VOLUME 14

PART

MEMOIRS

OF THE

QUEENSLAND MUSEUM

BRISBANE

VOLUME 14

PART 3

MEMOIRS

OF THE

QUEENSLAND MUSEUM

ISSUED BY THE AUTHORITY OF THE MINISTER FOR EDUCATION,
THE HON, J. C. A. PIZZEY

CONTENTS

☆ ☆

VOLUME 14 PART 3
(Issued 28th August, 1963)

Page

Revision of the Extinct Macropodid Genus Sthenurus Owen in

Queensland . . Alan Bartholomai 51

The Systematic Status of Certain Australian and New Guinean Gekkonid

Lizards . . Arnold G. Kluge 77

New Species of Trinoton Nitzsch (Mallophaga, Insecta) . . Theresa
Clay

87

REVISION OF THE EXTINCT MACROPODID GENUS STHENURUS
OWEN IN QUEENSLAND

Alan Bartholomai
Queensland Museum

While specimens of Sthenurus Owen are not common in Queensland Upper
Cainozoic deposits, the genus is moderately well represented in the collections of the
Queensland Museum. Much of this material formed the basis for the revision of
De Vis (1895), and this has been supplemented by recent collections from the Darling
Downs, south-eastern Queensland, A study of all this material together with the
relevant literature has enabled results of some taxonomic importance to be presented.
Evaluation of the incomplete locality data available supports previous stratigraphic
conclusions by Woods (1960b) regarding the possible existence of at least two different
sequences within the Upper Cainozoic sediments of the Darling Downs.

The genus Sthenurus was proposed by Owen (1873) for Macropus atlas , a
species described originally by Owen (1838) from the Wellington Caves in New South
Wales. S. brehus, described by Owen (1874), is now known to be referable to the
genus Protemnodon Owen. De Vis (1895) recognized the existence of additional species
in collections from the Darling Downs and described S. oreas and S. pales. A further
species, S. occidentalis, was named by Glauert (1910a) from the Mammoth Cave,
Western Australia, while recently Marcus (1962) has described S. andersoni from the
Pleistocene clay deposits near Bingara, north-eastern New South Wales. Most of the
recorded material is from Pleistocene sediments, although one Victorian specimen,
referred by Stirton (1957) to the Sthenurinae has been dated by Gill (1953) as Lower
Pliocene. Another Tertiary occurrence is the lower incisor referred to the Sthenurinae
by Stirton et al. (1961), from the Pliocene Mampuwordu Sands at Lake Palankarinna,
north-eastern South Australia.

De Vis (1895) amalgamated the genus Procoptodon Owen with Sthenurus, a
union which is no longer tenable in the light of the present conception of the genera.
A new subfamily, the Sthenurinae, was proposed by Raven and Gregory (1946) to
include, not only Sthenurus and Procoptodon, but also Palorchestes Owen. Tate (1948)

B

52

MEMOIRS OF THE QUEENSLAND MUSEUM

treated Procoptodon as a subgenus of Sthenurus. Palorchestes was removed from the
Macropodidae by Woods (1958), who showed that its affinities are with the
Diprotodontidae. Supporting Simpson (1930, 1945) and earlier workers, Ride (1959)
assigned Procoptodon to the Macropodinae because of the type of tooth replacement
and the basic pattern of the molars. These macropodine features, as noted by Woods
(1960a), are also present in Sthenurus.

The author wishes to express his appreciation to Mr. J. T. Woods of the
Queensland Museum for his assistance, to Dr. J. W. Evans of the Australian Museum,
Sydney, for the loan of material, and to Mr. W. Dunmall of Dalby, for the donation
of recently collected specimens from Pilton, on the Darling Downs.

Measurements throughout are in millimetres.

Family MACROPODIDAE
Subfamily MACROPODINAE
Genus STHENURUS Owen

Sthenurus Owen, 1873, Proe. Roy. Soc. Lon., 21, p. 128 ; Owen, 1874, Phil. Trans., 164,
pp. 265-274.

Type Species. — Macropus atlas Owen.

