Memoirs

OF THE

Queensland Museum

Brisbane Volume 16

30 June, 1971 Part 1

Volume 16
Part 1

Memoirs

OF THE

Queensland Museum

Published by Order of the Board

Mem . Qcl Mus. (1971) 16(1): 1-18.

MORPHOLOGY AND VARIATION OF THE CHEEK TEETH IN
MACROPUS GIGANTEUS SHAW AND MACROPUS AGILIS (GOULD)

Alan Bartholomai
Queensland Museum

ABSTRACT

The cheek teeth in the Grey Kangaroo, Macropus giganteus Shaw, and the
Sandy Wallaby, M. agilis (Gould) have been described and the variation investi-
gated. The results generally indicate considerably less variation in molar teeth
and in the deciduous, molariform premolar, DPij, than in the remaining premolar
teeth in the series, P' and especially P®. In the cheek teeth, sexual dimorphism in
metrical features is shown to be a factor warranting consideration in macropodid
species.

The present investigation was initiated to provide adequate information on cheek teeth
variation in large, sexed samples of selected, extant macropodids. The necessity for this
type of study has become increasingly apparent in connection with investigations on extinct
macropodid faunas. To facilitate an assessment of possible size and morphological variation
and the determination of species limits associated with this largely Upper Cainozoic taxo-
nomic research, an accurate knowledge of variation present in extant species is of para-
mount importance.

The study has been restricted to cheek teeth because these are relatively abundant and
well preserved elements in the fossil deposits and, for the most part, fossil species are defined
by the morphology of their cheek teeth. These skeletal elements are frequently in sufficiently
large numbers to allow meaningful statistical analyses and comparisons to be undertaken
(Bartholomai, 1967).

The species M. giganteus and M. agilis have been selected for study for a number of
reasons. Most important is the existence of closely related material in Pleistocene deposits,
particularly in the fluviatile deposits of the Darling Downs area, southeastern Queensland,
and the results will be of direct value for comparison in this regard. Large, sexed samples

2

MEMOIRS OF THE QUEENSLAND MUSEUM

of M. giganteus and M. agilis are available, mainly as a result of investigations on living
macropodids currently being undertaken by the Queensland Department of Primary
Industries. Samples considered have been collected from restricted areas, M. giganteus from
the environs of Warwick, southeastern Queensland, and M. agilis from the Townsville area,
mid-eastern Queensland. Use of these has obviated the necessity to consider possible
geographical variation. Results are thus of greater value for use in comparisons with geo-
graphically and stratigraphically restricted fossil samples. Finally, the selection has allowed
an assessment of sexual dimorphism to be undertaken within the ‘kangaroo’ and ‘wallaby’
groups within the Family Macropodidae.

Limited information has previously been available on the subject (Tedford, 1967), but
for the most part the statistical results were based on small, geographically diversified
samples, reducing their value for the purposes intended.

Primary data sheets have been lodged in the Library of the Queensland Museum, and
the specimens from which data were derived are being progressively transferred and
registered into the neontological collections of the Queensland Museum. All measurements
throughout are in millimetres.

The author wishes to express his appreciation to Dr T. H. Kirkpatrick of the Queens-
land Department of Primary Industries, for making available for study the skull remains
here considered.

CHEEK TEETH IN MACROPUS GIGANTEUS SHAW, 1790

P 2 relatively elongate, broader posteriorly than anteriorly, markedly constricted
mesially in occlusal view. Paracone and metacone well developed, high, connected by
secant ridges ascending into mesial constriction, giving longitudinal crest a well defined
bifid appearance in lateral view; slight cuspule frequently present towards limit of anterior
metacone ridge, accompanied by slight, vertical labial and lingual ridges; protocone low,
moderately well defined, with low anterior ridge usually curving labially to unite with
anterior ridge from paracone at anterior limit of crown; posterior ridge from protocone
less well defined, ascending at lingual limit of mesial crown constriction, usually uniting
with low, broad ridge which descends from postero-lingual portion of paracone; protocone
ridge occasionally unites directly at that point; hypocone relatively strongly developed,
well defined, occasionally connected labially by weak ridge to base of metacone; connecting
ridge more frequently ascends from anterior hypocone ridge which curves to postero-lingual
base of paracone; posterior ridge from hypocone curves labially to unite with posterior
ridge from metacone at posterior margin of crown. Lingual basin irregular, divided into
anterior, mesial, and posterior pockets; occasionally additional minor pocket developed

CHEEK TEETH IN MACROPUS

3

mesially. Slight cuspule or ridge sometimes present at antero-lingual base of hypocone.
Small cuspule occasionally present labiad to and below metacone; where absent, cuspule
frequently replaced with slight ridge to metacone.

DP 3 molariform, subrectangular in basal outline, slightly constricted across median
valley; lophs relative high, but metaloph higher than protoloph, bowed anteriorly, with
metaloph broader than protoloph. Anterior cingulum moderately low, usually broad, short,
ascending lingually, with only slight indication of presence of forelink usually seen near
axis of crown. Ridge usually ascends from paracone to unite with labial limit of cingulum;
occasionally anterior paracone ridge descends within labial cingular limit. Midlink strong,
moderately high, curving postero-labially from protocone, uniting with short ridge from
point on metaloph slightly linguad to axis of crown; junction frequently puckered. Median
valley Y-shaped, occasionally more broadly U-shaped in lingual moiety; slight ridges
acsend towards base of valley from paracone and metacone; lingually, valley occasionally
with low, broad ridge transverse to crown axis. Ridge from hypocone strong, ascending
posteriorly to near postero-labial base of crown; posterior ridge from metacone weak.
Slight fossette developed above axis of crown; slight vertical groove often present on
hypocone ridge, close to hypocone.

P 3 comparatively small, subtriangular in basal outline being broader posteriorly than
anteriorly, with slight mesial constriction lingually. Paracone and metacone well developed,
relatively high, connected by secant ridges which normally ascend into mesial cleft, giving
crest a marked bifid appearance in lateral view ; posterior portion frequently better developed
than anterior; subsidiary cuspule occasionally present, associated with a pair of vertical
labial and lingual ridges, towards limit of anterior metacone ridge; this cuspule rarely better
developed ; slight pockets also very rarely developed on both anterior and posterior moieties
of crests. Anterior ridge from paracone ascends towards base of crown, associated with low
ridge around anterior margin where this developed, while posterior ridge from metacone
curves lingually to below posterior base of crown. Hypocone less well defined than other
cusps, usually connected labially to base of metacone by relatively strong ridge; where this
ridge absent, base of metacone connected to anterior ridge from hypocone which ascends
into lingual constriction of crown; posterior ridge from hypocone curves labially to below
posterior margin of crown. Low, variable ridge developed basally, postero-lingual to
paracone. Slight posterior fossette frequently present, but lingual basin largely undeveloped.

M 4 <M 2 <M 3 <M 4 ; molars subrhomboidal in basal outline, slightly constricted across
median valley; lophs relatively high, bowed anteriorly, frequently puckered at unworn
crest ; metaloph broader than protoloph in M 1 and M 2 , often approximately equal in M 3 ,
and narrower in M 4 ; metaloph higher than protoloph. Anterior cingulum relatively low,
moderately broad, short, ascending lingually, with forelink well developed, frequently very
strong, linguad to axis of crown. Slight but variable ridge usually ascends from paracone
to unite with labial limit of cingulum; occasionally ridge may disappear before reaching
base of protoloph. Slight, vertical accessory ridges sometimes present on anterior surface

4

MEMOIRS OF THE QUEENSLAND MUSEUM

of protoloph. Midlink strong, moderately high, curving postero-labially from protocone,
uniting with short ridge from point on metaloph slightly linguad to axis of crown ; junction
frequently puckered. Median valley V-shaped, occasionally more broadly U-shaped in
lingual moiety; ridges ascending into valley from paracone and metacone usually poorly
developed, especially that from metacone; lingual base of valley occasionally with low,
broad ridge transverse to crown axis; lingual margin of median valley rarely with low ridge.
Ridge from hypocone strong, ascending postero-labially to above postero-labial base of
crown; posterior ridge from metacone weak. Slight fossette developed above axis of crown,
while slight vertical groove often present on hypocone ridge, close to hypocone.

P 2 relatively small, subtriangular in basal outline, broader posteriorly than anteriorly,
slightly constricted mesially in occlusal view. Longitudinal crest usually markedly bifid in
lateral view, comprising secant ridges descending posteriorly from anterior cuspid and
slightly curving lingually from postero-labial cuspid ; occasionally additional cuspule present
at anterior limit of posterior moiety of crest, this rarely elevated to produce near trenchent
crest; cuspule associated with variable set of vertical labial and lingual ridges. Anterior
ridge from anterior cuspid descends slightly lingually to variable but sometimes well
elevated cuspule above base of crown; basal swellings associated with cuspule better
developed lingually, with small basal tubercle frequently present postero-lingual to anterior
cuspid. Postero-lingual cuspid well defined, high, with strong ridge descending labially to
posterior cuspid of crest; anterior ridge from postero-lingual cuspid descends mesially to
curve labially and unite with base of crest at position of cleft, defining relatively deep,
posterior basin; slight variable ridges frequently present into basin. Occasionally small
basal tubercle developed antero-lingual to internal cuspid.

DP 3 molariform, subtriangular in basal outline, moderately constricted across talonid
basin, with lophids relatively high, strongly convex posteriorly. Hypolophid much broader
than protolophid. Trigonid basin relatively narrow, its length being less than distance
between lophids, with lingual portion near planar but with basin strongly descending
labially. Forelink moderately strong, relatively low, descending slightly lingually from
protoconid then anteriorly to antero-labial margin of high anterior cingulum. Midlink
high, strong, descending antero-linguaily from hypoconid and curving anteriorly, frequently
abruptly, uniting with moderately well defined ridge from protoconid above talonid basin;
junction often plicate; midlink crosses talonid slightly labiad to axis of crown. Talonid
basin near planar and V-shaped lingually somewhat descending and U-shaped labially;
labial moiety occasionally with slight, broad transverse ridge. Posterior surface of hypo-
lophid generally convex but usually more angular postero-lingually; slight groove descends
postero-lingually from position mesiad to hypocone.

P 3 relatively small, subtriangular to subovate in basal outline, usually broader posterior-
ly and slightly constricted mesially. Anterior margin of crown near vertical. Anterior cuspid
with trenchent ridge usually descending to unite with descending ridge from postero-labial
cuspid slightly posterior to mid-point of crown; crest usually bifid in lateral view, but

Fig. 1 : Some variation in the morphology of deciduous and permanent premolars in the Grey Kangaroo,
Mcicropus giganteus Shaw. P 2 : 1, J20368; 2, J20847; 3, J20901 ; 4, J20983. P 2 : 5, J20847; 6, J 20901 ;
7, J20368. P 3 : 8, J20388; 9, J20390; 10, J20396; 11, J20389; 12, J20397. P 3 : 13, J20397; 14, J20385;
15, J20388; 16, J20398; 17, J20384; 18, J20389; 19, J20390. a = occlusal view; b = lateral view.
All specimens in the Queensland Museum.

6

MEMOIRS OF THE QUEENSLAND MUSEUM

occasional low cuspid developed at base of cleft, associated with set of vertical labial and
lingual ridges; this cuspule rarely better developed to give crest a trifid appearance in lateral
view; posterior ridge from anterior cuspid occasionally with pocket developed along
occlusal surface. Posterior ridge from postero-labial cuspid curves lingually, usually to
unite with high, poorly defined postero-lingual cuspid, but occasionally descending to
postero-lingual base of crown.

