and
dorsally; fringe broad or narrow and usually
terminating in a well defined tip which may be
very short or long and tapering.
Rows of Tubercles on Eyelids: Usually one
row, but may be none or two or the tubercles may
be irregularly arranged. All are very small.
Colour: Live specimens of P. platurus have not
been examined for this study. Green (1973, p. 21)
notes that this species is ‘usually light brown or
grey, similar to the sandstone in which it lives’.
Preserved specimens are light brown or grey and
most bear darker brownish flecks or striations all
over the dorsal surface. Ventrally specimens are
pale cream to grey.
Habitat
Green (1973) has discussed the habitat of this
species in detail. P. platurus is almost invariably
confined to sandstone areas.
Distribution
P. platurus occurs only in mid-eastern New
South Wales and is confined to the ‘Sydney-
Hawkesbury Sandstone’ as it is defined by Breed-
en (1972, p. 6). Queensland records of this species
(e.g. Museum of Comparative Zoology, Harvard
specimen 10259, Mt Tamborine, S.E.Q. of Love-
ridge 1935, p. 298 and the small specimen of
Lonnberg and Anderson (1915, p. 3), also from
Tamborine) are almost certainly based on speci-
mens of the southern form of P. cornutus.
Phyllurus cornutus (Ogilby)
(Figs. 1, 3, 4; Plates 36A, 37D, 38D, 39D, 40C)
Gymnodactylus cornutus Ogilby, 1892, p. 8. (Bellenden
Ker Ranges, northeastern Queensland; syntypes AM
R748 50, R752-3, R1094).
Phyllurus lichenosus Gunther, 1897, p. 405, pi. 12.
(Mount Bartle Frere, northeastern Queensland,
holotype presumed lost).
Gymnodactylus sphyrurus Barrett, 1950, p. 31. (non G.
sphyrurus Ogilby).
Material Examined
Syntypes: Bellenden Ker, NE.Q., AM R748-50,
R752-3, R1094.
Other Specimens: QM (NE.Q.): Shipton’s Flat, via
Cooktown, J 17801; 14-4 km SW. Mossman, J7936;
Atherton, J5704; Walsh Camp, via Atherton, J9532;
Millaa Millaa, J5508; Bellenden Ker, J3021; Innisfail,
J5323; Ingham, J3429; AM (NE.Q.): Millaa Millaa
R11375; Bartle Frere, R4769; Evelyn, approx, 1 4*4 km
from Ravenshoe, R26117 23. QM (SE.Q., NE.NSW.):
Tamborine J8183, J 12257; Mt. Tamborine (= Tam-
borine Mt.) J398, J2409, J2933— 4, J3254, J4439, J8359,
J8861, J 10440; Eagle Heights, (?) J4819; Mt. Cluny
J 8074-5, J8099; Binna Burra J8646; Mudgeeraba J4198,
J5690; Canungra J148, J3215; Mt. Ballow J23937;
Beechmont J5649; Tallebudgera J3313, Mt. Lindsey
J 10565; Lamington J5382; Tweed R. J1 143; N.S. W. J730.
AM(N.S.W.) Murwillumbah R2409, A233; Bulga Table-
land R6247, R1 1860; Huonbrook R2315; Nambucca R.
R6792; Nimbin R 1 1 553; Wauchope, R1 1621. Queens-
land. R1 1844. Girraween, near Stanthorpe RG10.
Diagnosis
A large leaf-tailed Phyllurus, distinguished from
P. salebrosus in having calcareous deposits only in
the skin covering the bones of the lower jar (vs
calcareous deposits all over throat) and by length
of attenuated tip of tail (greater than one third total
length of tail vs less than one third total length of
tail). P. cornutus is distinguished from the small
species P. platurus with which it has been confused
296
MEMOIRS OF THE QUEENSLAND MUSEUM
in some areas by tubercles at flank (long hooked
spines surrounded by smaller spines (P. cornutus
NE.Q. specimens) or shorter, unhooked, but
slender spines usually surrounded by smaller spines
(P. cornutus SE.Q.-NE.N.S.W. specimens) vs very
small, rounded tubercles without smaller tubercles
(P. platurus ). Specimens with original tails are
further distinguished by the presence of spines for
the full length of the tail tip (absent towards the tip
in most specimens of P. platurus examined).
Description
Size: P. cornutus is a large species being smaller
than only P. salebrosus. The largest specimen
examined, a northern specimen (J5704), has the
Fig. 1: Phyllurus cornutus. A-B, tail length; B-C, length
of attenuated tip of tail.
following dimensions SVL 140 0, HW 30-9, HL
40-5, tail missing. The dimensions of the largest
specimen with an original tail (R 12935, also a
northern specimen) are SVL 140 0, T 89-4, TT 38-4,
HW 30T, HL 41-6. Specimens from southeastern
Queensland and northeastern N.S.W. are ap-
parently slightly smaller (dimensions of largest
specimen, R11860, SVL 128 0, HW 26 0, HL 35-6,
tail regenerated). Only two specimens of the
Girraween, near Stanthorpe, SE.Q., population
have been examined (RG10; QM live specimen:
SVL 96-9-100-2, T (QM live specimen) 70-0, TT
30-0, HW 22-5-23-4, HL 27-8-28-0).
Flank Tubercles: All northern specimens have
long, slender, hooked, closely adjacent spines
surrounded by smaller hooked spines. In southern
specimens the flank tubercles range from long,
hooked spines surrounded by smaller hooked
spines (J2126) through weakly developed spines
(RG10) to small rounded or flattened, widely
spaced spines which occur in about half the
specimens examined. The absence of long hooked
spines in southern specimens has apparently led to
some of the confusion of P. cornutus with P.
platurus which has weakly developed flank tu-
bercles.
Original Tails: Length of attenuated tip of
tail/total length of tail is shown in Fig. 1 . The length
of the tip of the tail is approximately half its total
length. There are normally 12 (9-14) complete rows
of spines across the tail and the spines occur along
the total length of the tail tip.
Regenerated Tails: Very broad (J3021), or
relatively narrow (J5690, J2409) depending on the
width of the fringe. The tail is smooth, parallel
sided, and usually terminates in a well defined tip.
Rows of Tubercles on Eyelid: usually 2 rows,
rarely 1 or 3.
Colour: Very few live specimens have been
examined but both greenish and greyish specimens
with the typical ‘lichen’ pattern have been seen.
Spirit specimens vary from plain dark brown to
very light grey, and are patterned with fine brown
lines.
Habitat
P. cornutus is an inhabitant of closed forests,
areas formerly covered in closed forest, and
adjoining wet sclerophyll forests. It is believed to be
a tree-dweller as specimens are frequently found
where trees are being felled. P. cornutus is very
common in some areas of northeastern Queensland
(China Camp, approximately 80 km S. of Cook-
town) (J. James, pers. comm.) and southeastern
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
297
Queensland (Lamington National Park), speci-
mens being found sitting on trees at night,
apparently feeding on insects. The Stanthorpe
population is common in granite boulder country,
an area with rather sparse, open vegetation cover.
In the Girraween National Park 30 specimens were
observed in 24 days. The bulk of these were seen
under exfoliations on the boulder sites and the
remainder in deep cracks in the boulders. A single
young specimen was found under an isolated rock
on a large flat boulder. Usually the specimens were
solitary but up to three adults were found in several
‘suitable’ crevices (T. Low, pers. comm.).
Distribution
P. cornutus occurs in northeastern Queensland,
southeastern Queensland, and north- and mid-
eastern New South Wales. No P. cornutus have
been collected or reported between Ingham and
Tamborine although it is likely that the species will
be found here with more intensive collecting,
especially in the closed forests of the ranges of
Mackay, mid-eastern Queensland. With only three
exceptions (Mudgeeraba, SE.Q; Ingham, NE.Q.;
Innisfail, NE.Q.) the localities from which Queen-
sland P. cornutus have been collected are above 300
metres, so it appears that this species has a
preference for elevated, cool areas.
Discussion
Three forms of P. cornutus are recognisable in
the specimens examined — from northeastern
Queensland, southeastern Queensland-north-
eastern New South Wales, and from the Stan-
thorpe area — but, until more specimens are
available especially from mid-eastern Queens-
land, despite differences in size, flank tubercles
and habitat preference, it seems preferable to
regard them as a single, rather variable species
rather than three distinct taxa.
The detailed description of this species by Ogilby
(1892, pp. 8-10) is adequate to enable easy
recognition of northern specimens upon which it
was based but, although he noted this species to be
‘very distinct from G. platurus , its nearest ally’
Ogilby did not elaborate on his reasons for this. He
regarded one unspecified specimen, apparently
collected at the same time and place as the syntypes
of P. cornutus, as being specifically inseparable
from P. platurus. All northern P. cornutus in the
collection of the Australian Museum have all the
features regarded as diagnostic of the species.