Diagnosis. — Mandible with I x spatulate ; P 3 elongate, moderately strongly ridged and
tuberculate, with prominent lingual crest between paraconid and metaconid, and postero-labial
crest associated with protoconid. Molars with broad rectilinear lophids, with oblique ridges
descending anteriorly from cuspids ; links descending lingual to and below levels of protoconid and
hypoconid ; ornamentation never exceptionally coarse, and less so on posterior surfaces of lophids ;
breadth across trigonid basin reduced ; talonid basin centrally elevated ; lateral surfaces of lophids
weakly to moderately convex in anterior view. Maxilla with P 3 elongate, moderately strongly
ornamented. Molars with broad, slightly anteriorly bowed, sharp lophs ; ornamentation never
exceptionally coarse, labially with well developed ridges from paracone and metacone ; median
valley centrally and labially elevated ; metaloph posteriorly with broad fossette.

It has often been noted that Sthenurus and Procoptodon exhibit extreme
shortening, deepening, and increased massiveness of the crania and jaws. However,
as indicated by Marcus (1962), the Pleistocene species of Sthenurus are divisible into
short-jawed forms, somewhat resembling Procoptodon, and relatively long-jawed
forms. This distinction becomes more obscure with specimens from the presumably
older Chinchilla Sand. In the short-jawed members of the genus, the characters
approximating to those of Procoptodon include the increased lateral convexity of the
longitudinal axis of the ramus, the corresponding increase in the thickness of the
ramus, and the firmly ankylosed symphysis.

THE EXTINCT MACROPODID GENUS STHENURUS OWEN

53

Although basic tooth structure is similar in both genera, many points of
difference are readily apparent. In Sthenurus I 4 is relatively larger and much more
spatulate ; P 3 is relatively longer ; the lower molars are comparatively bro&der,
with more rectilinear lophid crests, but with a narrower trigonid basin, and the labial
tooth surfaces are convex in anterior view rather than perpendicular or convergent
towards the crests as in Procoptodon. Ornamentation in Procoptodon is generally
extremely coarse, although fewer ridges may be developed on the protolophid
surfaces. While the posterior surface of the hypolophid in Sthenurus is relatively
smooth, that in Procoptodon is characterized by coarse vertical ridging. In Procoptodon
the central portion of the talonid basin is not elevated.

The upper dentition is similarly distinct in Sthenurus, with P 3 being
comparatively more elongate ; the upper molars are less ornamented, with the lophs
less strongly anteriorly bowed and with somewhat broader, sharper crests. The central
and labial portions of the median valley in Procoptodon are not markedly elevated,
while the metaloph has a relatively narrower posterior fossette.

The suggestion of Tate (1948) that Procoptodon and Sthenurus are related
genera cannot be denied, and further it appears likely that Procoptodon has been
derived from a primitive Sthenurus stock. As Procoptodon has not yet been located
in the Chinchilla Sand or other Pliocene sediments in Australia, it also seems likely
it was a rapidly evolving genus, derived late in the geological history of the group.
Both genera are well represented in the Pleistocene fluviatile deposits of the Darling
Downs.

STHENURUS ANDERSONI Marcus
(Figures 1-3)

Sthenurus areas De Vis ( partim ), 1895, Proc. Linn. Soc. N.S.W., 10 (n.s.), pp. 96-97.

Sthenurus atlas De Vis {partim), 1895, Proc. Linn. Soc. N.S.W., 10 (n.s.), pp. 97-99, pi. 16,
figs. 9-10.

Sthenurus andersoni Marcus, 1962, Rec. Aust. Mus., 15, pp. 209—304, fig. 1.

Material. — Referred specimens in the collections of the Queensland Museum
are as follows : F. 2970, partial left mandibular ramus with P 3 -M 4 , Gowrie, Darling
Downs; F. 2971, partial left mandibular ramus with P 3 -M 4 , Darling Downs (figd.
in part, De Vis, 1895, pi. 16, fig. 10) ; F. 3809, partial left mandibular ramus with
M 4 , Darling Downs ; F. 3810, partial right mandibular ramus with P 3 -M 3 , Darling
Downs ; F. 3811, partial right mandibular ramus with P 3 -M 3 , Darling Downs ;

54

MEMOIRS OF THE QUEENSLAND MUSEUM

F.3812, partial left mandibular ramus with M 2 -M 4 , Darling Downs ; F. 3813, partial
left mandibular ramus with I 2 (incomplete), P 2 “M 3 , juvenile, Kings Creek, Pilton,
Darling Downs ; F. 2978, partial right maxilla with M 2 -M 3 , Darling Downs. (The
original of De Vis (1895, pi. 16, fig. 9) is not extant in the Queensland Museum
collections).

Figure 1 . — Sthenurus andersoni Marcus. Lateral and occlusal views of incomplete mandible ;
F. 2970, natural size.