M 1 <M 2 <M 3 <M 4 ; molars subrectangular in basal outline, slightly constricted across
talonid basin; lophids relatively high, bowed posteriorly, frequently puckered at unworn
crest; hypolophid broader than protolophid in M 4 and M 2 , often approximately equal in
M 3 and narrower in M 4 ; hypolophid higher than protolophid. Anterior cingulum relatively
high, usually indented anteriorly, moderately broad, its length approximately equalling
distance between lophids; cingulum descends slightly lingually and markedly labially.
Forelink high, strong, descending antero-lingually from protoconid then anteriorly or
slightly labially to point labiad to mid-point of cingulum. Trigonid basin better developed
lingually than labially, with antero-labial fossette normally present in labial moiety.
Anterior and posterior ridges from metaconid poorly developed. Midlink strong, relatively
high, descending antero-lingually from hypoconid then anteriorly above talonid basin to
unite with very short ridge from below or slightly linguad to protoconid; junction often
puckered. Labial moiety of talonid V-shaped, lingual portion U-shaped; occasional, low,
transverse ridge present in base of lingual portion of basin; talonid slopes labially and
lingually from midlink. Anterior ridge from entoconid poorly developed. Posterior surface
of hypolophid with moderately deep, broad diagonal groove from near hypoconid to near
postero-lingual base of crown.

CHEEK TEETH IN MACROPUS AGILIS (GOULD, 1842)

P 2 elongate, slightly broader posteriorly than anteriorly, slightly convex labially and
concave lingually in occlusal view. Paracone and metacone well developed, high, connected
by high secant longitudinal crest; crest straight or slightly concave labially in occlusal view,
transected about one-half way between cusps by well defined set of vertical labial and
lingual ridges, with production of cuspule at crest. Very weak second set of ridges some-
times present close to metacone. Occasionally labial ridge from cuspule bifurcates basally.
Anterior and posterior ridges from paracone and metacone act as continuation of longi-
tudinal crest, ascending and curving lingually towards base of crown. Protocone low,
poorly defined; anterior ridge from protocone curves labially below base of crown as
extension of lingual cingulum ; posterior ridge from protocone forms low lingual cingulum;
cingulum slightly sinuous in lateral view; lingual basin shallow, crossed by low transverse
ridges from cuspules on cingulum to base of longitudinal crest; hypocone better defined
but low, with its anterior ridge contributing to lingual cingulum; posterior hypocone ridge
curves labially below base of crown to unite with posterior metacone ridge; labially, low

CHEEK TEETH IN MACRO PUS

7

ridge connects hypocone to lingual ridge ascending from metacone. Postero-lingual fossette
shallow.

DP 3 molariform, subrectangular to subtriangular in occlusal view, slightly constricted
across median valley; lophs relatively low, but with metaloph higher than protoloph,
bowed anteriorly with metaloph considerably broader than protoloph. Anterior cingulum
relatively low and narrow, short, ascending lingually with no indication of presence of
forelink; ridge ascends from paracone to limit of cingulum forming moderately high secant
area in functional continuity with longitudinal crest of P 2 . Midlink relatively strong, low,
curving postero-labially from protocone to metaloph, near axis of crown; slight contribu-
tion to midlink from metaloph. Median valley V-shaped, occasionally more broadly

p 2

1 — 3 P 2

mm

Fig. 2: Some variation in the morphology of deciduous and permanent premolars in the Sandy Wallaby,
Macropus agilis (Gould). P 2 : 1, J14813; 2, J14606; 3, J14678. P 2 : 4, J14678; 5, J14813; 6, J14632.
P 3 : 7, J14597; 8, J14594; 9, J14579; 10, J14622. P 3 : 1 1, J14594; 12, J14602; 13, J14622. a = occlusal
view; b = lateral view. All specimens in the Queensland Museum.

8

MEMOIRS OF THE QUEENSLAND MUSEUM

U-shaped in lingual moiety. Posterior ridge from paracone and anterior ridge from metacone
curve into valley uniting to form accessory structure across labial moiety of valley. Posterior
ridge from hypocone strong, curving across posterior surface of crown towards postero-
lingual margin; posterior ridge from metacone weak; posterior fossette usually well defined.

P 3 relatively large, subovate to subtriangular in basal outline, being broader posteriorly
than anteriorly; crown slightly convex labially, slightly concave lingually. Paracone and
metacone well developed, high, connected by high secant longitudinal crest which is straight
or slightly concave labially; crest normally transected by two sets of vertical labial and
lingual ridges, with set near mid-point of crest usually best developed. Occasionally, third
set of very slight ridges present near metacone; slight cuspules present along crest, associated
with transecting ridges. Anterior and posterior ridges from paracone and metacone
respectively ascend towards base of crown, with that from metacone curving lingually.
Protocone poorly defined, low, with anterior ridge disappearing below antero-lingual base
of crown. Posterior ridge from protocone forms low lingual cingulum; cingulum normally
slightly sinuous in occlusal and lateral views. Lingual basin shallow, usually slightly broader
posteriorly, crossed occasionally by slight ridges to base of longitudinal crest; occasionally
transverse ridge close to hypocone stronger developed. Hypocone much stronger than
protocone, moderately high, with anterior ridge contributing to lingual cingulum; posterior
ridge curves labially to unite with posterior ridge from metacone ; labial ridge from hypocone
relatively strong, uniting with ridge from metacone at base of longitudinal crest. Postero-
lingual fossette well defined.

M 1 <M 2 <M 3 <M 4 ; molars subrectangular in basal outline, slightly constricted across
median valley; lophs relatively low, bowed anteriorly, frequently puckered at unworn loph
crests; metaloph broader than protoloph in M 1 and M 2 , often approximately equal in M 3 ,
and narrower in M 4 ; metaloph higher than protoloph. Anterior cingulum low, broad, short,
ascending lingually, with forelink normally absent; occasionally low ridge crosses as forelink
from base of protoloph to cingulum approximately below axis of crown, but this never
very strong. Strong ridge ascends from paracone to labial limit of cingulum. Midlink strong,
low, curving postero-labially from protocone, uniting with short ridge from near mid-point
of metaloph. Median valley usually V-shaped; ridges ascending into valley from paracone
and metacone usually weak, but occasionally these unite across labial moiety providing
subsidiary link. Posterior ridge from hypocone strong, ascending labially to point slightly
labiad to axis of crown above base of crown; posterior ridge from metacone somewhat
weaker, curving lingually to unite with hypocone ridge, forming margin to well defined
posterior fossette.

P 2 relatively elongate and comparatively large, subovate to subtriangular in basal
outline, usually slightly broader posteriorly than anteriorly; frequently crown slightly
constricted at posterior one-third, but occasionally labial surface convex and lingual
concave. Longitudinal crest well defined, high, secant, with anterior and posterior cuspids

CHEEK TEETH IN MACROPUS

9

well defined; crest usually transected mesially by a single set of vertical or near vertical
labial and lingual ridges but occasionally a less well developed set may be present close to
posterior cuspid; cuspules developed at crest associated with transecting ridges. Posterior
extension of crest curves lingually before descending to postero-lingual base of crown;
occasionally slight posterior ridge from posterior cuspid of crest also present. Base of crown
rarely tumid, particularly labially.

DP 3 molariform, subtriangular in basal outline, very slightly constricted across talonid
basin; lophids relatively low, bowed posteriorly, with hypolophid broader than protolophid.
Trigonid basin relatively narrow, its length being less than distance between lophids, with
lingual portion near planar but with basin strongly descending labially. Forelink low,
strong, descending labially from protoconid to point on anterior cingulum labiad to axis
of crown; anterior cingulum moderately low. Metaconid with moderately strong ridge
descending anteriorly into trigonid and occasionally to lingual limit of cingulum. Midlink
strong, low, descending anteriorly from hypocone to unite with short ridge from protolophid
somewhat labiad to axis of crown. Talonid basin near planar, V-shaped. Postero-lingual
margin of crown usually somewhat angular.

P 3 relatively large, elongate, subovate to subtriangular in basal outline, usually slightly
broader posteriorly than anteriorly; frequently crown slightly to moderately constricted at
posterior one-third. Longitudinal crest well defined, high, secant, with anterior and posterior
cuspids well defined ; crest normally transected by two sets of vertical or near vertical labial
and lingual ridges of which the anterior set are better developed ; cuspules are present at
crest associated with transecting ridges; rarely crest puckered in unworn teeth. Posterior
extension of crest slightly to strongly developed, either gently curving lingually to descend
to base of crown or verging to abruptly curving lingually before descending. Base of crown
rarely tumid labially.

M 1 <M 2 <M 3 <M 4 ; molars subrectangular in basal outline, very slightly constricted
across talonid basin; lophids relatively low, bowed posteriorly, rarely puckered at unworn
crest; hypolophid broader than protolophid in M 4 and M 2 , often approximately equal in
M 3 , and narrower in M 4 ; hypolophid higher than protolophid. Anterior cingulum mode-
rately low, relatively narrow, its length being slightly less than distance between lophids.
Trigonid basin with lingual portion near planar, but markedly descending in labial portion.
Forelink usually relatively low, strong, curving antero-lingually from protoconid to point on
cingulum slightly labiad to axis of crown; rarely slight suggestion of accessory link present
towards base of protolophid. Anterior ridge from metaconid usually very poorly developed
or absent. Midlink relatively low, strong, descending from hypoconid to unite with slight
ridge from protolophid labiad to axis of crown; junction occasionally incomplete or
puckered. Labial moiety of talonid basin V-shaped but lingual portion frequently more
broadly U-shaped. Rarely low fold present towards base of hypolophid close to midlink.
Posterior surface of hypolophid normally gently convex.

10

MEMOIRS OF THE QUEENSLAND MUSEUM

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TABLE 2

CHEEK TEETH IN MACROPUS

11

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14

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 5

Comparison of Sexed Samples of Macropus giganteus by Student’s Ltest

Maxillae

Mandibles

Character

t

P

Character

t

P

P 2 length . .

P 2 length

3.27

0.001-0.01

maximum breadth

3.96

<0.001

maximum breadth . .

2.63

0.01-0.02

DP 3 length

3.34

<0.001

DP 3 length . .

3.64

<0.001

breadth protoloph . .

5.03

<0.001

breadth protolophid

1.62

0. 1-0.2

P 3 length . .

1.93

0.05-0,1

P 3 length

3.67

<0.001

maximum breadth . .

2.72

0.001 0 01

maximum breadth . .

1.88

0.05-0.1

M 1 length . .

4.77

<0 001

M x length

4.84

<0 001

breadth protoloph . .

6.25

<0 001

breadth protolophid

4.88

<0 001

M 2 length . .

4.45

<0 001

M 2 length . .

4.69

<0 001

breadth protoloph. .

7.15

<0 001

breadth protolophid

7.58

<0 001

M 3 length . .

4.14

<0 001

M 3 length . .

5.51

<0 001

breadth protoloph. .

6.02

<0 001

breadth protolophid

7.68

<0 001

M 4 length

5.33

<0 001

M 4 length . .

7.85

<0 001

breadth protoloph . .

6.52

<0.001

breadth protolophid

7.07

<0.001

TABLE 6

Comparison of Sexed Samples of Macropus agilis by Student’s 1-test

Maxillae

Mandibles

Character

t

P

Character

t

P

P 2 length

2.45

0.02-0.05

P 2 length

0.21

0.8-0. 9

maximum breadth . .

—

—

maximum breadth . .

0.38

0.7-0. 8

DP 3 length

1.58

0. 1-0.2

DP 3 length

0.61

0.5-06

breadth protoloph . .

0.90

0. 3-0.4

breadth protolophid

2.51

0.01-0 02

P 3 length

0 39

0.7-0. 8

P 3 length

0.90

0. 3-0.4

maximum breadth

0.48

0 6-0.7

maximum breadth

1.23

0.2-0. 3

M 1 length

1.35

01-0.2

M 4 length ..