In his description of P. lichenosus from Bartle
Frere (the mountain adjoining Bellenden Ker, the
type locality of P. cornutus) Gunther separated P.
lichenosus from P. cornutus because it lacked the
‘strong spinate knob surmounted by a conical
tubercle behind the eye’ and from P. platurus
largely because it had the ‘series of slender tubercles
which flanks the abdomen’ and which are not
present in P. platurus. The conical tubercles on the
knob behind the eye of P. cornutus and the other
species of Phyllurus occur in a variety of sizes and
the size and pattern of tubercles on Phyllurus
specimens vary greatly. Gunther’s illustration
shows a definite cluster of spines, if not a knob,
behind the eye. This feature is not diagnostic of the
species. The original tail of the specimen illustrated
has the attenuated tip greater than l 3 total length of
tail — a feature diagnostic of P. cornutus. No
specimens of P. cornutus from mid-eastern Queens-
land are known and the unnumbered specimen
from Port Curtis, mid-eastern Queensland, re-
ferred to P. cornutus by Gunther is probably P.
salebrosus sp. nov. the only large Phyllurus which
occurs in the area. Garman (1901, p. 2) briefly
described P. cornutus , again from northern mat-
erial, and noted for the first time the difference in
the lateral spines of this species and P. platurus.
Only two other reptiles have a recorded distri-
bution similar to that of P. cornutus. These are
Tropidechis carinatus Krefft 1863 (Trinca, Gray-
don, Covacevich and Limpus, 1971 , pp. 803-4) and
Leiolopisma challenged (Queensland Museum re-
ference collection). Two frogs have similar
distributions — Lechriodus fletcheri (Boulenger)
1 890, Litoria chloris (Boulenger) 1 893 (Queensland
Museum reference collection). Several genera of
reptiles with different northern and southern closed
forest species have been noted — Goniocephalus,
Cacophis (Queensland Museum reference col-
lection) and Mixophyes (Liem and Hosmer, 1973,
p. 455). Specimens of species of these genera from
the intervening closed forests of mid-eastern
Queensland are not known.
Phyllurus caudiannulatus sp. nov.
(Figs. 2, 3, 4; Plates 36B, 37C, 38A, 39A, 40A)
Material Examined
Holotype: Queensland Museum J 15619, J (with
original tail), Bulburin State Forest, via Many Peaks,
ME.Q. Coll. 3rd Sept., 1968, J. Covacevich, T. P. Tebble.
Paratypes: J22286, $ (with original tail) and J22287,
? (with regenerated tail), both from Granite Creek,
Bulburin State Forest, via Many Peaks, ME.Q. Coll.
1st April, 1972, G.B. and S.R. Monteith; J24132, £
(with regenerated tail), Forest Station, 700 m, Bulburin
State Forest, via Many Peaks, ME.Q. Coll. S.R.
Monteith, 29th April, 1974; J25411, <3 (with original
tail), Eungella National Park, via Mackay, ME.Q.
Coll. Feb, 1975, K. McDonald.
298
MEMOIRS OF THE QUEENSLAND MUSEUM
Diagnosis*
A small Phyllurus which differs from other
Phyllurus in having distinct white bands on the
original tail, which may be cylindrical or slightly
leaf-shaped. Regenerated tails are cylindrical (vs
leaf-shape in all other species of Phyllurus). Speci-
mens without tails are very similar in size and shape
to P. platurus. They differ only in being slightly
more spinose and in being restricted to closed forest
of mid-eastern Queensland (vs more open forest in
rocky, especially sandstone areas of mid-eastern
New South Wales).
Description of Holotype
SVL 74-2, TL 1 38-4, T 64-2, HL 22 0, HW 17-5.
Head large, depressed, elongate, triangular, very
distinct from neck; skin of the head more or less
confluent with cranial ossification; ear opening
elliptical, less than half as large as eye. Body
moderate. Limbs long; digits strong; subcylindrical
at base, compressed distally. Head covered in small
granules, intermixed with larger rounded to conical
tubercles; a distinct row of small conical tubercles
across the head midway between ear and eye, one
distinct row of tubercles on eyelid which also bears
another row of smaller, irregularly placed tu-
bercles; labials Body and limbs covered
dorsally and ventrally with small granules; dorsally
these are intermixed with larger conical tubercles
which are largest laterally on the body and quite
small along the dorsal mid-line and on limbs close
to digits. Tail not depressed, slender, tapering,
covered dorsally and ventrally by fine granules
intermixed dorsally with large, conical tubercles;
tubercles in regular rows, smaller towards tip. In
spirit, basically fawn (between Tilleul-Buff and
Pale-Olive Buff of Ridgway, 1912) dorsally and
cream (close to Ivory Yellow of Ridgway) ventrally
darker, brownish (Close Brown of Ridgway)
blotches irregularly over head, back and limbs. Tail
bears six distinct creamish (Cartridge Buff of
Ridgway) bands.
Variation in Paratypes
Variation is very slight. The flank tubercles in the
four specimens of P. caudianulatus are similar.
The first row is very small, rounded at the base and
TABLE 1: Dimensions of Paratypes of P. caudian-
nulatus
Specimen
SVL
TL
T
HL
HW
J22286*
75-8
139-0
53-2
215
17-6
J22287f
76-5
122-6
46-1
21-5
17-1
J241 32 1
8L8
133-8
52-0
23-7
18-9
J25411*
84-8
145-0
60-0
23-4
18-0
*original tail f regenerated tail
Fig. 2: Phyllurus caudiannulatus (J 15619), showing two
rows of tubercles on the eyelid.
*Since this paper went to press a large series of Phyllurus has been collected in the Eungella area by the Australian and
Queensland Museums supported financially by the Australian Biological Resources Survey.
Although these specimens have slightly flared to distinctly leaf-shaped original and regenerated tails unlike P.
caudiannulatus previously described, they resemble P. caudiannulatus in size, body proportions, colour pattern
(including the presence of light bands on the tail), and spination. Like P. caudiannulatus previously described,
specimens from Eungella without tails are difficult to distinguish from P. platurus although light tail bands are not
present in any P. platurus examined and P. caudiannulatus and the Eungella specimens are generally more spinose
than P. platurus.
Snout (mm) Head Width (mm) Attenuated Tip of Tail (mm)
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
299
40 _
30 _
20 _
10 _
□
□
□
□ O
□
□ o □
□ □
□ O
0^0 °
o
o
_l i I I I I I 1 L.
10 20 30 40 50 60 70 80 90
Total Length of Tail (mm)
40
30
20
B
D ’b
s* □
□ □ o
□
• •
* o
• *o
•o
o
20
30 40
Head Length (mm)
50
20 .
Q OSD'
if- - •
10 .
□ n
dP
n n^n
#x D
I I 1
20 30 40
Head Length (mm
Fig. 3: Head and tail measurements in Phyllurus spp.
• P. salebrosus; O P. cornutus; □ P. platurus; X P. caudiannulatus
300
MEMOIRS OF THE QUEENSLAND MUSEUM
sharp. Dorsally they are larger and very spinose.
Tubercles on the eyelids of all specimens are very
small and are irregularly placed. On J22286 the
dark blotches of the dorsum form distinct striations
on the head, giving an impression of a slightly
darker specimen. The tail bands on this specimen
were white in life. The regenerated tails of J22287
and J24 1 32 taper conventionally and are covered in
small granules which form uniformly small tu-
bercles dorsally and which are smooth ventrally.
The regenerated tails lack the distinct light bands of
the original tails of J 156 19 and J22286 but are
marked with fine dark lines forming an irregular
pattern. The original tail of J25411 is slightly
expanded to form a ‘leaf and bears three white
bands and one white blotch instead of the 5-6
distinct light bands of the tails of J 156 19 and
J22286.
Habitat, Distribution
P. caudiannulatus is known only from closed
forest in two localities in mid-eastern
Queensland — Bulburin State Forest, via Many
Peaks and Eungella National Park, via Mackay.
The holotype (J 15619) was collected during the day
under the bark of a lichen-covered dead tree stump.
Two of the paratypes (J22286-7) were collected at
night on a dead tree. The third paratype (J24132)
was found at night on the buttress of a live tree.
J25411 is one of six specimens seen at night by
spotlight approximately 12m above the ground on
trees in the Broken River area of the Eungella
National Park. P. caudiannulatus and P. salebrosus
are sympatric in the Bulburin State Forest but
apparently occupy different niches here. One
specimen of P. salebrosus (J22288) collected at the
same time and in the same place as two paratypes of
P. caudiannulatus (J22286-7) was found on granite
rocks close to the ground.
Phyllurus salebrosus sp. nov.
(Figs. 3, 4; Plates 36C, 37A, 38B, 39B, 40B)
Material Examined
Holotype: Queensland Museum J8 142, S (with orig-
inal tail), Monto. SE.Q.
Paratypes: J2879, $ (with original tail), Roma, SC.Q.:
J4474 (with original tail), Coongoola, SC.Q.; J4897, $
(with original tail) Coomooboolaroo, via Duaringa,
ME.Q.; J9770. c3 (with original tail) Lowmead, ME.Q.;
J5390, S, J6198, $ (with regenerated tails), Goodnight
Scrub, Wallaville, SE.Q.; J8377, $ (with regenerated
tails); Injune, SE.Q.; J22288, ^ (with regenerated tail),
Bulburin State Forest, via Many Peaks, ME.Q.; J6328, $
(tail missing), roof of cave, Cracow Creek, Cracow,
ME.Q.