Diagnosis. — Moderately small, long-jawed ; P 3 comparatively elongate, narrow, with
the median groove at anterior end of postero -labial crest inconspicuous. Lower molars
comparatively small ; with lophids low, weakly ornamented on anterior surfaces ; trigonid basin
very narrow. Upper molars comparatively small, weakly ornamented.

Measurements

THE EXTINCT MACROPODID GENUS STHENURUS OWEN 55

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Maxilla

Specimen

M 1

M*

M*

Univ.

Aust.

of Calif. UC. 60008, (Marcus, 1962)
Mus. F. 49662, (Marcus, 1962)

11-0 x 11*2

13-0 x 12-8

13-6 x 13 0

F. 2978

—

12-9 x —

13-7 x 12-9

Supplementary Description. — P 2 small, subovate in basal outline, anteriorly with
small cuspule at basal margin ; crown slightly constricted mesially ; lingual crest transected
by three sets of vertical ridges ; curved postero-labial crest flanked anteriorly by inconspicuous
groove ; intervening basin moderately deep, ornamented.

Figure 2. — Sthenurus andersoni Marcus. Lateral and occlusal views of juvenile mandible, with
unerupted P 3 exposed by fenestration of the ramus ; F. 3813, natural size.

DP 3 molariform, subrectangular in basal outline, constricted across talonid basin, with
high rectilinear lophids ; hypolophid broader than protolophid. Trigonid basin narrow, its length
approximately three-fourths distance between lophids ; forelink low, descending from near
protoconid anteriorly to near mid-point of anterior margin, turning abruptly lingually as curving

THE EXTINCT MACROPODID GENUS STHENURUS OWEN

57.

transverse ridge above low anterior cingulum ; very weak accessory ridges on anterior surface of
protolophid. Hypolophid with mid-link strong, descending across elevated central portion of
talonid basin from near hypoconid to point labial to mid-point of posterior surface of protolophid ;
accessory ornament of strong oblique ridge from entoconid ; posterior surface very finely grooved.
Posterior cingulum ill-defined.

The present specimens are morphologically in general agreement with the
original material described by Marcus (1962) from the Pleistocene clay deposits,
15 miles east of Bingara, north-eastern New South Wales. However, in the sizes
and proportions of the cheek teeth, the Queensland sample appears to form a more
variable assemblage, with measurements falling, in many examples, towards or even
slightly outside the observed extremes in the original material. Attrition is directly
responsible for many of the apparent discrepancies in lengths. Only in the case of
F. 3813 is the variation possibly significant. However, compared with the Bingara
material, this specimen is identical in all morphological features, except that the
lateral surfaces of the lophids of anterior molars are not so markedly convex in
anterior view. P 2 and DP 3 , retained in F. 3813, appear to be better preserved than
in the Australian Museum specimen, MF. 10, mentioned by Marcus (1962) and enable
the presentation of supplementary descriptive matter relating to the former.

There is also some departure in the form of the premolar. In F. 3811 it is
much narrower above the anterior root and has the lingual crest transected by a
very weakly developed set of vertical labial and lingual ridges near the paraconid,
in addition to the four sets typically present. However, Marcus (1962) notes that
the cuspules along the lingual crest may be subdivided. As in the Bingara material,
P 3 exhibits a basal cingulum-like structure below the paraconid, together with a
variably developed cuspule at its anterior extremity.

The specimen figured as 8. atlas by Owen (1874, pi. 22, figs. 5-8), from the
Pleistocene deposits of Queensland, is most probably referable to 8. andersoni because
of the size of the cheek teeth and the structure of the molars. P 3 appears to be basall} 7
swollen below the paraconid but its true structure is obscured owing to its intermediate
stage of eruption. Another specimen, a partial left maxilla from the Condamine
JRiver, Darling Downs, originally listed by Owen (1845) as Macropus atlas , and later
figured (Owen, 1874, pi. 23, figs. 7-9) as Protemnodon anak, is also apparently
referable to S . andersoni . In the case of the mandibles referred by De Vis (1895) to
S. atlas, Marcus (1962) was correct in his suggestion that the range in morphology
indicates that more than one species of Sthenurus is present and further that some
of the material belongs to S. ajidersoni. De Vis (1895) included the single maxilla
here referred to S. andersoni in S. oreas. Structurally, the teeth are identical with
those described by Marcus (1962) and, as the accompanying table of measurements
shows, the dimensions are likewise comparable.