1.07

0.2-0. 3

breadth protoloph. .

0.46

06-0.7

breadth protolophid

1.54

0. 1-0.2

M 2 length

2.14

0.02-0.05

M 2 length

1.20

0.2-0. 3

breadth protoloph . .

1.28

0.2-0. 3

breadth protolophid

1.51

0. 1-0.2

M 3 length

1.27

0 2-0.3

M 3 length

2.13

0.02-0.05

breadth protoloph. .

1.65

0,05-0.1

breadth protolophid

1.76

0.05-0.1

M 1 length

1.29

0.2-0. 3

M 4 length

2.53

0.01-0.02

breadth protoloph. .

1.49

0. 1-0.2

breadth protolophid

1.42

0. 1-0.2

CHEEK TEETH IN MACROPUS

15

DISCUSSION

Early palaeontological investigations, such as those of Owen (1874) or De Vis (1895),
which frequently present the best comparative discussions of the dentition in the subfamily
Macropodinae, have tended to stress the value of the premolars in defining the species
described. More recent investigations involving dentition also tend to emphasize the value
of the morphology and size of the premolars in taxonomic considerations. In this connec-
tion, Tate (1948) has argued largely on the basis of the permanent premolar for the inclusion
of the wallabies within the fossil genus Protemnodon Owen, a conclusion no longer tenable.
In the present study, species are referred to Macropus , following Calaby (1966).

The emphasis on premolars has largely resulted from the greater interspecific variation
usually exhibited by these teeth. The molars are normally adequately described but they are
frequently considered to possess insufficient features showing significant interspecific
differences to justify any great value being placed on them. Where fossil species have been
defined on the morphology of their molar teeth, the importance of the work has often been
reduced by inadequate knowledge of intraspecific variation both in the species being
described or in the living species drawn upon for comparison.

In examining in detail the morphology and size of the cheek teeth in the living Grey
Kangaroo, Macropus giganteus , and the Sandy Wallaby, M. agilis, the most obvious
impression is that intraspecific variation is fairly consistently high in the deciduous and
permanent premolars, P| and to a greater extent in P 3 . Variation is present in the molars
and may infrequently be more extreme in the deciduous molariform premolar and anterior
molar, but in these the basic hypsobrachyodont condition (hypsodont of Bensley, 1903) is
usually only slightly affected by accessory structures, and the differences are frequently
only a matter of degree of development. Notwithstanding these considerations, P| and Pf
do exhibit marked interspecific variation and there appears to be every advantage in
maintaining a measure of dependence upon the taxonomic value of the morphology of
these teeth. It is believed, however, that as many features as possible of the remaining cheek
teeth should also be considered conjointly, together with an assessment of the variation
likely to be encountered in the teeth and other morphological characters in the species
concerned.

Apart from purely morphological variation encountered in the cheek teeth (see figures
1 and 2) in M. giganteus and M. agilis, intraspecific size variation is considerable and this
must be taken into account in any taxonomic assessment. The summaries of measurements
provided for each species includes an evaluation of size variation in each sample as a whole
as well as for individual sexed samples, treated as if each was drawn from a natural popula-
tion. In these, variation as defined by the Coefficient of Variation, V, is calculated to be
from 3 .44-11 .36 in M. giganteus and 2.77-13.21 in M. agilis. By far the bulk of values
fall between 4 . 00 - 6 . 00 in each species.

16

MEMOIRS OF THE QUEENSLAND MUSEUM

Fig. 3:

cpT-'d-r^-o CO to co
cor^S-r^-coobcboo

i i i i i i

Histograms for representative sexed samples of A, Macropus giganteus and B, M. agilis, illustrating
the distribution of some dimensions in the cheek teeth.

CHEEK TEETH IN MACROPUS

17

Bartholomai (1967) has shown that in the fossil species Troposodon minor (Owen),
V mostly falls between values of 4 .00-7.00, but in this case the specimens comprising the
sample were drawn from differing stratigraphic levels within the Upper Cainozoic deposits
of the Darling Downs area and this fact most likely accounts for many of the slightly higher
values than are present in the living macropodines. Studies on other Australian fossil
macropodid samples by Merrilees (1965, 1967), Bartholomai (1970), and, in part, Tedford
(1967) have yielded basically similar results for V to those in T. minor. Tedford’s (1967)
results for V in M. ferragus, however, are generally much higher than have so far been
observed in recent or fossil macropodids suggesting mixing of this sample. Simpson et al.
(1960) state that ‘as a matter of observation, the great majority of them (values for Y in
mammals) lie between 4 and 10, and 5 and 6 are good average values.’ Thus the bulk of
the results for the living species of macropodines here considered conform with values of V
for linear dimensions calculated for other mammal groups, and these are closely approx-
imated by fossil results.

Values for V presented for some molar teeth dimensions in recent M. canguru in
Tedford (1967) are generally higher than those for the present sample of M. giganteus.
Differences apparent most probably relate to the fact that the material considered by
Tedford was drawn from what is now believed to be two species (Kirsch and Poole, 1967),
M. fuliginosus the Kangaroo Island and western mainland species of Grey Kangaroo and
M. giganteus , the eastern Grey Kangaroo. Further, the material considered was collected
from widely separated localities and some geographical variation may have been included.

An interesting situation exists between M. giganteus and M. agilis when the results of
tables 5 and 6 are compared. These represent a comparison of means of the sexed samples
for each species by use of Student’s t test. Results for M. agilis , a member of the ‘wallaby’
group, approach significance at the 5% level in the lengths of P a , M 3 , and M 3 and at the
2% level in the protolophid breadth of DP 3 and the length of M 4 . In M. giganteus ,
representing the ‘kangaroo’ group, the results are generally highly significant at the 0 . 1 %
level and only rarely are these not significant or do they only approach significance. Sexual
dimorphism is therefore generally present in the Grey Kangaroo at least in the characters
considered. This point could have significance where initial sorts of fossil material are being
made utilizing frequency distributions. Bimodality in such distributions is thus likely within
a single species but as can be seen in figure 3 the degree of separation of the modes is not
necessarily excessive.

Kirkpatrick (1965) has shown that it is not uncommon for individuals of M. giganteus
to possess a fifth molar tooth in the cheek teeth series. No evidence of the existence of
was found in the sample used in the present study and results are not believed to have been
complicated by this additional factor. While progression of the tooth row is evident in
M. agilis , the P l are retained until late in life and identification of teeth is uncomplicated.
In M. giganteus , however, there is actual loss of anterior teeth at an early age (Kirkpatrick,
1964), necessitating greater care in determination of molars after the permanent premolars
have been ejected.

18

MEMOIRS OF THE QUEENSLAND MUSEUM

LITERATURE CITED

Bartholomai, A., 1967. Troposodon, a new genus of fossil Macropodinae (Marsupialia). Mem. Qd Mus.
15: 21-33.

1970. The extinct genus Procoptodon Owen (Marsupialia: Macropodidae) in Queensland. Mem. Qd
Mus. 15: 213-33.

Bensley, B. A., 1903. On the evolution of the Australian Marsupialia; with remarks on the relationships
of the marsupials in general. Trans. Linn. Soc. Land. ( Zool .) (2) 9: 83-217.

Calaby, J. H., 1966. Mammals of the upper Richmond and Clarence Rivers, New South Wales. C.S.I.R.O.
Div. Wildl. Res. Pap. 10: 3-55.

De Vis, C. W., 1895. A review of the fossil jaws of the Macropodidae in the Queensland Museum. Proc.
Linn. Soc. N.S.W. (2) 10: 75-133.

Gould, J., 1842. Notes. Proc. Zool. Soc. Lond. 1841: 80-3.

Kirkpatrick, T. H., 1964. Molar progression and macropod age. QdJ. Agric. Sci. 21 (1): 163-5.

1 965. Studies of Macropodidae in Queensland. II . Age estimation in the Grey Kangaroo, Red Kangaroo ,
Eastern Wallaroo and Red Necked Wallaby, with notes on dental abnormalities. Qd J. Agric.
Anim. Sci. 22: 301-7.

Kirsch, J. A. W., and Poole, W. E., 1967. Serological evidence for speciation in the Grey Kangaroo,
Macropus giganteus Shaw, 1790 (Marsupialia: Macropodidae). Nature 215: 1097-8.

Merrilees, D., 1965. Two species of the extinct genus Sthenurus Owen (Marsupialia, Macropodidae) from
south-eastern Australia, including Sthenurus gilli sp. nov. J. roy. Soc. W. Aust. 48: 22-32.

1967. South-western Australian occurrences of Sthenurus (Marsupialia, Macropodidae), including
Sthenurus brownei sp. nov. J. roy. Soc. W. Aust. 50: 65-79.

Owen, R., 1874. On the fossil mammals of Australia. Part VIII. Family Macropodidae: Genera: Macropus
Osphranter, Phascolagus , Sthenurus and Protemnodon. Phil. Trans. 164: 245-87.

Shaw, G., 1790. ‘Nat. Miscell.’ pi. 33 and text.

Simpson, G. G., Roe, A., and Lewontin, R. C., 1960. ‘Quantitative Zoology’. Rev. Ed. pp. viii -f 440
(Harcourt Brace and Co.: New York).

Tate, G. H. H., 1948. Results of the Archbold expeditions, No. 59: Studies on the anatomy and phylogeny
of the Macropodidae (Marsupialia). Bull. Amer. Mus. Nat. Hist. 91: 237-351.

Tedford, R. H., 1967. The fossil Macropodidae from Lake Minindee, New South Wales. Bull. Dept. Geol.
Uni. Calif. 64: 156.

Mem. Qd Mus. (1971) 16 ( 1 ): 19-26, pl.l.

DASYURUS DUNMALLI, A NEW SPECIES OF FOSSIL MARSUPIAL
(DASYURIDAE) IN THE UPPER CAINOZOIC DEPOSITS OF QUEENSLAND

Alan Bartholomai
Queensland Museum

ABSTRACT

A new species of fossil marsupial native cat, Dasyurus dunmalli, is described
from the Chinchilla Sand of possible Pliocene age, while further material derived
from the Pleistocene fluviatile deposits of the eastern Darling Downs is referred
to the extant D. viverrinus Shaw, 1800. The presence of Dasyurus is noted in the
cave and fissure-fill deposits at Cement Mills, Gore.

Small dasyurids are poorly represented in collections of fossil marsupials from the
Upper Cainozoic deposits of Queensland. A number are present however, which are refer-
able to the genus Dasyurus Geoffroy. Lydekker (1887) recorded the presence of D. viverrinus
Shaw from the Pleistocene fluviatile deposits of the Darling Downs area, southeastern
Queensland, but no definitive study has been undertaken on the limited sample available.
The present study was prompted by the recent discovery of additional material and the
existence of previously unreported specimens in the collections of the Queensland Museum
and is in keeping with a general study of the Upper Cainozoic faunas of Queensland.
Although part of the sample is poorly localized, it is evident that the specimens have been
derived both from the Chinchilla Sand of possible Pliocene age and from the Pleistocene
fluviatile deposits. A single specimen has also been recovered from the Pleistocene cave and
fissure-fill deposits at Cement Mills, Gore, southeastern Queensland.

Studies in recent years in the Diprotodontidae, Macropodidae and Phalangeridae have
indicated that there are frequently, but not invariably (Bartholomai, 1967), taxonomic
differences at the specific level between samples from the possibly Pliocene and Pleistocene
deposits in the Darling Downs area and it was considered valuable to ascertain if any
distinction exists within the genus Dasyurus of the Dasyuridae.

The author wishes to express his appreciation to Mr W. Dunmall of Dalby, southeastern
Queensland, for the donation of recently collected specimens from Chinchilla to the
Queensland Museum.