Other Material: AM Pt Lincoln [?], R5586; Sydney
[?], R300. QM Goodnight Scrub, Wallaville J25360.
Diagnosis
A large (the largest) Phyllurus , P. salebrosus is
distinguished from all other species of Phyllurus by
its extremely rough throat. Tubercles are present
all over the chin, not only on the scales below the
jaw bones as in P. cornutus, the species it resembles
most. The original tail of P. salebrosus further
distinguishes it from P. cornutus (attenuated tip of
tail total length of tail vs <1 total length of
tail).
Description of Holotype
SVL 1391, TL 242-7, T 75-7, TT 20- 1, HL 40-4,
HW 30-6. Head large, depressed, triangular, very
distinct from neck; skin of head almost confluent
with cranium; ear opening elliptical, vertical, less
than half as large as the eye. Body moderate. Limbs
long; digits strong; subcylindrical at base, com-
pressed distally. Head covered with small granules,
intermixed with larger, rounded to conical tu-
bercles; one and a half rows of tubercles on eyelid;
labials body and limbs covered with small
granules, intermixed with conical, almost spinose
larger tubercles which are surrounded by smaller
conical tubercles towards the sides. Chin, throat
and ventral surfaces covered with flat granules
intermixed with larger, but still small tubercles.
Tail depressed, very broad, leaf-like, contracted at
the base, attenuated at the tip; fringed with larger,
hooked tubercles; very large conical tubercles in a
cluster near contracted base of tail, irregularly
towards fringes of leaf, and in eight regular rows
over posterior one third of tail; ventral surface of
tail smooth. In spirit, basically gray (between Pearl
Gray and Light Gull Gray of Ridgway) dorsally
and cream (close to Ivory Yellow of Ridgway)
ventrally. Dorsally there is a series of brown (close
to Burnt Umber of Ridgway) lines forming roughly
shaped hexagons. The ventral surface is flecked
with a lighter brown (close to Road’s Brown of
Ridgway) which forms a series of irregular trans-
verse lines on the chin.
TABLE 2: Dimensions of Paratypes of P. salebrosus
Specimen
SVL
TL
T
TT
HL
HW
J2879*
1190
183-8
64-5
16 6
34-5
27-4
J4474*
120-4
196-7
64-4
15-2
33-8
26-7
J4897*
93-5
167-2
55-3
16-0
29-1
23-4
J9770*
1150
198-4
61-2
17-0
32-8
24-1
J5390 1
133-0
187-2
47-8
—
36-7
29-1
J6 1 98 1
133-2
173-3
39-6
—
38-0
29-2
J8377f
125-4
194-0
47-6
—
35-0
27-4
J22288f
141-0
197-3
53-2
—
38-8
32-0
J6382 t
128-5
—
—
—
39-0
34-5
* original tail fregenerated tail t tail missing
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
301
Fig. 4: Distribution of Phyllurus spp.
302
MEMOIRS OF THE QUEENSLAND MUSEUM
Variation of Paratypes
Variation is slight. Flank tubercles may be quite
long, rounded at the base and surrounded by
smaller tubercles (J4474, J6198) or smaller, narrow,
and without smaller tubercles (J8377). The cal-
careous deposits in the throat may be large and well
developed (J22288) or smaller and less obvious
(J8377). There are two rows of tubercles on the
eyelid in five specimens, one is present in two
specimens (J22288, J9770), and in J4897 and J6198
there are one and a half rows. Rows of spines across
tail in specimens with normal tails vary between 6
(J2879, J4897) and 9 (R5586). Four of the para-
types have regenerated tails, two of which are
illustrated. These differ from the regenerated tails
of P. cornutus in lacking the tip which is usually
present in this species and gives the regenerated tail
a true ‘leaf shape.
Distribution
All P. scilebrosus examined are from mid-eastern
and south-central Queensland, with the exception
of two specimens, R5586 (a completely faded
specimen with original tail from Pt. Lincoln, S.A.)
and R300 (a tailless specimen from Sydney).
Although these could be reliable records of a
species which is apparently uncommon wherever it
occurs, these localities are 1200 and 900 km
respectively from the nearest Queensland locality
in which P. salebrosus is known, and there are no
other records of this species outside Queensland.
Detailed collection data is available for only a
few specimens of P. salebrosus. J6328 was collected
from the roof of a cave adjoining Cracow Creek,
Cracow, mid-eastern Queensland and J22288 was
found at night on granite rocks in closed forest of
the Bulburin State Forest, via Many Peaks, mid-
eastern Queensland. With the exception of the
Bulburin State Forest the localities from which this
species has been collected are generally quite dry
and rocky. Most are sandstone or granite areas.
J22288 was collected at the same time and in the
same place as two of the paratypes of P. caudian-
nulatus in the Bulburin State Forest, a moist area
bounded on either side by drier granite ridges in
which P. salebrosus has been collected (J8142,
Monto and J9770, Lowmead). Three specimens
(J5390, J6198, J25360) have been collected in the
Good Night Scrub, near Wallaville, southeastern
Queensland. The last, one of two specimens seen by
spotlight in similar situations, was found eight
metres above the ground on a Bottle Tree
( Brachychiton rupestre). The Good Night Scrub
where these specimens were found is an Araucarian
Vine Thicket (Webb 1 968, p. 308) on "stony, light to
medium clay with many quartzite fragments’
(Cannon et. al. 1962 p. 121). The only other
specimen collected recently (held live in the col-
lection of Mr T. Low) was found in Robinson
Gorge, via Taroom, southeastern Queensland on
the wall of a small open sandstone cave. The species
is either uncommon or difficult to find in both
areas.
ACKNO WLEDGM ENTS
Mr A. Easton prepared the plates and Miss S.
Hiley prepared the map. Mr T. Low, Mr J. James,
Mr G. Monteith and Mr K. McDonald provided
data on Phyllurus populations. Mr B. Campbell
and Dr H. Cogger have given helpful advice and
criticism throughout this work. Dr H. Cogger and
Mr P. Rawlinson have provided data on type
specimens and supplied obscure references. Mr J.
Tracey and Mr E. Zillman supplied data on the
Good Night Scrub.
LITERATURE CITED
Boulenger, G. A., 1885. "Catalogue of the lizards in the
British Museum (Natural History)’. Vol. 1. (British
Museum: London).
Breeden, S. and Breeden, K., 1972. ‘Australia’s
Southeast. A Natural History of Australia: 2’.
(William Collins Ltd.: Sydney).
Cannon, J. R., Corbett, N. H., Haydock, K. P.,
Tracey, J. G., and Webb, L. J., 1962. An in-
vestigation of the effect of the dehydroangustione
present in the leaf litter of Backhousia angustifolia on
the germination of Araucaria cunninghami — an
experimental approach to a problem in rain forest
ecology. Aust. J. Bot. 10 (2): 1 19-28.
Garman, S., 1901. Some reptiles and batrachians from
Australasia. Bull. Mus. Comp. Zool. Harv. 39: 1-14,
pis. 1-2.
Gray, J. E., 1845. ‘Catalogue of Lizards in the British
Museum (Natural History)’. (British Museum: Lon-
don).
Gray, G. R., 1845. Reptiles. In "The Zoology of the
Voyage of H.M.S. Erebus and Terror’. (Eds. J.
Richardson and J. E. Gray). Vol. 2 (E. W. Janson:
London).
Barrett, C., 1950. 'Reptiles of Australia’. (Cassell:
London).
Green, D., 1973. Observations on the southern Leaf-
tailed Gecko ( Phyllurus platurus). Herpetofauna 6
(2): 21-4.
Gunther, A., 1897. Descriptions of new species of
lizards and of a tree frog from north-eastern
Queensland. Novit. Zool. 4 : 403-6, pis. 11-12.
Kluge, A. G., 1967. Systematics, phylogeny, and
zoogeography of the lizard genus Diplodactylus
Gray (Gekkonidae) Aust. J. Zool. 15 : 1007-1108,
pis. 1-19.
Liem, D. S. and Hosmer, W., 1973. Frogs of the genus
Taudactylus with descriptions of two new species
(Anura: Leptodactylidae). Mem. Qd Mus. 16 (3):
435-57, pi. 29.
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
303
LOnnberg, E. and Anderson, L. G., 1915. Results of Dr
E. Mjoberg’s Swedish Scientific Expeditions to
Australia 1910-13. (part) 7. Reptiles collected in
northern Queensland. Kungl. Svenska Vetensk.
Handlingar 52 (?): 1-9.
Loveridge, A., 1934. Australian reptiles in the Museum
of Comparative Zoology, Cambridge, Massa-
chusetts. Bull. Mus. comp. Zool. Harv. 77 (6):
243-383, pi. 1.
Merrem, B., 1820. ‘Versuch eines Systems der
Amphibien — Tentamen Systematis Amphibiorum’.
( — : Marburg).