58

MEMOIRS OF THE QUEENSLAND MUSEUM

Figure 3 . — Sthenurus andersoni Marcus. Lateral and occlusal views of maxillary fragment ;
F. 2978, natural size.

Specimens of S. andersoni from Queensland are from scattered localities in
the Pleistocene fluviatile deposits of the Darling Downs, whereas the material upon
which Marcus (1962) based his description is all from the one locality, Bone Camp
Gully, near Bingara, New South Wales. The slight structural and size variations
presented by the Queensland specimens may be accounted for in terms of geographical
distribution or could be due to slight differences in age between the various localities.

Among the material referred by De Vis (1895) to 8. atlas, there are two
mandibular specimens, F. 813 and F. 814, without locality data, but with preservation
suggesting derivation from the Chinchilla Sand. These show close affinity with
8. andersoni , but are significantly smaller than individuals of this species, and, as well,
exhibit some differences in morphological detail. The ramus is shallower, but remains
approximately the same thickness as in 8. andersoni ; all of the cheek teeth are
significantly smaller ; P 3 is lower crowned ; and the vertical median groove at the
anterior limit of the postero-labial crest is moderately deep and extends towards the
crown base. The lower molars are structurally similar to those in 8. andersoni, but

THE EXTINCT MACROPODID GENUS STHENURUS OWEN

59

the lateral surfaces of the lophids, when viewed anteriorly, are only very slightly
convex. Compared with the partial left mandibular ramus from the Lower Pliocene
of Victoria, referred by Stirton (1957) to the Sthenurinae, the lophids in the present
specimens are approximately the same width, but the teeth are shorter and have
much narrower trigonid basins.

It appears likely, that when better preserved and more accurately localized
material from the Chinchilla Sand becomes available, a taxonomic distinction may
have to be recognized.

STHENURUS OREAS De Vis

(Figures 4-5)

Sthenurus oreas De Vis ( partim ), 1895, Proc. Linn. Soc. N.S.W., 10 (n.s.), pp. 96-97, pi. 16,
figs. 5-6.

Material. — F. 2923, holotype, associated mandibular rami with all cheek
teeth, Darling Downs (figd. in part, De Vis, 1895, pi. 16, figs. 5-6).

F. 3814, partial left maxilla with M^M 3 , Cement Mills, Gore, S.E. Queensland ;
F. 3815, isolated left M 2 , no locality.

Diagnosis. — Short-jawed ; ramus very wide, with longitudinal axis slightly convex
laterally ; masseteric crest high ; masseteric, mandibular, and mental foramina extremely large.
P 3 relatively short, broad, moderately coarsely ornamented. Molars with wide, low, rectilinear
lophids, simply but moderately coarsely ornamented on anterior surfaces ; hypolophid with
accessory ridge close to mid-link generally well developed ; posterior lophid surfaces weakly
ornamented ; trigonid basin moderately wide. Upper molars somewhat coarsely ornamented.

Revised Description. — Mandible strong, thick, comparatively shallow, with

longitudinal axis slightly convex laterally. Symphysis relatively short, ankylosed, at an angle of
approximately 40° to base of mandible ; produced postero-ventrally resulting in decided ventral
extension of basal margin of ramus below P 3 ; geniohyal pit deep, rather high, well forward of
posterior symphysial limit. Diastema short ; ventral margin of ramus broadly rounded between
symphysis and diagastric ridge. Mental foramen extremely large, nearly round, antero- ventral
to P 3 and close to diastemal crest ; accessory foramen well-defined, halfway between posterior root
M 2 and ventral margin of ramus. Ramus with deep labial groove between mental foramen and
point below posterior root of M x . Diagastric process postero- ventral to M 4 , moderately developed,
separated from base of angle by shallow, post-diagastric sulcus, bounded above by shallow
diagastric fossa ; this fossa separated above from shallow depression opening posteriorly into
pterygoid fossa. Post-alveolar shelf short, leading to post-alveolar ridge, ascending gradually
posteriorly to disappear on mesial wall of coronoid process, immediately above extremely large
mandibular foramen. Masseteric crest raised almost to level of occlusion of cheek teeth, with
production of an extremely large masseteric foramen ; masseteric fossa deep. Anterior margin of
coronoid process inclined beyond vertical.

C

60

MEMOIRS OF THE QUEENSLAND MUSEUM

Figure 4 . — Sthenurus orecis De Vis. Lateral and occlusal views of mandible ; F. 2923, holotype,
natural size.