20

MEMOIRS OF THE QUEENSLAND MUSEUM

Family DASYURIDAE
Subfamily DASYURINAE
Genus Dasyurus Geoffroy
Dasyurus dunmalli sp. nov.
(PI. 1 , figs. 1-4)

Material: F6579, holotype, partial left mandibular ramus with base of Ci, Pi-P 3 , M r M 3 with
protoconid shattered in M 2 and M 3 , adult, Chinchilla Sand at Chinchilla Rifle Range (Rifle Range number
78, Par. of Chinchilla), from side gully leading into middle gully system, Darling Downs, SE.Q.

F742, partial right mandibular ramus with C L , alveoli for Pj-Pjj, M r M 3) M 4 broken, adult. Darling
Downs. F6580, partial left mandibular ramus with alveoli for P 2 -P 3 , M 1( Chinchilla Sand at Chinchilla
Rifle Range, from side gully leading into middle gully system, Darling Downs.

Diagnosis: This species is morphologically very similar to the living Dasyurus viverri-
nus Shaw but is distinguished by the presence of an additional, very small premolar imme-
diately anterior to Mj. Although this tooth is readily lost, an alveolus is consistently present
in specimens in which that portion of the ramus is preserved.

Description: Ramus shallow anteriorly, deepest below posterior molars. Symphysis
elongate, relatively deep, ovate, extending posteriorly to below anterior root M 2 ; geniohyal
pit shallow. Mental foramina usually positioned below P 2 and M t closer to ventral margin
of ramus than dorsal; occasionally third foramen present below M r M 2 .

Canine large with strongly upcurved root and recurved, piercing crown. Laterally,
crown rounded, but lingual surface markedly angular anteriorly and posteriorly, with
antero-lingual portion more strongly angular; mesially, lingual surface convex. Enamel
margin raised higher antero-lingually than elsewhere around tooth, with slight basal
cingulum present paralleling enamel margin.

Pi<P 2 >P 3 ; all premolars basically similar morphologically, the major differences
being in size and in the presence of only a single root in P 3 compared with a divided root
in P! and P 2 . First premolar relatively small, in close juxtaposition with postero-labial
surface of C 2 and positioned with its anterior margin more anterior than posterior surface
of the canine. Single high cuspid present, positioned above anterior moiety of crown;
anterior ridge from cuspid relatively steep, but posterior ridge descends more gently to
near horizontal posterior occlusal surface of tooth; slight basal cingulum present labially
and lingually, strongest postero-labial and postero-lingual to cuspid. In occlusal view,
crown moderately convex labially but only slightly convex or flattened lingually. P 2 morpho-
logically similar to P t but cuspid positioned slightly more anteriorly. P 3 very small, more
ovate in occlusal view, with basal cingulum well defined.

NEW SPECIES OF FOSSIL MARSUPIAL

21

M 1 <M 2 <M 3 >M 4 ; molars high crowned with sharp, generally distinct cuspids and
well defined basins. Protoconid best developed, strong, generally widely separated from
paraconid and united with this cuspid by strong, secant ridge; Mj with paraconid reduced
to slight cuspule towards base of anterior ridge from protoconid. Metaconid generally well
defined, lower than protoconid, usually well separated from that cuspid ; metaconid weak
in M 1; positioned close to protoconid. Trigonid basin almost non-existent in M t , but well
developed in M 2 -M 4 . Hypoconid somewhat stronger than entoconid, but both cuspids low
compared with protoconid ; hypoconid connected to base of protoconid by relatively strong,
secant ridge; similar ridge curves postero-lingually to unite usually with hypoconulid then
to entoconid; hypoconulid very weak in M x , becoming stronger posteriorly, but apparently
absent or very weak in M 4 ; entoconid less distinctly united to base of metaconid. Talonid
basin low, well defined. Slight antero-labial cingulum ascends from below protoconid to
below paraconid; usually less well defined antero-lingual cingulum present. Posterior
cingulum slight.

TABLE 1

Measurements (mm) for Dasyurus dunmalli sp. nov.

Specimen

Pi

P 2

P 3

M,

m 2

m 3

m 4

F6579*

F740

F741t

3. 4x1. 9
3.5x 1.8

3. 8x2.1
4.1 x2.0

1.8x1. 5

4. 8x2. 6
4. 4x2. 3

5. 7x3.0
! 5. 1x2. 8

6.1 x3. 6
5. 5x3. 2

5 .2 x —

4. 9x2. 6

* holotype f Doubtfully referred to D. dunmalli.

Discussion: Dasyurines are poorly represented as fossils in collections from the Upper
Cainozoic deposits of Queensland. Smaller forms are particularly poorly known, but larger
forms are comparatively better represented. Two species of Sarcophilus Geoffroy and Cuvier
have been recorded from Queensland, S. laniarius Owen, originally described from the
Wellington Caves, New South Wales (Owen, 1838) and locally present in Pleistocene
fluviatile and cave deposits and S. prior de Vis, 1884, based on the proximal articular surface
of a right tibia with portion of the shaft, from the Chinchilla Sand at Chinchilla. The
smaller dasyurine, Dasyurus viverrinus Shaw has been recorded from Gowrie, Darling
Downs by Lydekker (1887), but at that time was known only from a single mandibular
fragment, British Museum (Natural History) specimen number Ml 906. Longman (1925)
indicated the presence of Antechinus flavipes Thomas in the Pleistocene cave earth deposits
at Marmor, mid eastern Queensland.

22

MEMOIRS OF THE QUEENSLAND MUSEUM

TABLE 2

Measurements (mm) for Dasyurus viverrinus Shaw (Fossil Sample)

Specimen

Pi

P 2

M*

m 2

m 3

m 4

F737

4. 8x2. 6

5. 4x2. 9

5. 5x2. 9

F738

—

—

—

—

5. 8x3.0

5. 6x2. 8

F739

3.8 ;< 1.4

3. 9x1. 7

4. 9x2. 4

5. 6x2. 8

5. 8x3.0

5. 4x3.0

D. dunmalli sp. nov. is currently known only from fragmentary mandibular rami and
as yet, no maxillary specimens referable to the genus Dasyurus Geoffroy have been recovered
from the Upper Cainozoic deposits of Queensland. Although only portion of the sample
referred to D. dunmalli is accompanied by locality information more specific than ‘Darling
Downs’ preservation of the remaining specimens is typical of derivation from the Chinchilla
Sand of possible Pliocene age (Woods, 1956). It is thus highly likely that the species is
restricted to that Formation.

One aged specimen, F741, a partial left mandibular ramus with M 2 -M 4 , from the
Chinchilla Sand at Chinchilla has been doubtfully referred to D. dunmalli. This specimen
does not present that portion of the ramus containing the diagnostic feature of the species.
Measurements for this ramus are included in table 1.

In addition to the material described as D. dunmalli, there exists in the collections of
the Queensland Museum, a series of mandibular fragments numbered F736-9 inclusive
which are morphologically identical with the living D. viverrinus. Again, the specimens
generally possess limited locality information with the exception of F736, from Clifton,
Darling Downs. The remainder all have preservation consistent with their derivation from
the Pleistocene fluviatile deposits of the eastern Darling Downs. A single mandibular
fragment, F3703, referable to the genus Dasyurus has been recovered from the cave and
fissure-fill deposits at Cement Mills, Gore, southeastern Queensland. Although this specimen
lacks teeth, the proportions of the ramus and its morphology suggest that it also is most
likely of D. viverrinus. Measurements for the series of specimens from the Pleistocene
fluviatile deposits are presented in table 2, while table 3 presents for comparison, mandibular
measurements from a recent sample of D. viverrinus.

Apart from the presence of the minute additional third premolar, or its alveolus, in
all specimens of D. dunmalli in which that portion of the ramus has been preserved, no other
consistent morphological differences are evident between this species and the fossil and
recent samples of D. viverrinus. The proportions of the first and second premolars are

NEW SPECIES OF FOSSIL MARSUPIAL

23

TABLE 3

Measurements (mm) for Dasyurus viverrinus Shaw (Recent Sample)

Specimen

Sex

Pi

P 2

Mi

M a

m 3

m 4

J17769

<3

4. lxl.5

4. 6x1. 9

5. 4x2. 4

5. 8x3.0

5.7x3. 1

6. 0x3.0

J7996

3. 9x1. 4

4. 3x1. 7

5. 0x2. 3

5. 5x2. 8

5. 7x3. 2

5. 7x3.1

J8787

<?

4. Oxl. 5

4. 3x1. 5

4. 6x2. 3

5. 5x2. 8

5. 7x3.0

5. 6x3.0

J20313

<3

4. 0x1. 7

4. 5x1. 9

4. 9x2. 4

5. 3x3.1

5. 7x3.0

5. 7x3.1

J20414

—

3. 7x1. 7

4. 2x1. 8

4. 7x2. 4

5. 6x2. 8

5. 8x3.1

5. 4x2. 9

J20413

—

4. lxl. 7

4. 2x1. 8

4. 8x2. 3

5. 8x2. 9

5. 8x3. 3

—

J20379

<y

3. 6x1. 4

3. 9x1. 5

4. 4x2. 2

5. 3x2. 5

5. 6x2.7

5. 2x2. 4

J 203 80

?

3. 7x1. 3

4. 0x1. 6

4. 6x2. 2

5. 0x2. 6

5. 2x2. 8

5. 0x2. 6

J20381

—

4. 2x1. 5

4. 3x1. 6

5. 0x2. 2

5. 7x2. 8

6. 0x3.0

5. 6x2. 6

J20382

$

4. 0x1. 5

4. 2x1. 6

4. 7x2. 4

5. 1x2. 9

5.6x3. 1

5. 5x2. 8

J20383

—

3.6x1 .4

4. 0x1. 5

4. 2x2. 2

4. 8x2. 4

5. 3x2. 7

5. 0x2. 5

somewhat different but the sample of D. dunmalU is too small to ascertain whether this
difference can be accommodated by normal variation within the species. Comparison of
the tables of measurements indicates that in other metrical features, differences are
negligable. Unlike many of the Pleistocene forms, gigantism was not a feature of Dasyurus.

Two Quaternary fossil species of Dayurus are known from elsewhere in Australia;
D. affinis McCoy, defined by McCoy (?1862) from Victoria, and D. bowlingi Spencer and
Kershaw from King and Deal Islands, Bass Strait (Spencer and Kershaw, 1910). The
syntypes of D. affinis have been figured by Gill (1953) and lack any trace of the presence
of P 3 . The species appears to be morphologically close to D. maculatus, but Ride (1964)
suggests that while it is nearly as large as the living species, it differs in its proportions.
Spencer and Kershaw (1910) provide adequate information to show that D. bowlingi is
larger than D. maculatus and morphologically distinct. Size considerations alone serve to
distinguish both southern fossil species from D. dunmalU and the fossil specimens of
D. viverrinus.

That fossil representatives of the genus Dasyurus should be found containing reduced
third premolars is not surprising when the dental characteristics of the family as a whole are
considered. In fact, complete loss of P 3 is relatively rare in living members of the family,
as shown by Tate (1947). He indicates that this condition is found only within Dasyuroides
Spencer, Dasycercus Peters, Myoictis melas (Muller and Schlegel), Pseudantechinus Tate,
Dasyurus (including Dasyurops Matschie, Dasyurinus Matschie and Satanellus Pocock;
see Ride, 1964, and Simpson, 1945) and Sarcophilus. The majority of living dasyurids trend
towards reduction of P 3 , with strongest development in P 2 . Relatively few forms retain P 3
in its normal, ‘primitive’ condition, larger than P 2 .