Ogilby, J. D., 1892. Descriptions of three new Australian
lizards. Rec. Aust. Mus. 2 : 6-11.
Ridgway, R., 1912. Color Standards and Color No-
menclature. (author: Washington D.C.).
Swainson, W., 1839. The natural history of fishes,
amphibians and reptiles, or monocardian animals. In
‘Cabinet Cyclopaedia’. (Longman, Orme, Brown,
Green and Longmans: London).
Trinca, J. C., Graydon, J. J., Covacevich, J., and
Limpus, C., 1971. The Rough-scaled Snake ( Tropi -
dechis carinatus), a dangerously venomous Aus-
tralian snake. Med. J. Aust. 2: 801-9.
Webb, L. J., 1968. Environmental relationships of the
structural types of Australian rain forest vegetation.
Ecology. 49 (2): 297-311.
Wermuth, H., 1965. Liste der rezenten Amphibien und
Reptilien Gekkonidae, Pygopodidae, Xantusidae.
Das Tierreich 80: 1-246.
White, J., 1790. ‘Journal ofa voyage to New South Wales
with sixty-five plates on non descript animals, birds,
lizards, serpents, environs, cones of trees and other
natural productions’. (J. Debrett: Piccadilly).
Plate 36
Queensland species of Phyllurus.
A. Phyllurus cornutus Girraween (released)
B. P. caudiannulatus Bulburin State Forest (J22286)
C. P. salebrosus Robinson Gorge (released)
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
Plate 36
Plate 37
Variation in flank tubercles of Phyllurus spp.
A. Phyllurus salebrosus J22288, J8377, J2879
B. P. platurus R 10220, R 10387
C. P. caudiannulatus J22287
D. P. cornutus R261 19, R 12935, R2126, R4769, R261 18, J2934,
J5690, A233
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
Plate 37
Plate 38
Variation in original tails of Phyllurus spp.
A. Phyllurus caudiannulatus J22286, J2541 1
B. P. salebrosus J4474, J8142
C. P. platurus R10387, R9274, R27324, R19084, R3793
D. P. cornutus J8075, J8099, R12935, J8359
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
Plate 38
Plate 39
Variation in regenerated tails of Phyllurus spp.
A. Phyllurus caudiannulatus J22287
B. P. salebrosus J22288, J8377
C. P.platurus R4814, R 10220, R28308, R 11621, R6284, R8253
D. P. cornutus J5690, R2409, A233, R4769, R26117
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
Plate 39
Plate 40
Throats of Phyllurus spp.
A. Phyllurus caudiannulatus J 1 5619, J22286
B. P. salebrosus J22288
C. P. cornutus R11375
D. P. platurus R4814, R28308
COVACEVICH: A REVIEW OF THE GENUS PHYLLURUS
Plate 40
Mem. QdMus. 17 ( 2 ): 305-10, pi. 41. [1975]
THE DISTRIBUTION OF THE CANE TOAD, BUFO MARINUS , IN AUSTRALIA
AND ITS EFFECTS ON INDIGENOUS VERTEBRATES
J. COVACEVICH
and
M. Archer
Queensland Museum
ABSTRACT
Bufo marinus was released in cane growing districts of Bundaberg, Mackay, and northeastern
Queensland in 1935 and 1936. At present this species occurs in eastern Queensland and
northeastern N.S.W. between Coen and Ballina, across Cape York Peninsula south of Coen, and
widely in northwestern Queensland. It is highly toxic to several native vertebrates and the results
of mouthing or ingestion of Bufo marinus by thirty-three native species are summarised.
Bufo marinus occurs naturally in North and
South America between southern Texas and cen-
tral Argentina (Mungomery 1936, p. 72). It has
been used as a biological control of insects, snails
and rats (Honegger 1970, p. 453) and has been in-
troduced to southern Florida and Louisiana, most
islands of the Caribbean, Hawaii, and many
western Pacific islands. A consignment of Bufo
marinus from Hawaii was released in sugar cane
growing districts of northeastern Queensland in
June, 1935’ . . . giving definite promise of yielding
some permanent measure of relief in many of the
(cane growing) areas stricken by ‘white grubs’
(larvae of the Grey back Beetle, Dermolepida
albohirtum Waterhouse and the Frenchi Beetle,
Lepidiota frenchi Blackburn) . . .’ (Mungomery
1936, p. 63). In late 1935 the introduction of the
species beyond the Cairns-Tully area, northeastern
Queensland was banned briefly but B. marinus was
soon released in other sugar cane growing districts
between Mossman, northeastern Queensland and
Isis, southeastern Queensland (Mungomery 1937,
p. 12).
Although some local protests were made prior
to the introduction of Bufo marinus into
Australia, it was not until 1938 that doubts about
the wisdom of this introduction were published.
Kinghorn (1938, p. 411) suggested that . . it
would . . . have been wiser to postpone liberation
of the toads until more searching inquiries . . . had
been made.’. Such caution was countered by other
views and it was seriously suggested that B. mari-
nus should be protected to enable it to adapt to its
new environment and control insects, especially
pests to sugar cane, as planned. By 1941 it was
apparent that the role B. marinus was playing in
controlling Greyback and Frenchi Beetles in
Queensland was a minor one. Greyback Beetles
are only rarely in contact with the ground (and
hence with B. marinus ). Contact occurs when a
few adults fall from trees after mating, or when
females move into cane fields to oviposit. B. mari-
nus does not frequent the cane fields at the time
of the Frenchi Beetles flights owing to lack of
cover in the fields.
In early 1974, following repeated press reports of
the spread and potential danger of Bufo marinus ,
we decided to examine the distribution of this
animal and some of its effects on the native fauna.
Our own observations were supplemented with
information gathered from associates, veterin-
arians, and the return of 2,500 circulars sent, with
the cooperation of the Department of Education,
to Primary Schools, Secondary Schools, and Prim-
ary Correspondence Schools. Other circulars were
distributed through the National Parks Branch of
the Forestry Department.
DISTRIBUTION
The present known distribution of Bufo marinus
in Queensland and northeastern New South Wales
is shown in Fig. 1 . Circulars reporting the presence
of B. marinus in particular areas were assessed with
regard to the possibility of identity confusion with
the native Giant Burrowing Frog, Cyclorana aus-
306
MEMOIRS OF THE QUEENSLAND MUSEUM
Fig. 1: Distribution of Bufo marinus in Queensland.
COVACEVICH AND ARCHER: DISTRIBUTION OF THE CANE TOAD
307
trails, which is a large species known as a ‘toad’ in
many western areas of Queensland. This assess-
ment has resulted in the reliability line in Fig. 1.
Without specimens to confirm extreme western
reports of Bufo marinus , we assume these reports
probably refer to Cycloranci australis. Bufo marinus
and Cyclorana australis are compared in Plate 41.
Small numbers of the species have recently been
released accidently (‘Time’, August 5th, 1974) in
the Northern Territory (Darwin) and in Western
Australia (Perth). It is not yet known whether these
unfortunate introductions will result in the estab-
lishment of B. marinus in these areas.
It is probable that the spread of B. marinus as
shown in Fig. 1 was aided by accidental or
deliberate transportation by humans. It was used
by medical practitioners in pregnancy tests, and
was believed to eradicate many species of insects
and snakes. Many universities use these animals in
large numbers, importing them into areas where
they have not yet been established. Private col-
lectors also maintain live B. marinus in many cities.
Reports of deliberate introductions of B. marinus
to Nambour SE. Q., Fraser Island SE. Q., and
Normanton NW.Q. have been received although
most correspondents could provide no information
on how or when the species first appeared in their
areas.
The precise factors controlling the spread of B.
marinus are not understood. There can be no
doubt, however, that temperature, water, food,
and availability of shelter, especially during dry
periods, are controlling factors. Straughan (1966,
p. 230) reports that mating will not occur unless the
water temperature reaches 25-6° C. There is no
information about minimal water temperature
requirements to enable B. marinus tadpoles to
develop to the stage of metamorphosis but this
normally occurs in water 25-6° C and above.
The success of B. marinus in establishing itself in
Australia is attributable to its adaptability, re-
silience, high breeding potential, and relative
freedom from predators. It has been found in every
vegetation zone excluding very high altitude closed
forests and extremely arid districts of the far west.
Specimens have been observed recently in the
following varied habitats — frontal dunes of ocean
beaches (Mon Repos, via Bundaberg, SE.Q.;
Walker Bay, via Cooktown, NE.Q.); the inland
border of coastal mangroves (Amos Bay, via
Cooktown); highly acidic areas of coastal wallum
(Cooloola, SE.Q.); Melaleuca swamps (Cook-
town, NE.Q.); open grassland (Marina Plains,
near Princess Charlotte Bay, NE.Q.); open
sclerophyll forest (Lily Creek, via Cooktown and
Mt. Molloy, NE.Q.); highland closed forest (Mt.
Glorious, SE.Q. and Dawes range, ME.Q.); dry,
sparse open plains adjoining Mitchell and Mor-
eland Rivers, N.Q.; wet or dry creek beds, and
towns and cities between Cooktown, NE.Q. and
Ballina, NE.N.S.W.