24

MEMOIRS OF THE QUEENSLAND MUSEUM

Tate (1947) suggests that reduction and loss of P 3 (P 4 in Tate) is evidence of specialisa-
tion. This suggestion follows that of Thomas (1887) and Bensley (1903). On the basis of
comparison with other dasyurids in which P 3 is vestigial, it appears likely that when maxillary
remains of D. dunmalli are located the third upper premolar may be more strongly
developed.

The species Glancodon ballaratensis Stirton, 1957, from the Pliocene of Victoria, has
been suggested by Ride (1964) to be structurally ancestral to Sarcophilus. It nevertheless
possesses some characters which Ride interprets as suggesting a more remote relationship
with Dasyurus. The presence of D. dunmalli in possibly Pliocene sediments, a form which
is only slightly more ‘primitive’ than living Dasyurus supports Ride’s conclusions on the
relationships within the group. Although no definite information is available on earlier
representatives of the Dasyurus line, it appears likely that their ancestry must have been at
least as early as the Middle Tertiary on the basis of the apparent stage of evolution of
D. dunmalli. An early dasyuroid fossil has been recorded by Stirton at al. (1961) from the
possibly Oligocene Etadunna Formation of the Lake Eyre Basin. However, in the absence
of additional information on its morphology, no purpose can be served by further specula-
tion on the relationships of this form to support or reject Ride’s (1964) treatment of it
within the Thylacinidae.

LITERATURE CITED

Bartholomai, A., 1967. Troposodon, a new genus of fossil Macropodinae (Marsupialia). Mem. Qd Mus.
15 : 21-33.

Bensley, B. A., 1903. On the evolution of the Australian Marsupialia; remarks on the relationships of the
marsupials in general. Trans. Linn. Soc. Loud. ( Zool .) (2) 9: 83-217.

De Vis, C. W., 1884. On tooth-marked bones of extinct marsupials. Proc. Linn. Soc. N.S.W. 8: 187-90.

Gill, E. D., 1953. Catalogue of Quaternary types and figured specimens in the National Museum of Victoria.
Mem. Nat. Mus. Vic. 18 : 168.

Longman, H. A., 1925. Fossil marsupials from Marmor. Mem. Qd Mus. 8: 109-10.

Lydekker, R., 1887. ‘Catalogue of the fossil Mammalia in the British Museum (Natural History).’ Part V
XVI + 345 pp. (Taylor and Francis: London).

McCoy, F., 71862. Note. Quarter Sheet N.W., Geological Survey of Victoria.

Owen, R., 1838. Fossil Marsupialia from the caves of Wellington Valley. In Mitchell, ‘Three Expeditions
to the Interior of Eastern Australia.’ Vol. 2, pp. 359-63 (T. and W. Boone: London).

Ride, W. D. L., 1964. A review of Australian fossil marsupials. J. roy. Soc. W.A. 47: 97-131.

Shaw, G., 1800. ‘General Zoology’, I, 513 pp. (London).

Simpson, G. G., 1945. The principles of classification and a classification of mammals. Bull. Amer. Mus.
Nat. Hist. 85: I-XVI, 1 350.

NEW SPECIES OF FOSSIL MARSUPIAL

25

Spencer, B., and Kershaw, J. A., 1910. A collection of sub-fossil bird and marsupial remains from King
Island, Bass Strait. Mem, Nat, Mus. Vic. 3: 5-35.

Stirton, R. A., 1957. Tertiary marsupials from Victoria, Australia. Mem. Nat. Mus. Vic. 21 : 121-34.

Stirton, R. A., Tedford, R. H., and Miller, A. H., 1961. Cenozoic stratigraphy and vertebrate palaeon-
tology of the Tirari Desert, South Australia. Rec. S. Aust. Mus . 14 : 19-61.

Tate, G. H. H., 1947. On the anatomy and classification of the Dasyuridae (Marsupialia). Bull. Amer.
Mus. Nat. Hist. 88: 101-55.

Thomas, O., 1887. On the homologies and succession of the teeth in the Dasyuridae, with an attempt to
trace the history of mammalian teeth in general. Phil. Trans, 128 : 443-62.

Woods, J. T., 1956. The skull of Thylacoleo carnifex. Mem. Qd Mus. 13: 125-40.

MEMOIRS OF THE QUEENSLAND MUSEUM

Plate 1

Dasyurus dunmalli sp. nov.

Fig. 1 : Lateral view of holotype, F6579, Chinchilla Rifle Range (Rifle Range No. 78,
Par. of Chinchilla), SE.Q., x2.

Fig. 2: Stereopair of F6579, x2.

Fig. 3 : Stereopair of F742, Darling Downs, SE.Q., x 2.

Fig. 4: Lateral view of F742, x2.

NEW SPECIES OF FOSSIL MARSUPIAL

Plate 1

Mem . Qd Mus. (1971) 16(1): 27-48, pis. 2-3.

NEW RECORDS AND NEW SPECIES OF CRABS (CRUSTACEA: BRACHYURA)
TRAWLED OFF SOUTHERN QUEENSLAND: DROMIACEA, HOMOLIDEA,
GYMNOPLEURA, CORYSTOIDEA, AND OXYSTOMATA

B. M. Campbell
Queensland Museum

ABSTRACT

Twenty-seven species trawled off southern Queensland are listed and three
new species belonging to the genera Mursia, Ebatia, and Cryptocnemus are
described. Within the groups considered, offshore southern Queensland shows
much greater affinity with Japan than with the adjacent shallower Moreton Bay.

While the crabs of Moreton Bay have been intensively surveyed (see Campbell and
Stephenson, 1970) the area immediately outside the Bay has received little attention.
Several expeditions have collected from northern Queensland (see Dali and Stephenson,
1953, p. 22) but until recently only the F.I.S. ‘Endeavour’ had undertaken exploratory
offshore dredging and trawling in southern Queensland. From this survey, Rathbun (1923)
recorded three crabs belonging to the families under consideration (see List of Species).

In July and August, 1968, the ‘Nimbus’ under the direction of Dr. A. J. Bruce, collected
by trawl from 60 stations off southern Queensland between Double Island Point and Cape
Moreton at distances of up to 40 miles offshore. The depths of the areas investigated ranged
from 25 to 204 fm. The portunid crabs collected by Dr. Bruce were recorded by Stephenson
and Cook, 1970, and this collection also forms the basis of the present report. This material
is supplemented by collections made by Mr. D. Harris on commercial trawling grounds off
Cape Moreton in 60-65 fm, by Mr. F. Wallace from off Caloundra, and by Mr. B. Beutel.

All material is housed in the Queensland Museum; station numbers refer to the
‘Nimbus’ 1/68 cruise; dimensions unless otherwise stated are of carapace width; all drawings
were made with the aid of a camera lucida; and abbreviations are used for crapace width
(cw.) and carapace length (cl.).

28

MEMOIRS OF THE QUEENSLAND MUSEUM

LIST OF SPECIES TRAWLED FROM SOUTHERN QUEENSLAND

Species in bold face have not previously been recorded from off the Queensland coast
south of Bustard Head (24°S.) and are dealt with in this paper. In the case of species not
dealt with the bracketed reference refers to a previous record from this area. Bracketed
numerals and letters following all species refer to their known overall distribution using
the notation listed by Campbell and Stephenson, 1970, p. 295. Species marked thus “f”
were not taken in the present collections.

Tribe DROMIACEA
Family DROMI1DAE

Dromia intermedia Laurie

f Dromidiopsis edwardsi Rathbun [Rathbun, 1923, p. 145
SE. of Double Island Pt., 33 fm]

Cryptodromia areolata Ihle

Tribe HOMOLIDEA
Family HOMOLIDAE

Homola orientalis Henderson
Family LATREILLIDAE

Latreillia australiensis Henderson

Tribe GYMNOPLEURA
Family RANINIDAE

Lyreidus tridentatus de Haan [Griffin, 1970, pp. 94-104,

figs, pi.: N. of Cape Moreton, 20-100 fm

Ranina ranina (Linnaeus)

Tribe CORYSTOIDEA
Family CORYSTIDAE
Gomeza bicornis Gray
Jonas leuteanus Ward

Tribe OXYSTOMATA
Family DORIPP1DAE

Dorippe frascone (Herbst) [Rathbun, 1923, p. 138 (as

D. dorsipes): SE. of Double Island Pt., 33 fm] [2, a.j.]

Family CALAPPIDAE

Calappa japonica Ortmann [2, a.j.]

C. philargius (Linnaeus) [Campbell and Stephenson,

1970, p. 246: Moreton Bay] [l,j.]

[2, a.j.]

[5]

[3, h.j.nz.]
[1, a.j.h.]

[2, a.j.]

[5]

[2, j-]
[ 2 ]

[4,j.]

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

29

Mursia australiensis sp. nov. [5]

Matuta planipes Fabricius [Rathbun, 1923, p. 138: off

Point Inskip, 10 fm] [2, j.]

Family LEUCOSIDAE

Ebalia brevimana sp. nov. [5]

E. longimana Ortmann [4, j.]

Merocryptus lambriformis A. Milne Edwards [3, j. nz.]

Myra kessleri (Paul’son) [2, nc.]

M. affinis Bell [Campbell and Stephenson, 1970,

p. 250: Moreton Bay] [2]

Arcania ( ?) heptacantha (de Haan) [2, j.]

A. undecimspinosa de Haan [2, j.]

A. elongata Yokoya [4, j.]

Pa tidal lia eburnea Alcock [2, j.]

Ixa inermis Leach [4]

Jxoides cornutus MacGilchrist [2, j.]

Leucosia unidentata de Haan [2, j.]

Cryptocnemus hemispheroides sp. nov. [5]

Family DROMIIDAE
Dromia intermedia Laurie

Dromia intermedia Laurie, 1906, p. 351. Ihle, 1913, p. 23, pi. 1, figs. 1-3. Sakai, 1936, pp. 10-11, pi. 6,
fig. 1.

Material: Female, off Caloundra, trawled, R. Elks, W3386.

This specimen agrees well with published figures and descriptions, differing only in
having the anterolateral teeth of the carapace more acute and forwardly projecting.

Distribution: From Ceylon and India to Japan and now Australia.

Cryptodromia areolata Ihle

Cryptodromia areolata Ihle 1913, p. 47, pi. 2, figs. 10-11. Sakai, 1936, p. 26, pi. 1, fig. 1; 1965, pp.
8-9, pi. 3, fig. 4. Takeda and Mijake, 1970, pp. 202-3.

Material: Female, 27°00' S., 153°39' E., trawled 100 fm, ‘Nimbus’ stn 26, 28.vii.1968, W3330. Male,
2 females, off Cape Moreton, trawled 65 fm, D. Harris, W3329. Female, 26°49' S., 153°37' E., trawled
100 fm, ‘Nimbus’ stn 19, 27.vii.1968, W3322.

Distribution: From Timor I. (Ihle, 1913) and Japan (Sakai, 1936, 1965), and now
southern Queensland; 15-100 fm.

30

MEMOIRS OF THE QUEENSLAND MUSEUM

Family HOMOLIDAE
Homola orientalis Henderson

Homola orientalis Henderson, 1888, pp. 19-20, pi. 2, fig. 1. Rathbun, 1923, pp. 143-4, pi. 37. Sakai,
1936, pp. 46-7, pi. 9, fig. 1.

Thelxiope orientalis (Henderson): Sakai, 1965, p. 15, pi. 6, figs. 3, 4 (synon.)

Material: One male, one female, 26°27' S., 153°50' E., trawled 148-9 fm, ‘Nimbus’ stn 55, 5.viii,1968,
W3324.

Distribution: From east Africa to the Philippines, Japan and Australia; previously
within Australia from Bass Strait and Victoria; 50-300 fm.

Family LATREILLIDAE
Latreillia australiensis Henderson

Latreillia australiensis Henderson, 1888, pp. 24-5, pi. 2, fig. 4. Whitelegge, 1900, p. 165. Rathbun,
1923, pp. 139-40.

Material: Male (fragmented), two females, off Cape Moreton, trawled 65 fm, D. Harris, W3327.

Distribution: Southeastern Australia from Bass Strait to Port Jackson and now
southeast Queensland; 30-150 fm.

Family RANIN1DAE

Ranina ranina (Linnaeus)

Cancer raninus Linnaeus, 1758, p. 625.

Ranina dentata de Haan, 1841, p. 139, pis. 34, 35, figs. 1-4. Haswell, 1882, p. 144.

Ranina ranina (Linnaeus): Sakai, 1937, p. 178, pi. 16, fig. 4; 1965, p. 4, pi. 2, fig. 1 . Barnard, 1950, pp.
397-8, fig. 75a-d. Healy and Yaldwyn, 1970, p. 76, pi. 36.

Material: Male, between Cape Moreton and Mooloolaba, trawled, F. Wallace, W3389. Male, 8 miles
SE. of Cape Moreton, W1964.

Although this species was not recorded from Moreton Bay by Campbell and Stephen-
son, 1970, there are several specimens in the Queensland Museum collections taken from
Moreton Bay, Bribie I., and Amity.

Distribution: From South Africa to Japan, Hawaii, and Australia. Within Australia
from western, northern, and eastern Australia to southern New South Wales.

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

31

Family CORYST1DAE
Gomeza bicornis Gray

Gomeza bicornis Gray, 1831, p. 39. Hale, 1927, pp. 145-6, fig. 147. Barnard, 1950, p. 305, fig. 57d-g.

Material: Female, between Cape Moreton and Mooloolaba, trawled, F. Wallace, W3390. Female,
5-6 miles E. of Pt Cartwright Light, trawled 19-21 fm, sand, coral and shell, 6.iii.l970, F. Wallace, W3391 .

Distribution: From South Africa to Ceylon, Japan, and within Australia from
South Australia and southeast Queensland.

Jonas leuteanus Ward

Jonas leuteanus Ward, 1933, pp. 379-80, pi. 23, fig. 8.

Material: Female, 5-6 miles E. of Pt Cartwright Light, trawled 19-21 fm, sand, coral and shell,
7.iii.l970, F. Wallace, W3392.

Distribution: Lindeman I., Cumberland Group, Queensland (type locality) and
southeast Queensland.

Family CALAPPIDAE
Calappa japonica Ortmann

Calappa japonica Ortmann, 1892, p. 566, pi. 26, fig. 8. Sakai, 1937, p. 96, pi. 18, fig. 4; 1965, p. 57, pi.
23, fig. 1. Barnard, 1950, pp. 352-3, fig. 66 n-p.

Calappa exanthematosa Alcock and Anderson, 1894, pp. 177-8. Alcock, 1899, pp. 21-2.

Material: Male, off Cape Moreton, trawled 70 fm, B. Beutel, March 1966, W2414. Female, off Cape
Moreton, trawled 65 fm, D. Harris, W3362 (dry coll.).

Distribution: South Africa to India, Japan, and now Australia; 17-70 fm.

Mursia australiensis sp. nov.

(Fig. 1 ; pi. 2A, B)

Mursia annata de Haan: Whitelegge, 1900, pp. 160-1.

[non] Mursia armata de Haan, 1837, p. 73, pi. 19, fig. 2.

[?] Mursia spinimanus Rathbun: Rathbun, 1911, p. 198, pi. 15, fig. 3.

[non] Mursia spinimanus Rathbun, 1906, p. 888, pi. 16, fig. 1.

Holotype: Male, 26 mm cl., 42.8 mm cw. including spines, 31 mm cw. excluding spines, off Cape
Moreton, trawled, April 1964, D. Harris, W2379.

32

MEMOIRS OF THE QUEENSLAND MUSEUM

Paratypes: Two males, 21, 22 mm cl., 26°31' S., 153 °40' E. ( c . 30 miles NE. of Caloundra), trawled
75.5 fm, ‘Nimbus’ stn 10, 26.yii.1968, A. J. Bruce, W3297. Male, 25.5 mm cl., 26°48' S., 153°32.5' E. (c.
20 miles E. of Caloundra), trawled 55 fm, ‘Nimbus’ stn 17, 27.vii.1968, A. J. Bruce, W3298. Three males,
24, 25, 29 mm cl., 1 8 miles N, of Cape Moreton, trawled 62-5 fm, sand and shell, FV ‘Gemini’, 19-20.iii.1970,
F. Wallace, W3299. Three males, 28-29 mm cl., off Cape Moreton, trawled 65 fm, D. Harris, W3300 (dry
coll.). Ten males, 16.5-27. 5 mm cl., female, 23 mm cl., off Cape Moreton, trawled 65 fm, D. Harris, W3301 .

Description

Carapace broader than long (cw. excluding lateral spines 1.15 to 1.18 times cl. in
males, 1.13 times in only available female); coarsely granulate over entire surface (except
frontal region which is less distinctly granulate) with 7 rows of tubercles radiating back-
wards from behind fronto-orbital region ; front narrow (5 . 4-5 . 9 times in cl., 2 . 5-2 . 8 times
in fronto-orbital width), three lobed, middle lobe projecting well forward of lateral lobes
variably separated from the lateral lobes (see fig. lc); anterolateral margins very convex,
sub-parallel in posterior fifth, with some ten distinct sharp granules decreasing in size pos-
teriorly ; lateral spine moderately long (3 . 4-4 . 2 times in cl. in males, 4.75 times in female),
slender, directed slightly posteriorly (posterior margin of spine lies in front of a line joining
tips of spines), curved markedly or only slightly upwards (maximum of c. 30° to horizontal
in posterior view, almost horizontal in female); posterolateral margins slightly shorter

Fig. 1 : Mursia australiensis . A-C, variation in posterior carapace margin (A), abdominal crest (B), and
front (C); D, third maxilliped; E, ventral view of front and orbit; F, second (right) male pleopod;
G, first (right) male pleopod.

Specimens illustrated: A-C, 1 = W3298, 2 = W3301, 3 = W3299, 4 = W3300; D-G, holotype.
Scale divisions 1 mm.

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

33

than or subequal to anterolateral margins; posterior margin short (4. 6-5. 4 times in cl.),
with three spines of which the laterals are largest (larger than anterolateral spinules) the
median varying from only slightly smaller than the laterals to a low obtuse protruberence.

Inner suborbital lobe triangular, with outer margin straight; separated from outer
orbital cup by a V-shaped sinus.

Merus of cheliped with three spines distally, the upper smallest but always distinct
(larger than anterolateral spinules), the lower largest (at least 2/3 lateral spine). Upper face
of wrist and outer face of chela coarsely granulate ; outer face of chela with 9 tubercles in
3 rows and 3 further tubercles near base of dorsal serrate crest; lower row with proximal
tubercle very acute and highest (triangular with concave sides), second tubercle lower and
broader based (triangular with convex sides), distal tubercle low, rounded, and granulate;
lower margin of chela finely serrate, becoming coarser distally ; inner face of dactyl with
row of some 25-30 stridulatory tubercles becoming smaller proximally and distally.

Ambulatory legs long (first leg 1 .5-2. 1 times cl.).

Abdomen of both sexes with three subequal flattened lobes on second segment well
separated by V- or U-shaped sinus and with finely granulate margins. Penultimate segment
of male abdomen as long as wide, lateral margins subparallel but slightly sinuous with slight
protruberences at latero-distal angles; ultimate segment longer than wide, triangular, with
slightly concave lateral margins.

First male pleopod stout, evenly tapering to a blunt tip, with fine spinules distally.
Second male pleopod long, slender, with distal third abruptly narrower, horny, sinuous,
with extreme tip conical, pointing laterally.

Colour, after alcohol preservation, pale biscuit with varying amount of rose pink on
carapace sometimes concentrated in granules and often darker towards the lateral spines,
on the frontal region, and in the upper outer face of the chelae. Inner face of chelae with
darker red patch at base of dactyl.

Discussion

Sakai (1965, pp. 51-5) lists the 7 known Indo-Pacific species and subspecies of this
genus and describes or figures 5 of these.

M. armata differs from other species, including the present one, in having the antero-
lateral carapace margins only slightly convex, the lateral teeth more than half the carapace
length. In M. curtispina, M. trispinosa, M. aspera , and M. hawaiiensis the second male
pleopod is considerably recurved distally, forming a complete loop in M. trispinosa . In the

34

MEMOIRS OF THE QUEENSLAND MUSEUM

present species the second pleopod is slightly undulating distally, outcurved at the tip.
The ornamentation of the outer face of the chela of the present species differs markedly
from Sakai’s figures (fig. 8a-d) of these four species.

M. bicristimana Alcock and Anderson (see Alcock, 1899, pp. 23-4, pi. 3, fig. 3) differs
from M. australiensis in having 2, not 3, teeth on the posterior carapace margin, the lateral
carapace teeth forwardly directed ; the three lobes of the second abdominal segment barely
separated by narrow grooves, the median lobe much wider than the laterals; and the three
ventral teeth of the outer face of the chela joined by a crest.

M. spinimanus differs in having the posterolateral margins of the carapace much more
convergent posteriorly, the surface of the carapace less distinctly tuberculate and more
finely granulate, the carapace broader (cw. 1 . 3 times cl.) and the suborbital notch V-shaped.
Rathbun (1911, p. 198) identified a small female from Saya de Malha (Indian Ocean) as
M. spinimanus. This specimen differs from M. spinimanus in granulation and tuberculation
of carapace and in carapace shape. It is tentatively placed in synonymy with the present
species, the only difference observable from Rathbun’s figure (pi. 15, fig. 3) being the
straight (as opposed to slightly curved) lateral carapace teeth.

M. australiensis is closest to M. curtispina Miers but comparison with Miers’s figure
and description (1886, pi. 24, fig. 1, pp. 291-2) shows the following differences:

(1) Miers’s figure shows a much broader front (3 .75 times in cl. as against 5 .4 to

5.9 times in M. australiensis ).

(2) The lateral carapace spines are shorter in Miers’s figure (6 times in cl. as against

3 .4 to 4.7 times in M. australiensis ) and are not obliquely directed backwards.

(3) The suborbital notch is narrow and V-shaped in Miers’s fig. 2a. In M. australien-

sis it is broadly V-shaped and the suborbital tooth is triangular.

(4) The teeth on the posterior carapace margin are smaller and more widely

separated in Miers’s figure, the posterior margin is shorter (3 .9 times in cl. as
against 4.6 to 5.4 times in M. australiensis).

Sakai (1965) has compared Miers’s holotype with specimens from Japan but although
he identifies the Japanese specimens with Miers’s species his figure (pi. 21, fig. 2) shows
the following differences from that of Miers.

(1) Front 5.0 times in cl.

(2) Lateral carapace spines 4.6 times in cl.

(3) Posterior surface of carapace indistinctly granulate.

(4) Posterior margin of carapace shorter (4.9 times in cl.).

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

35

These differences could be due to subspecific variation within M. curtispina or to in-
accuracies in Miers’s illustration.

The Japanese specimens described by Sakai differ from M. australiensis in the following
particulars:

(1) Front 5 .0 times in cl. as against 5 .4-5.9 times.

(2) Posterior surface of carapace indistinctly granulate.

(3) Posterior carapace teeth smaller and more widely separated.

(4) Two distal spines of lower series of outer face of chela are more acute and more

prominent.

(5) The second male pleopod is strikingly recurved distally,

(6) The lateral carapace spines are less obliquely directed backwards.