Almost any water is used by B. marinus for egg
deposition and fertilization. Queensland Museum
slide MF321 shows a pair in amplexus and a string
of freshly laid eggs in the salt water of Rainbow
Beach, Cooloola, SE.Q. No data is available on the
survival potential of such eggs but eggs, tadpoles,
and newly metamorphosed young have been
observed on a tidal flat at Amos Bay, 30km S. of
Cooktown (J.C. early 1973; January, 1974). Here
water from heavy rain drains across the flat and
accumulates in tyre tracks. The whole flat is
regularly covered in salt water. B. marinus has
apparently used the hot, shallow, slightly brackish
pools to breed successfully in great numbers
although adults have been seen only rarely in the
surrounding area. Schultze-Westrum (1970, p. 37)
has reported the ability of B. marinus in New
Guinea to breed in brackish water and to survive
water temperatures of 40 °C in the larval stage.
Straughan (1966, p. 230) notes that B. marinus will
utilize ‘. . . any waterhole, transient pool, ditch, or
sluggish creek backwater . . irrespective of
whether the water is clear or muddy, whether water
weeds are present or absent, and whether the
substrate is of sand, mud or rock. This remarkable
adaptability as far as breeding sites is concerned
contrasts sharply with the habits of many species of
native frogs which are highly selective in their
choice of breeding sites.
The high breeding rate of B. marinus has also
assisted its spread in Queensland. J.H.B. (1936, p.
25) reported the production of 125,000 eggs by nine
females in one morning and notes that females may
breed more than once in a season. At Barrett’s
Lagoon 8 km west of Cooktown, NE.Q., at the
height of the breeding season in 1970, the shallow
edges of the lagoon appeared to be ‘alive’ with
pairs of B. marinus in amplexus for several
successive nights. Many of the millions of eggs
produced in such permanent water presumably
develop, metamorphose, and grow to adulthood
because they have few predators. In some areas
(e.g. Endeavour R., Cooktown, NE.Q.) teeming
thousands of newly metamorphosed B. marinus
have been observed leaving the water and gradually
moving into the shelter of grass or flood debris on
the banks.
Similarly, its catholic choice of shelter sites
indicates that this requirement for establishment is
only a minor one. We have seen or collected B.
marinus from shallow depressions, in logs, drain
308
MEMOIRS OF THE QUEENSLAND MUSEUM
pipes, debris, hollows at ends of well constructed
burrows (greater than 25cm deep) in soft stony and
clayey soils, under cement slabs, rocks, and sheets
of roofing iron. Presumably by using these shelters
they are able to survive prolonged hot and dry
periods in areas of relatively low rainfall (e.g.
Laura, Charters Towers, and Mt. Garnet). The
remarkable endurance of B. marinus to extremes of
temperature is demonstrated by individuals which
were observed moving slowly away from a
bushfire, having survived massive burning (D.
Crossman, pers. comm.).
B. marinus also exhibits catholic food habits.
Several studies have shown that the major part of
the diet of B. marinus in Queensland is insects,
largely beetles (Mungomery 1936, pp. 63-74;
Straughan 1966, p. 322; van Tets and Vestjens
1973, p. 52) but survey circulars indicate that small
snakes of several species (including Amphiesma
mairii ), lizards, and frogs are also eaten and
recently a small marsupial ( Planigale maculata)
was found in the gut of a toad collected near
Samford, SE.Q. (C. Limpus, pers. comm.). These
tastes, which include its only known common
predator (Amphiesma mairii), ensure a food supply
for B. marinus in any Australian habitat where cold
or extreme aridity do not prevent its survival.
Another adaptation facilitating the spread of B.
marinus is its relative freedom from predation.
With the possible exception of the freshwater
Snake (Amphiesma mairii), which has been re-
ported as eating young B. marinus regularly (Lyon
1973, p. 4) and observed eating tadpoles, B.
marinus has no major predators in Australia. It is a
highly toxic species (Meyer and Linde 1971, p.
522). The bulk of the venom is contained in the
parotid glands and the skin but much of the
remainder of the animal is also apparently toxic to
Australian vertebrates.
EFFECTS ON ENDEMIC VERTEBRATES
The effect B. marinus has had on endemic
vertebrates since its introduction forty years ago
cannot be assessed in detail from the data presented
here and may never be completely known because
little information is available on species pop-
ulations before 1935 and because so many other
possibly detrimental changes have taken place in
the environment since B. marinus was introduced.
Schultze-Westrum (1970, p. 37) in reviewing the
effects of B. marinus on native New Guinean
vertebrates has examined its detrimental effects in
terms of (1) predation, (2) food competition, (3)
competition for resting places.
Many of the reports of predation upon toads are
inconclusive because the subsequent fate of the
predator was not or could not be determined. In the
case noted of the Western Native Cat ( Dasyurus
geoffroii), death was not immediate, and it is
conceivable that such a predator could be seen to
attack a toad, and leave without apparent upset.
There is no substantiated evidence that any ende-
mic mammal can prey successfully upon B.
marinus. We regard B. marinus as a potential
threat to the existence of some terrestrial endemic
predators such as frog-eating snakes and native
cats (Dasyurus spp.) with which it is sympatric. One
of us (M.A.) kept a Western Native Cat (Dasyurus
geoffroii) in captivity for three years. During this
time it was never observed to hesitate in attacking
any small vertebrate. Although it often exhibited
caution in the method of killing these prey, it never
hesitated to bite. It was the result of a single bite (an
accidental encounter) on an individual B. marinus
that caused its death. We have no reason to believe
that other individuals of Dasyurus spp. would
behave any differently if confronted by a toad. It is
of interest to note that the only recent undoubted
reports of the common occurrence of D. viverin-
nus and D. maculatus coming to the attention
of the Queensland Museum are from areas where
toads are known not to occur (e.g. the Lamington
Plateau and the Wyberba district near Stanthorpe).
Predation by toads on some native vertebrates is
noted above. The instance of a whole Planigale
maculata being found in the stomach of a B.
marinus is not a positive indication of predation.
The toad may conceivably have found the animal
dead. However, considering that B. marinus nor-
mally (we have no evidence to the contrary) eats
live prey, this record strongly suggests predation.
Competition involving toads concerns food and
living space. It is now common, when searching for
reptiles and small mammals in some areas of
eastern Queensland, to find only B. marinus,
sometimes in groups, in niches occupied by small
native vertebrates in adjoining toad-free areas. B.
marinus is probably the most common small
vertebrate in eastern Queensland and apparently
survives where native vertebrates cannot. In late
1973 a search for reptiles on a small island in the
partly constructed North Pine Dam near Petrie,
southeastern Queensland, was unsuccessful.
With the exception of a single Litoria fallax there
were no native vertebrates on the island. Every
log and grass-shaded depression housed num-
erous adult and young B. marinus all of which
appeared to be in excellent condition. Com-
petition for and domination of breeding grounds
have probably* been instrumental in reducing
populations of some native frogs along with
COVACEVICH AND ARCHER: DISTRIBUTION OF THE CANE TOAD
309
clearing of habitats. Reports of the disap-
pearance of certain species of frogs ( Limnodyn-
astes peroni) and snakes ( Pseudechis por -
phyriacus, Acanthophis antarcticus, and Pseu-
donaja textilis) following the arrival of B. marinus
in several areas are too numerous to be ignored
although it is impossible to prove B. marinus
responsible.
Effects of Mouthing or Ingesting B. marinus
Kuhlia rupestris (Jungle Perch)
Ingesting; no ill effects. (Remains of B , marinus
found in gut; observed feeding, Daintree R.,
NE.Q., — J. Grimes, pers. comm.)
‘Jew (= Cat) Fish’*
Ingesting eggs, young; no ill effects.
‘Eel’ (probably Anguilla sp.)*
Ingesting eggs, young; no ill effects. (Populations
reported to have been affected adversely initially.)
‘Perch’*
Ingesting eggs, young; no ill effects.
Chelodina sp. (Long-necked Tortoise)
Ingesting; no ill effects. (Specimen observed feeding
on dead B. marinus, Gunn et. al. 1972, p. 1 10.)
Egernia bungana (Land Mullet)
Mouthing; death. (B. marinus entered cage and
was apparently bitten on hind leg- R. Latcham.
pers. comm.)
Varanus spp, (Goannas)
Ingesting whole; death. ( Varanus populations re-
ported repeatedly to have declined in numbers. Captive
specimen died within a few minutes — P. Douglas, pers.
comm.)
Crocodilus porosus (Estuarine Crocodile)
Mouthing whole; no ill effects. (One 2m wild
specimen found near Edward R,, NW.Q. with B.
marinus in mouth; B. marinus released when specimen
captured — J. Bredl, pers. comm.)
Ingesting whole; no ill effects. (Hungry captive
specimens have eaten many B. marinus reluctantly on
several occasions; well-fed specimens refused
them — C. Tanner, pers. comm.)
Carpet Snake' (probably Morelia spilotes variegaia)*
Ingesting whole; effects not reported. (Reported
frequently, but doubtful because M. spilotes variegata
is normally a mammal and bird eater.)