Until material from a wider range of localities throughout the central Indo-west Pacific
is compared, and the presence or absence of intergradation between Japanese and Fijian
M. curtispina and M. australiensis is demonstrated it could be considered that the latter
is best regarded as a subspecies of the former. The decision to treat M. australiensis as
specifically distinct was based, in part, on the degree of difference which exists between the
five species of Mursia recorded by Sakai (1965) as occurring together in Japan. Although
two of these (M. trispinosa Parisi and M. hawaiiensis Rathbun) were regarded by Sakai as
subspecies of M. curtispina , their coexistence without apparent intergradation requires
their recognition as distinct species. The differences between these five species in such
features as carapace shape, chela ornamentation and second male pleopod configuration
are no greater than the differences between M. australiensis and M. curtispina listed above.

Family LEUCOSJDAE

Ebalia brevimana sp. nov.

(Fig. 2A-H; pi. 3 A)

Holotype: male, cw. 8.4 mm, 26°27' S., 153°50' E., trawled 148 fm, ‘Nimbus' stn 55, 5,viii.l968,
A. J. Bruce, W3395.

Description

Carapace slightly longer than broad (cl . 8 . 6 mm, cw. 8 . 4 mm) with regions indistinctly
defined by shallow grooves, with six tubercles disposed as in figure 2A (one intestinal, one
cardiac, two branchial and two gastric); hepatic regions with low rounded swelling overlying
the small anterolateral tooth; branchial regions swollen, rounded, giving the carapace an

36

MEMOIRS OF THE QUEENSLAND MUSEUM

Fig. 2 : A-H. Ebalia brevimana, holotype. A, carapace, dorsal view ; B, carapace, lateral view ; C, mushroom-
shaped granules on carapace; D, third maxilliped; E, front; F, first male pleopod (1, abdominal
face, 2, median face; 3, median face; 4, sternal face; 5, left pleopod in situ, latero-abdominal face);
G, second male pleopod; H, male abdomen. Scale divisions 1 mm.

I, male pleopod, E. tuberculosa , 8.5 mm cw., Aust. Mus. P5524.

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

37

almost circular outline in dorsal view; front with two rounded lobes separated by shallow
median depression; frontal lobes very rounded in lateral view; posterior margin with two
rounded lobes laterally, little wider than frontal lobes, shallowly separated by the concave
posterior margin which is partly obscured in dorsal view by the large intestinal protruber-
ance. Surface of carapace with large scattered fimbriate mushroom-like granules which are
confluent on the larger carapace tubercles, interspersed with fine scattered hairs on low
granules.

Chelipeds short, 1 .25 times carapace length, propodus as long as merus, dactylus only
slightly shorter than palm, length of palm c. 1.5 times height. Surface of all segments
smooth, formed by confluent discoid fimbriate tubercles. Both fixed and movable fingers
with thin keel on outer edges, more prominent on third finger; and with raised longitudinal
row of granules extending to hooked tips ; teeth fine, interlocking distally with narrow gape
proximally. Male abdomen with first and second segments distinct; third to sixth fused,
with sharp spinous tubercle distally; seventh segment elongate, with lateral margins sub-
parallel for most of their length, tip paraboloid.

Third maxilliped merus approximately half total length of ischium, c. 0.65 times
marginal length. Maxillipeds (and whole ventral surface) covered with mushroom-shaped
granules not elongated into spines anteriorly. First male pleopod with almost straight shank,
tapering evenly, and flattened distally proximal to the asymmetrically bilobed aperture;
with moderate setae on the lateral face and very short setae distally on the sternal face.
Second male pleopod short, with terminal process almost as long (c. 0 .9 x) as shank.

Colour, after alcohol preservation, faded to pale biscuit with few small pale orange
spots scattered on carapace.

Discussion

This species, with its six dorsal carapace tubercles, is close to E. tuberculosa (A. Milne
Edwards) (see Haswell, 1880, p. 54, pi. 6, fig. 3, as Phlyxia granulosa; Miers, 1886, p. 306,
pi. 25, figs. 1, la; Hale, 1927, p. 197, fig. 198; Rathbun, 1923, p. 134, pi. 35, figs. 1-2;
Sakai, 1937, pp. 111-2, fig. lla-d; 1965, p. 28, pi. 13, fig. 2; Barnard, 1950, p. 368, figs.
70h-k; Takeda and Mijake, 1970, pp. 211-2 (synon.)). It differs from that species in the
following particulars.

(1) Carapace more rounded in dorsal view, front less produced (variable in E.

tuberculosa but not as in present species, see Barnard, 1950, and compare
Sakai, 1965 with Miers, 1886).

(2) Posterior carapace lobes indistinctly separated, with intestinal prominence over-

lapping posterior margin in dorsal view (variable in E. tuberculosa but not
approaching present species; compare Sakai, 1965, with Hale, 1927).

38 MEMOIRS OF THE QUEENSLAND MUSEUM

(3) Chelipeds are shorter, chelae shorter in relation to finger length and to height

of chelae (variable in E. tuberculosa but not approaching present species;
compare Sakai, 1937, with Miers, 1886, Hale, 1927, Sakai, 1965).

(4) Male abdomen with sixth segment fused to preceding segments, with tubercle

on sixth segment. (Sakai, 1937, and Barnard, 1950, fig. j (side view) show
distinct sixth segments whereas Haswell, 1880, and Barnard, 1950, fig. j (ventral
view) show fused segments. Sakai and Barnard show a tubercle on the posterior
margin of the seventh segment. Haswell shows one on the sixth segment.
While Barnard’s ventral view figure is obviously in error (see also Stebbing,
1920, pi. 26a; 1921, pi. 18a) Haswell’s figure suggests a close agreement in this
feature with the present species although in all other respects his figure re-
presents typical E. tuberculosa. The type material of Haswell’ s Phlyxia granu-
losa consists of four dried specimens mounted on glass plates. Dr D. J. G.
Griffin of the Australian Museum, Sydney, reports that the abdomen of the
single male can not be examined because of the mounting medium; that these
specimens appear to be conspecific with specimens of E. tuberculosa from the
same locality (off Sydney) in which the abdominal formula of males is
1 H~2 + R+ 6 + T; and that the type specimens are not conspecific with
the present species. It would appear that Haswell’s drawing of the male
abdomen is incorrect).

(5) First male pleopod with straight, evenly tapering shaft, with bilobed tip. (Sakai,

1937, and Barnard, 1950, illustrate male pleopods of specimens of E. tuber-
culosa from Japan and South Africa respectively. These agree essentially with
each other and with specimens of E. tuberculosa from off Botany Bay, N.S.W.,
illustrated in fig. 21).

E. salamensis Doflein, 1904; E. japonica Rathbun, 1932; Nursia scandens Stebbing,
1920; and N. postulans Stebbing, 1921, have been synonymised with E. tuberculosa (see
Sakai, 1965). Examination of the original descriptions of these nominate species show
that they do not belong to the present species and are correctly synonymised with E.
tuberculosa.

Ebalia longimana Ortmann
(Fig. 3)

Ebalia longimana Ortmann, 1892, p. 578, pi. 26, fig. 13. Sakai, 1937, pp. 109-11, fig. 10, pi. 13, fig. 6;
1965, pp. 29-30, pi. 13, figs. 5, 6 (synon.).

Material: male, 4.5 mm cl., with deformity of left branchial region, 26°30' S., 153 c 44' E., trawled
100 fm, ‘Nimbus’ stn 49, A. J. Bruce, 30.vii.1968, W3396.

This small specimen agrees well with figures and descriptions of Ortmann and Sakai,
differing in having the male pleopods less slender, the chelipeds shorter than in Sakai’s

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

39

illustration of male specimens, approaching those of the illustrated female (Sakai, 1965,
pi. 13, fig. 6). Both differences are considered due to the smaller size of this individual.

Distribution: Previously recorded only from Japan.

Merocryptus lambriformis A. Milne Edwards

Merocryptus lambriformis A. Milne Edwards, 1873, p. 85, pi. 13, figs. 1-1 c. Whitelegge, 1900, p. 162
Rathbun, 1923, p. 133, pi. 32, figs. 2, 3. Sakai, 1937, pp. 113-4, pi. 13, fig. 7; 1965, p. 30, fig. 30,
pi. 14, figs. 1, 2. Serene, 1955, p. 145, figs. 1, 2. Bennett, 1964, p. 22, fig. 108.

Material: Male, 26°49' S., 153°37' E., trawled 100 fm, ‘Nimbus’ stn 19, 27.vii.l968, W3353. Female,
off Cape Moreton, trawled, 22.vii.1965, B. Beutel, W2255.

Distribution: Japan to Samoa, Australia, and New Zealand. Within Australia from
the Great Australian Bight, Bass Strait, Port Hacking, Crowdy Head, and now Queensland;
22-120 fm.

Myra kessleri (Paul’ son)

Callidactylus kessleri Paul’son, 1875, p. 80.

Myra kessleri (Paul’son): Ihle, 1918, p. 260 (synon.). Tyndale-Biscoe and George, 1962, p. 89, figs. 7, 9.
Myra darnleyensis Haswell, 1880, p. 52, pi. 5, fig. 4.

Fig. 3: Ebalia longimana. A, carapace; B, male abdomen; C, first male pleopod; D, second male pleopod.
Scale divisions 1 mm.

40

MEMOIRS OF THE QUEENSLAND MUSEUM

Material: Male, 25°or S., 153°30 / E., 29 fm, Bureau of Mineral Resources stn 1377, W3384.

Distribution: From Red Sea (Paul’son 1875) and Seychelles to Indonesia, Australia,
and New Caledonia. Previous Australian records are from Dampier Archipelago (Western
Australia) and Torres Strait (fide Tyndale-Biscoe and George, 1962).

Arcania (?) heptacantha (de Haan)

(PI. 3B)

Jphis heptacantha de Haan, 1861, p. 27.

Arcania heptacantha (de Haan): de Man, 1907, pp. 398-9, pi. 31, figs. 8-10. Sakai, 1937, pp. 126-7,
fig. 18; 1965, p. 41, pi. 16, fig. 6.

Material: Female, 26°30' S., 153°15' E., trawled 26 fm, ‘Nimbus’ stn 8, 26.vii.1968, W3359. Female,
Arnhem Bay, Northern Territory, 10 fm, sand and mud bottom, V. Wells, W2604.

Published figures and descriptions demonstrate some confusion as to the relative
status of A. heptacantha and A. septemspinosa (Fabricius). De Man (1907, pp. 398-9) in
attempting a comparison could only suggest that the relative lengths of carapace spines
(although shown to be variable by Alcock), and perhaps the depth of the groove separating
branchial from cardiac and intestinal regions, were the most likely features that could be
used to distinguish the two species. Rathbun (1910, p. 314) described her A. siamensis
(synonymised with A. heptacantha by Sakai, 1937, 1965) as differing from A. septemspinosa
(which she recorded from the same locality) in having the carapace more subglobular, the
branchial regions more swollen, and the marginal spines shorter than in A. septemspinosa.

Barnard (1950, p. 385, fig. 71, 7 If, g) describes and illustrates specimens which, al-
though he identifies them as A. septemspinosa , have short carapace spines. Barnard’s
specimens are also unusual in having the lateral spines situated on the posterior third of
the carapace, the lower posterior spines very close together under the base of the median
posterior spine, and the anterolateral margins straight or almost concave. The four very
short posterolateral spines of Barnard’s illustration bear no resemblance to the long spines
figured for A. septemspinosa by Sluiter (1881, fig. 1) and by Herbst (1790, pi. 20, fig. 1 12).