‘Black-headed Python’ (probably Aspidites melano-
cephalus)*
Ingesting whole; effects not reported. (Doubtful,
normally a reptile and mammal eater.)
Stegonotus cucullatus (Slatey Grey Snake)
Mouthing whole; death. (One specimen found dead
with B. marinus only partly ingested, near Cairns,
NE.Q. C. Tanner, pers. comm.)
Amphiesma mairii (Common Keelback)
Ingesting eggs, tadpoles, young; no ill effects. (Lyon
1973, p. 4. A. mairii thrives in captivity on diet of B.
marinus — C. Tanner, pers. comm. Only species
known to utilize B. marinus regularly as food source.)
‘Green Grass Snake’ (probably Green Tree Snake,
Dendrelaphis punctulatus )
Ingesting tadpoles, young; no ill effects. (Captive
specimen — R. Latcham, pers. comm.)
Boiga irregularis (Brown Tree Snake)
Ingesting whole, whole minus skin; death. (Two
adult specimens were force-fed on B. marinus minced
whole and minus skin; both died overnight — C.
Tanner, pers. comm.)
Pseudechis porphyriacus (Red-bellied Black Snake)
Ingesting whole; death. (Populations reported re-
peatedly to have declined in several areas since arrival
of B. marinus ; Covacevich, 1974, p. 23.)
Acanthophis antarcticus (Death Adder)
Mouthing; death.
Ingesting; death. (Two adults found dead with B.
marinus partly ingested, Deighton R., NE.Q. — F.
Woolston, pers. comm. Populations reported to have
declined since arrival of B. marinus ; Shipton’s Flat, via
Cooktown, NE.Q. — J. Roberts, pers. comm.; Mt.
Molloy, NE.Q. — F. Little, pers. comm.)
Pseudonaja textilis (Brown Snake)
Ingesting; death. (Dead adult found in Bauple
State Forest, SE.Q., with B. marinus in gut. — D.
Crossman, pers. comm. Young P. textilis from
Mclvor R., NE.Q. refused young B. marinus in cap-
tivity. Populations reported to have declined in some
areas following arrival of B. marinus, Mclvor R..
NE.Q., — C. Tanner pers. comm.)
Notechis scutatus (Tiger Snake)
Mouthing; death.
Ingesting; death. (J24594 captive juvenile, died
suddenly with B. marinus only partly ingested. Clutch
of approximately 30 captive young were fed young B.
marinus ; all died overnight, some with 2 B. marinus in
gut but most with only 1 partly ingested — C. Tanner,
pers. comm.)
'Ducks’*
Ingesting young; no ill effects. (Domestic ducks not
affected according to survey reports and fowls reported
with no ill effects following ingestion. No indication of
whether ducks are native or domestic.)
‘Ibis’*
Ingesting intestine only; no ill effects.
‘Crane’*
Ingesting; no effects reported.
‘Swamp Hen’*
Ingesting tongue only; no ill-effects.
‘Pheasant’*
Ingesting young; no ill effects.
Corvus sp. (Crow)
Mouthing; death.
Ingesting stomach, tongue, old road killed speci-
mens; no ill effects. Frauca, 1974, p. 1 12-4. (Pet Corvus
sp. died suddenly after pecking B. marinus — R. Lat-
cham, pers. comm. Two circulars report the eating of B.
marinus tongues and stomachs; many report pecking at
old road-kills.)
Dacelo gigas (Kookaburra)
Mouthing; death.
Ingesting freshly killed young adults; no im-
310
MEMOIRS OF THE QUEENSLAND MUSEUM
mediate ill effects. (015549, D. gigas with B. marinus
in mouth found dead with no apparent injuries on
roadside, Woodgate State Forest, SE.Q. Young
freshly killed adult B. marinus consumed readily by
D. gigas at Mt. Nebo, SE.Q. — F. Dale, pers. comm.)
‘Owl’*
Ingesting tongue only; no ill effects.
Podargus strigoides (Tawny Frogmouth)
Ingesting; no ill effects. (Filmer 1974, p. 2).
‘Kite Hawk’*
Ingesting; no ill effects. (Observed following tractor
and eating disturbed B. marinus.)
‘Whistling Kite’*
Ingesting gut only; no ill effects.
Koel
Ingesting gut only; no ill effects. (Cassels 1970, p. 16).
Hydromys chrysogaster (Water Rat)
Ingesting possibly stomachs only; no ill effects.
(Carcasses of B. marinus minus stomachs have been
found in several areas along river banks. H. chrysogas-
ter believed to be predator along Endeavour R.,
NE.Q. — C. Tanner, pers. comm.; many survey re-
ports.)
Dasyurus geoffroii (Western Native Cat)
Mouthing; death. (Captive individual observed to
bite B. marinus and drop it. Panting, rapid pulse,
tetanic contractions, convulsions, and death in 30
minutes.)
Sarcophiius harrisii (Tasmanian Devil)
Mouthing; death. (Captive specimen — P. Doug-
las, pers. comm.)
* Reported only in survey circulars.
DISCUSSION
B. marinus has not eradicated the pests it was
introduced to control; it is still actively spreading in
Australia, and it is not clear what limiting factors
will eventually control this spread. It has a
deleterious effect on some native vertebrate fauna,
which may result in the regional extinction of
native terrestrial predators, although several spec-
ies can apparently ingest B. marinus or its remains
without ill effects.
At present there is no effective biological control
of the species here despite the fact that the snake
Amphiesma mairii regularly consumes B. marinus
larvae and young and is common in many of the
areas supporting B. marinus. Prolonged cold
temperatures and frost and, to a lesser extent,
dryness may restrict its spread much further into
inland Queensland and southern New South
Wales. While B. marinus has thrived in all the areas
to which it has been introduced, there are no
records of successful large scale eradications.
ACKNOWLEDGMENTS
This report has been compiled with the assis-
tance of the Queensland Departments of Educ-
ation and of Forestry, and of L. Bird, C.
Cameron, Dr H. Cogger, D. Crossman, D. Dale,
P. Douglas, C. Farlow, D. Fleay, J. Grimes, R.
Gunson, A. Hall, R. Hobson, J. Hodge, J.
Hutchings, G. Ingram, A. Jeffries, R. Latchan, C.
Limpus, F. Little, B. Lyon, K. McDonald, C.
Morris, E. Reye, J. Roberts, R. Roberts, C.
Tanner, M. Tyler, E. Wagner, J. Winter, F.
Woolston. Bruce Campbell has given helpful
advice on the manuscript. Sue Hiley drew the
distribution map and Allan Easton took the
photographs.
LITERATURE CITED
B , J. H., 1936. How frequently do toads produce
eggs? Cane Gr. quart. Bull., July 1st, p. 25.
Cassels, M., 1970. Another predator on the Cane Toad.
Nth. Qd Nat. 37(151): 6.
Covacevich, J., 1974. An unusual aggregation of snakes
following major flooding in the Ipswich-Brisbane
area, south-eastern Queensland. Herpetofaunal (1):
21-4.
Filmer, I., 1974. Bird Notes. Queensland Ornithological
Newsletter 5 (7): 2.
Frauca, H., 1974. Crows, Channel-bill Cuckoos and
Cane Toads. Aust. Birdlife 1 (6): 1 12-4.
Gunn, R. H„ Galloway, R. W., Walker, J., Nix, H.
A., McAlpine, J, R., and Richardson, D. P., 1972.
‘Shoalwater Bay Area, Queensland’. Technical
Memorandum 72/10 (CSIRO Division of Land
Research: Canberra).
Honegger, R., 1970. Eine Krote erobert die Welt Natur
u. Mus. 100 : 447-53.
Kinghorn, J. R., 1938. The Giant Toad Bufo marinus in
Australia. Aust. Mus. Mag., Dec. 31st, pp. 410-1.
Lyon, B., 1973. Observations on the Common Keelback
Snake, Natrix mairii in Brisbane, south-eastern
Queensland. Herpetofauna 6(1): 2-5.
Meyer, K., and Linde, H., 1971. Collection of toad
venoms and chemistry of the toad venom steroids. In
‘Venomous Animals and their Venoms. W. Bucherl
and E. E. Buckley, (Eds)., Vol. 2, Ch. 40, (Academic
Press: New York).
Mungomery, R. W., 1936. A survey of the feeding habits
of the giant toad, ( Bufo marinus L), and notes on its
progress since its introduction into Queensland.
Proc. Qd Soc. Sug. Cane Tech. 1936 : 63-74.
1937. The present situation regarding the giant Amer-
ican toad in Queensland. Cane Gr. quart. Bull. 5: 12.
Schultze-Westrum, T. G., 1970. ‘Conservation in
Papua and New Guinea.’ Final Report on the 1970
World Wildlife Fund Mission. 46 pp.
Straughan, I. R., 1966. The natural history of the ‘Cane
Toad’ in Queensland. Aust. Nat. Hist. 15 (7): 230-2.
Van Tets, G. F., and Vestjens, W. J. M., 1973. ‘Birds and
their food at Mackay airport, Queensland.’ Div.
Widl. Res. Tech. Mem. No. 8 (CSIRO: Canberra).
Plate 41
Fig. A: Bufo marinus (Brisbane)
Fig. B: Cyclorano australis (Cooktown)
COVACEVICH AND ARCHER: DISTRIBUTION OF THE CANE TOAD
Plate 41
Mem. QdMus. 17 ( 2 ): 313-18. [1975]
THE WOLF SPIDERS OF AUSTRALIA (ARANEAE: LYCOSIDAE): 5.
TWO NEW SPECIES OF THE BICOLOR GROUP
R. J. McKay
Queensland Museum
ABSTRACT
Two new species Lycosa snelli and Lycosa duracki from north Western Australia are added
to the bicolor group of Australian Wolf Spiders.
McKay (1973) defined the ‘bicolor group’ of
lycosid spiders as being similar in coloration, form
of epigynum, and internal genitalia. Two new
species found on open gravel areas in the Pilbara
and Kimberley regions of Western Australia are
described and are placed in the bicolor group of
species. The concept of this species group was
originally morphological in nature, but the species
are also characterised by a number of behavioural
and ecological parameters. An expanded definition
of the bicolor group can be given:
1. The carapace is uniformly coloured and
lacks median or lateral stripes.
2. All species are robust with heavy legs and
well developed eyes.
3. The female epigynum and internal genitalia
are similar in morphology in all species
(McKay 1973, figs. 2, 3).
4. Capture of prey is made at the burrow
entrance and females rarely if ever leave the
immediate vicinity of the burrow; the char-
acteristic positions are: straddling above,
straddling to one side with a leg in contact, or
draping over the mouth of the burrow. All
species rapidly retreat into the burrow when
disturbed, and appear highly sensitive to
ground vibrations or movement.
5. The burrow is usually vertical with an open
entrance flush with the soil surface; more
rarely closed with a pebble, a silk and sand-
grain lid, or a well constructed hinged door.
6. The habitat is always open, largely un-
vegetated areas, with compact soils, usually
in arid or semi-arid regions.
In most details these spiders agree well with the
American species of the genus Geolycosa. Their
behaviour is remarkably similar (see Kaston 1948,
p. 316). All the American species lack dorsal spines
on tibia 4 of females, this is considered by most
authors to be important in the diagnosis of the
genus Geolycosa. 1 + 1 dorsal spines are present on
tibia 4 of females of Australian species as on the
males of Australian and American Geolycosa, so
the generic significance of such spines is open to
doubt. The genus Geolycosa can be defined by
using behavioural data, but not yet in a completely
satisfactory manner by using morphological char-
acters. A redefinition of the genus Geolycosa is
considered to be premature at this stage of our
knowledge of the Australian Lycosinae.
The male palpal organs of Lycosa snelli and
Lycosa duracki have been figured but the structure
of the palpal organs of other members of the
bicolor group is unknown. I have placed these new
species in the older genus Lycosa pending a generic
revision of the Australian lycosid spiders.
Lycosa snelli sp. nov.
(Figs. 1 a-g)
Material Examined
Holotype: Western Australian Museum, WAM
69-797, $ M, C.L. 12 0 mm, Towera Station, north of
Lyndon River, W.A., collected by Mr A. Snell, January,
1952. In spirit.
Paratypes: Western Australia; Barradale 18 km
south, 29.V.1972, RJM, 1 $ P, 1 2 > 4 > 1; inner
and outer palmar tubercles small but distinct.
Large flange along inside edge of second finger.
Hind limbs short; toes cylindrical, with small
tubercles at proximal joints; toes in decreasing
order of length are4>3>5>2> 1; inner
metatarsal tubercle small and at base of first toe; no
outer metatarsal tubercle. Skin smooth.
Dimensions: SV 25 0; TL 110; HW 81; EN 2 0;
IN 2-9; TL/SV - 0-44; HW/SV = 0-32; EN/IN =
0-70.
Colouration in life: On body, dorsal surface
bright purplish-red (Pompeian red of Ridgway
336
MEMOIRS OF THE QUEENSLAND MUSEUM
1912), with two black (Dull violet black of
Ridgway) V-shaped markings on back originating
medially, and extending posteriorly towards in-*
guinal regions; lips edged yellow, with fine black
barring; heavy black band extends from nostrils to
eye, and from eye towards arm; black patch on
lateral surface between arm and leg; ventral
surfaces bright yellow (Strontian yellow of Ridg-
way), with an extensive diffuse red patch on throat;
cloaca, inguinal and axillary regions yellow. On
forelimbs, dorsal surface black with yellow round
base of arm; ventral surface yellow; palm brown
and fingers yellow with black bands on third and
fourth. On hindlimb, dorsal surface black; ventral
surface yellow; posterior surface of tarsus and sole
of foot black. In alcohol all yellow regions have
turned white.
Variation
Female paratypes (AM R38194, NMV D33826,
QM J23945) do not differ much from holotype in
dimensions and proportions: SV 23-8-25-4 (mean
24-5); TL 9-8-10-4 (10 0); HW 8-6-9 2 (8-8); EN
1 -8—1 -9 (1-9); IN 2-8-31 (2-9); TL/SV 0-34-0-43
(0-39); HW/SV 0-36-0-39 (0-38); EN/IN 0-61-0-68
(0-64). Dimensions of male paratypes are similar to
those of females: SV 23-8-29-9 (26 2); TL 9 9-119
(10-8); HW 7 8-9-9 (9 0); EN 1 -6-2-2 (19); IN
2 6-3-4 (3-0); TL/SV 0-38-0-44 (0-41); HW/SV
0-31 0 39 (0-34); EN/IN 0-53-0-74 (0 63).
In most specimens the vomerine series do not
extend to the medial edges of the choanae. Males in
breeding condition show extensive dark brown
nuptial pads on the dorsal surfaces of the first
fingers. (Fig. IB). Breeding females show flanges
on second fingers, as in holotype (Fig. 1 A). Fingers
of males in decreasing order of length are 3 > 2 = 4
> 1. The head stripe is variable in extent and
intensity. The dorsal colouration varies from one
extreme where the entire dorsal surface is bright
red, to the other where the red is replaced by black.
J22680 1 and J23945-6 have turned brown from
preservation.
In most individuals the red colouration on the
throat is confined to a small diffuse patch, but in
some the throat is entirely red and sharply
delimited from the rest of the under-surface.
Larval Morphology
The following description applies to typical
larvae in stages 31 to 33. Eight such larvae had
total lengths between 1 6-6 and 19-0 mm (mean
18-0).
The body is widest across the mid-region of the
abdomen, and at this point it is slightly wider than
deep. The snout is evenly rounded from both dorsal
Fig. 1: A. Ventral view of hand of female in breeding
condition (AM R38194). B. Ventral view of hand of
breeding male. C. Ventral view of foot of female (AM
R38194). (Scale line 5 mm).
INGRAM AND CORBEN: A NEW SPECIES OF KYARRANUS
337
Fig. 2: A, Mouthparts of stage 31 larva. B, Lateral view of stage 31 larva (Scale line 5 mm).
and lateral views. The nares are widely spaced,
being positioned dorso-laterally in line with the
eyes. They open in an antero-lateral direction. The
eyes are dorso-lateral in position and variable (in
diameter) from quite small to fairly large. The
spiracle is sinistral, lateral in position and visible
from above. It opens posteriorly and increases in
diameter from the opening to the origin. The anal
tube is median and quite prominent, opening well
out from the body at the edge of the ventral fin.
The mouth is antero-ventral in position and
opens anteriorly. There is a single row of peripheral
papillae around all but the upper median two-
thirds of the disc where a partially involuted flap of
skin forms a pocket (Fig. 2A). There are no labial
tooth rows. In some specimens, the lower labium
bears broken papillae-like ridges arranged some-
what in rows.
The curved beaks both have fine serrations. The
upper beak is a little less massive than the lower.
In preserved specimens, the dorsal surface is
lightly pigmented, the density increasing over the
brain and spinal cord regions and over parts of the
intestinal mass. In lateral view, the pigment partly
extends over the gill region and the intestinal mass.
The ventral surface is clear. The dorsal surface of
the tail musculature is lightly stippled over most of
its length, the pigment decreasing posteriorly.
Laterally it scatters over the anterior half of the
musculature. The dorsal fin is sparsely stippled
while the ventral fin is clear or with occasional
flecks. The iris appears black.
Field Notes
Kyarranus kundagungan is a ground-dwelling
frog of sub-tropical rainforest in mountainous
areas. It is known only from very damp situations,
particularly saturated leaf-litter and mud in soaks
or small creek beds. In such sites, males call from
water-filled cavities covered with rocks or leaf-
litter. Calling takes place from late August to mid-
February.
Egg masses have been found in late November
and early December. The foam mass resembles that
described by Moore (1961) for K, loveridgei. The
albumen is very wet and sticky with air bubbles
mainly at the top and the large cream-coloured
eggs concentrated at the bottom. The eggs are
about 3T mm in diameter and are contained in
individual jelly capsules of approximately 4 9 mm
diameter.
Tadpoles have been found in typical water-filled
cavities, and groups of newly metamorphosed
larvae in drier hollows. Juvenile frogs have been
located in January and late August, the latter
presumably being progeny from the previous
season. Very young individuals are blackish with
variable amounts of white speckling on the ventral
surfaces.
Vertebrates found synchronosympatric with K
kundagungan were: Lechriodus ftetcheri, Mixophyes
fasciolatus, M. balbus, Litoria pearsoni , Lamp-
ropholis challengeri (Challenger Skink), Tropi-
dechis carinatus (Rough-scaled Snake), and Melo-
mys cervinipes (Mosaic-tailed Rat) at
Cunningham’s Gap; Adelotus brevis , Lechriodus
ftetcheri, Mixophyes fasciolatus, Litoria pearsoni,
L. leseurii, and an undescribed Litoria belonging to
the ewingi group, and Anomalopus truncatus at
Teviot Falls; Adelotus brevis at Mistake Moun-
tains.
Comparison with other Species*
Kyarranus kundagungan is markedly different
from K. sphagnicolus and K. loveridgei in exhibiting
the distinctive red, black and yellow colouration.
Kyarranus sphagnicolus has a more rounded, wider
head and longer tibia (TL/SV 0-45-0-50), the dorsal
ground colouring is grey to reddish brown, the
*Based on examination of 29 specimens of K. loveridgei in
the Queensland Museum collections (from Lamington
Plateau, 35 km S. of Boonah, and Mt. Warning) and 2
specimens of K. sphagnicolus (from Pt. Lookout).
338
MEMOIRS OF THE QUEENSLAND MUSEUM
ventral surface white to brown and the throat
darkly mottled. Breeding females have flanges on
both first and second fingers. The call of K.
sphagnicolus is described by Moore (1961) as a soft
growl 4 gurr-r-r\ quite different from that of K.
kundagungan , which is a deep guttural ‘ork\ It is
difficult to distinguish between the calls of K.
loveridgei and K. kundagungan.
Kyarranus loveridgei and K. kundagungan are
similar in shape, size and habitat. Kyarranus
loveridgei has SV 21-7-30-2 (26-4), TL/SV
0-35-0-46 (0-41), HW/SV 0-31-0-39 (0-35) and EN
/IN 0-50 0-83 (0-63). However, K. kundagungan is
more robust and pear-shaped, and the canthus
rostralis is usually less defined than that of K.
loveridgei. The dorsal colouring of K. loveridgei is
grey to brown, the ventral surface is whitish to grey
with darker speckling, and the throat has brown
mottling, while K. kundagungan has a dorsal
ground colouring of bright red to black, the ventral
surface is immaculate yellow, and the throat has a
diffuse red patch or is completely red. Male K.
loveridgei may be found calling from cavities
anywhere on the forest floor, especially along
creeks, whereas K. kundagungan is confined to wet
patches in creeks and soaks. The cavities of K.
loveridgei are smooth-walled, in moist earth, and
contain no water. The eggs are placed in these, and
the tadpoles when present, are in liquified jelly.
(Moore, 1961, reports similar observations). The
cavities of K. kundagungan are filled with water
which can freely seep in and out, and in which the
eggs are laid and the tadpoles swim during
development.
Moore (1961) suggests that K. loveridgei could
lack nuptial pads, however breeding males of both
it and K. kundagungan possess similar nuptial pads
on the dorsal surfaces of the first fingers.
Etymology
The name kundagungan is derived from the
words ‘kunda’, mountain, and ‘gungan’, frog, from
the dialect of the Kabi tribe, that once lived in
south-east Queensland.
Distribution
Currently known from Mistake Mountains (the
type locality) in the north, south along the Great
Dividing Range to Teviot Falls.
DISCUSSION
Spencer (1901) named Philoria frosti from Mt.
Baw Baw, Victoria. Parker (1940) described Philo-
ria loveridgei from the McPherson Range, SE.
Queensland. Moore (1958) concluded that loverid-
gei was generically distinct from P. frosti and
referred the former and a new species (. sphagnicolus ,
from Pt. Lookout, near Ebor, New South Wales) to
a new genus, Kyarranus.
The status of Kyarranus has been the subject of
controversy. The similarity between Kyarranus and
Philoria in adult and larval morphology, and
ecology has been commented on by several auth-
ors. Brattstrom (1970) indicated his intention to
synonymise these two genera, (a move which
Watson and Martin, 1973, supported by evidence
from life histories). However, Lynch (1971) con-
sidered the two genera to be superficially similar,
reflecting parallel adaptation to a montane en-
vironment, and suggested that they represent the
result of independent divergence from a
Limnodynastes- like ancestor. Tyler (1972) re-
cognised both genera when describing the
superficial mandibular musculature and vocal sacs,
and reviewing the phylogeny of Australo-Papuan
leptodactylids. As currently recognised, Kyarranus
consists of two species groups, one containing K.
sphagnicolus, the other including K. loveridgei and
K. kundagungan. These groups resemble each other
considerably more than either resembles Philoria.
The authors consider that with the data presently
available it is best to recognise Kyarranus as a
distinct genus.
ACKNOWLEDGMENTS
The authors express their sincere gratitude to M.
Anstis of Penshurst, New South Wales, who
contributed the section on larval morphology
including the drawings and to M. J. Tyler (South
Australian Museum) for his helpful suggestions
and assistance in preparation of the manuscript.
They also thank J, Covacevich (Queensland
Museum), Dr D. S. Liem and Dr M. C. Bleakly for
their help. Mr A. Easton of the Queensland
Museum provided the photograph of the paratype.
The Queensland Forestry Department issued the
permit to collect in State Forests. Members of the
‘Wildlife Research Group’ (Queensland) assisted in
the field and in typing the manuscript.
LITERATURE CITED
Brattstrom, B. J., 1970. Thermal acclimation in
Australian amphibians. Comp. Biochem. Physiol. 35:
69-103.
Littlejohn, M. J., 1963. The breeding biology of the baw
baw frog, Philoria frosti Spencer. Proc. Linn. Soc.
N.S. W. 88: 273-6.
Lynch, J. D., 1971. Evolutionary relationships, os-
teology, and zoogeography of leptodactyloid frogs.
Univ. Kans. Mas. Nat. Hist., Publ. Misc. 53: 1-238.
Moore, J. A., 1958. A new genus and species of
INGRAM AND CORBEN: A NEW SPECIES OF KYARRANUS
339
leptodactylid frog from Australia. Amer. Mus. Novit.
1919 : 1-7.
1961. The frogs of eastern New South Wales. Bull.
Amer. Mus. Nat. Hist. 121 : 149-386.
Parker, H. W., 1940. The Australasian frogs of the
family Leptodactylidae. Novit. Zool 42 : 1 106.
Ridgway, R., 1912. ‘Colour Standards and Color
Nomenclature’. (Author: Washington),
Spencer, B., 1901. Two new species of frogs from
Victoria. Proc. R. Soc. Vic. (new ser.) 13 : 175-8.
Tyler, M. J., 1972. Superficial mandibular musculature,
vocal sacs and the phylogeny of Australo-Papuan
leptodactylid frogs. Rec. S. Aust. Mus. 16 : 1-20.
Watson, G. F. and Martin, A. A., 1973. Life history,
larval morphology and relationships of Australian
leptodactylid frogs. Trans. R. Soc. S. Aust. 97: 33-45.
Plate 42
Kyarranus kundagungan, live specimen, Cunningham’s Gap.
INGRAM AND CORBEN: A NEW SPECIES OF KYARRANUS
Plate 42
CONTENTS
Page
Bartholomai, Alan
The genus Macropus Shaw (Marsupialia : Macropodidae) in the Upper
Cainozoic deposits of Queensland .. .. . . . . . ■■ 195
Archer, M.
Ningaui, a new genus of tiny dasvurids (Marsupialia) and two new species.
N. timeaieyi and N. ride}, from arid Western Australia . . . . . . 237
Archlr, M.
Abnormal dental development and its significance in dasyurids and other
marsupials .. .. .. .. . . . • . . •• ..251
Jamieson. B. G. M.
The genus Digaster (Megascolecidae: Oligochaeta) in Queensland .. .. 267
Covacevich, J.
A review of the genus Phyllurus (Lacertilia: Gekkonidae) . . . . . . 293
Covacevich. J. and Archer, M
The distribution of the Cane Toad, Bufo marinus , in Australia and its effects
on indigenous vertebrates . . . . . . . . . . . . . . 305
McKay, R. J.
The wolf spiders of Australia (Araneae: Lycosidae): 5. Two new species of the
bicolor group . . .. .. .. .. ■■ . • •• ..313
McKay, R. J.
The wolf spiders of Australia (Araneae: Lycosidae): 6. The leuckartii group 319
Vernon, D. P. and Martin. J. H. D.
Birds of Moreton Island and adjacent waters .. .. .. .. .. 329
Ingram, G. and Corben, C.
A new species of Kyarranus (Anura: Leptodactylidae) from Queensland,
Australia . . . . • . • • ■ • ■ - ■ ■ • ■ ■ • 335