The Australian specimens can not be satisfactorily fitted into the variation shown by
these (or this) species without detailed comparisons of material from a range of localities
throughout the Indo-Pacific area. They appear closest to A. heptacantha as figured by
Sakai (1937, 1965) but differ from his specimens in having the lateral carapace spines
reaching to the end of the ambulatory meri, the front narrower and more anteriorly project-
ing (well forward of the lateral pterygostomial spines), and the carapace regions less
distinctly marked.

Distribution: A. heptacantha from the Gulf of Thailand to Japan, and now Australia.
A. septemspinosa from South Africa (? identity) to India and Hong Kong.

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

41

Arcania undecimspinosa de Haan

Arcania undecimspinosa de Haan, 1841, p. 135, pi. 33, fig. 8. Sakai, 1937, p. 123, fig. 15a, pi. 14, fig. 2;

1965, p. 40, fig. 6a, pi. 16, fig. 3.

Arcania granulosa Miers, 1877, p. 240, pi. 38, fig. 29.

Material: Two males, female, off Cape Moreton, trawled 65 fm, D. Harris, W3348 (dry collection).
Two males, 26°31' S., 153°43' E., trawled 100-102 fm, ‘Nimbus’ stn 11, 26.vii.1968, W3358. Four males,
two females, off Cape Moreton, trawled 65 fm, D. Harris, W3352. Two females, off Cape Moreton, trawled,
B. Beutel, 22.vii.1965, W2256. Female, 26°49' S., 153 °35' E., trawled 75 fm, ‘Nimbus’ stn 18, 27.vii.1968,
W3360. Female, 26°49' S., 153°37' E., trawled, 100 fm, ‘Nimbus’ stn 19, 27.vii.1968, W3356. Male, off
Caloundra, trawled, R. Elks, W3349.

Distribution: India to Japan and Australia. Previously recorded from Australia by
Miers (1877) from ‘Moreton B.\ but not recorded from within the bay by Campbell and
Stephenson, 1970.

Arcania elongata Yokoya

Arcania undecimspinosa elongata Yokoya, 1933, p. 132, fig. 47. Sakai, 1937, p. 124, figs. 15b, 16; 1965,
pp. 40-1, fig. 6b, pi. 16, fig. 2.

Material: Female, 26°30' S., 153°15' E., trawled, 26 fm, ‘Nimbus’ stn 8, 26.vii.1968, W3358. Male,
southern Queensland, E. C. Vallis, W3347 (dry collection).

These specimens differ from A. undecimspinosa in all features mentioned by Sakai
(1937, pp. 124-5; 1965, pp. 40-1), approaching A. novemspinosa in many respects but differ-
ing from that species in the sharper granulation (spinulation) of the carapace, more elongate
carapace, smaller marginal spines, less projecting front (which is finely spinulate not
granulate), absence of prominent proximal spine on posterior margin of arm of cheliped.

In view of the constancy of the distinguishing features over such a wide geographic
range A. elongata must be given full specific status.

Distribution: Previously known only from Japan (Sakai, 1965).

Randallia eburnea Alcock

Randallia eburnea Alcock, 1896, p. 196. Sakai, 1937, pp. 132-3, fig. 22; 1965, p. 42, pi. 17, fig. 1. Tyn-
dale-Biscoe and George, 1962, pp. 87, 94.

Material: Male, 30 miles due E. of Mooloolaba, 69-70 fm, R. Elks, W2849. Female, 26°31' S.,
153°33' E., trawled 56 fm, ‘Nimbus’ stn 9, 26.vii.68, W3355. Female, 26°49' S., 153°35' E., trawled 75 fm,
‘Nimbus’ stn 18, 27.vii.1968, W3354. Six males, southeast Queensland, trawled, D. Harris, W3344 (dry
collection). Female, off Caloundra, trawled, R. Elks, 1 3.viii. 1967, W3350. Two males, female, 28 miles
N. of Cape Moreton, trawled 62-5 fm, sand and shell, F, Wallace, 7.iv.l970, W3364. Male, 18 miles N. of
Cape Moreton, trawled 62-5 fm, sand and shell, F. Wallace, 19.iii.1970, W3368. Male, 5 females, off Cape
Moreton, trawled 65 fm, D. Harris, W3380.

42

MEMOIRS OF THE QUEENSLAND MUSEUM

These specimens agree with those of Tyndale-Biscoe and George in that the distal
third of the first to third dactyls have long hairs and the tooth on the penultimate segment
of the male abdomen is well developed. They differ, agreeing with Ihle’s description, in
having the fused 3rd to 6th abdominal segments all clearly recognisable.

Distribution: From India and the Laccadives to Japan, Western Australia, and now
Queensland; 15-80 fm.

Ixa inermis Leach

Ixa inermis Leach, 1817, p. 26, pi. 129, fig. 2. Haswell, 1880, p. 59. McNeill, 1942, p. 430; 1968, p. 40.
Holthuis and Gottlieb, 1956, pp. 291-6, pi. 5, fig. 1. Healy and Yaldwyn, 1970, p. 82, fig. 43.

[non] Ixa inermis Leach: Alcock, 1896, pp. 272-3.

Material: Male, 26°45' S., 153°2L E., trawled 25 fm, ‘Nimbus* stn 16, 27.vii.1968, A. J. Bruce, W3397.
Male, off Cape Moreton, trawled 65 fm, D. Harris, W3398. Male, female, off* Cape Moreton, trawled 65 fm,
D. Harris, W3399 (dry collection). Male, 6 miles NE. of Caloundra Light, trawled 21 fm, sand and shell,

27. 11. 1970, F. Wallace, W3374. Female, 6-7 miles NE. of Caloundra Light, trawled 22 fm, sand and shell,

3.111.1970, F. Wallace, W3373. Female, Hervey Bay, Capt. Hoult, G646.

Distribution : From the Malay Archipelago and Australia. Previously within Australia
from Thursday Island, Low Isles, Cape Grenville, Port Denison, and off Tin Can Bay,
25 fm.

Ixoides cornutus MacGilchrist

Ixoides cornutus MacGilchrist, 1905, p. 255. Sakai, 1937, pp. 137-9, pi. 19, figs. 1-4; 1965, p. 44, pi. 18,
fig. 3.

Material: Five males, off Cape Moreton, trawled, D. Harris, W3343 (dry collection). Male, female,
off Cape Moreton, trawled 65 fm, D. Harris, W3361. Male, female, off Cape Moreton, trawled, B. Beutel,
22.vii.1965, W2257. Male, 28 miles N. of Cape Moreton, trawled 62-5 fm, sand and shell, F. Wallace, 7.iv.
1970, W3365. Male, female, 18 miles N. of Cape Moreton, trawled 62-5 fm, sand and shell, F. Wallace,
1 S.iii. 1 970, W3369.

Distribution: From the Iranian Gulf to Hong Kong and Japan (Sakai, 1965), and
now eastern Australia.

Leucosia unidentata de Haan

Le ucosia unidenlata deHaan, 1841, p. 133, pi. 33, fig. 3. Haswell, 1882, p. 118. Alcock, 1896, pp. 215-6.
Sakai, 1937, p. 146, fig. 286, pi. 15, fig. 4; 1965, pp. 47-8, pi. 19, fig. 3.

Material: Female, 18 miles N. of Cape Moreton, trawled 62-5 fm, sand and shell, F. Wallace, 19.iii.
1970, W3367. Female, 28 miles N. of Cape Moreton, trawled 62-5 fm, sand and shell, F. Wallace, 7.iv.l970,
W3366. Male, 26°3L S., 153°33' E., trawled 56 fm, ‘Nimbus’ stn 9, 26.vii.1968, W3378. Male, 26°17' S.,
153°42' E., trawled 47 fm, ‘Nimbus’ stn 2, 25.vii.1968, W3383. Female, 26°48' S., 153°32.5' E., trawled

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

43

55 fm, ‘Nimbus’ stn 17, 27,vii.l968, W3379. Male, off Caloundra, trawled, R. Elks, W3377. Male and
female, off Cape Moreton, trawled 65 fm, D. Harris, W3376. Fifteen males, five females, trawled off
southern Queensland, D. Harris, W3388 (dry collection).

Alcock’s description and Sakai’s figures indicate that the posterior margin of the
carapace is very convexly rounded. While this is true in the females of the present series,
in the males this margin is concave, with two slight, well rounded lobes laterally. The pleo-
pods agree well with Sakai’s (1937, fig. 28b) illustration except that the distal bristles are
much longer (as long as the terminal projection) and the three spirals of the shaft of the
pleopod are not as distinctly divided to form six as in Sakai’s figure.

Distribution: From India to Hong Kong, Japan, Molluccas, Torres Strait, and now
southern Queensland.

Cryptocnemus hemispheroides sp. nov.

(Fig. 4; pi. 3C)

Holotype: female, 8 mm cw., 26°2T S., 153 c 50 / E., trawled 148 fm, ‘Nimbus’ stn 55, 5.viii. 1 968, W3363.
Description:

Carapace broader than long (cw. = 8 mm, cl. = 7 . 1 mm), smooth, dorsally rounded,
without ridges or granules on the dorsal surface; margins of carapace with lamellate
expansions originating at a distinct tooth on the anterolateral margin, increasing in width
posteriorly, edge composed of straight lines which meet to form distinct angles laterally,
posterolaterally and at each side of the slightly concave posterior margin.

Front markedly produced, deeply divided into two slightly upturned lobes separated
by U-shaped sinus in dorsal view; small but distinct subsidiary lobes halfway along length
of median margins of frontal lobes.

Median portion of front between frontal lobes markedly depressed to meet epistome,
separating antennulary fossae which extend well up under frontal lobes; antennulary fossae
separated from orbits by second segment of antennae; suborbital lobe large.

Anterior extremity of buccal cavern reaches well beyond anterior extremity of ptery-
gostomian region. Merus of third maxilliped much shorter than ischium.

Chela approx. 1 .6 times cw. Merus flattened in proximal half with thin flat laminate
carinae on both margins, trigonal in distal half with thin carinae on all three margins;
carpus trigonal with broadly lamellate carina on outer margin, ridge ending in small distal

44

MEMOIRS OF THE QUEENSLAND MUSEUM

tooth on upper inner margin, low ridge ending in larger tooth on lower inner margin;
propodus flattened, with broad lamellate carina on outer margin and on inner margin
extending along edge of fixed finger decreasing in width to near the tip; dactylus as long as
propodus excluding fixed finger, with lamellate carina on outer margin decreasing in width
to near the tip ; cutting edges of fingers with series of low teeth with larger ones interspersed
at regular intervals, these latter increasing in size towards the tip on the fixed finger,
subequal on the dactylus.

Walking legs decreasing in size posteriorly; merus and propodus flattened, with lamel-
late carinae on both margins ; carpus of first two legs with broad carina on outer margin,
of last two legs with two carinae, one on outer margin, one on posterior (upper) face;
dactyl very thin, styliform.

Abdomen of female with free segment followed by four discernable segments fused
in a circular plate, with terminal segment free, triangular, longer than broad.

Colour, after alcohol preservation, pale, with pale pink-orange spots, one in middle
of posterior margin, two posteriorly on dorsal surface with indistinct broken band running
anterolaterally from these towards the anterolateral teeth; small white speckles running
posterolaterally from orbits in band which ends in medial expansion at level of anterolateral
teeth, similar white spots on upper face of cheliped.

Fig. 4: Cryptocnemus hemispheroides. A, carapace, frontal and lateral view; B, carapace, dorsal view;
C, front; D, third maxilliped; E, female abdomen. Scale divisions 1 mm.

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

45

Discussion

Ihle (1918, p. 317) listed the known Indo-Pacific species of Cryptocnemus and (p. 286)
provided a key to these species. To this list must be added the subsequently described
C. vincentianus Hale, 1927, C. planus Ward, 1933, and C. kamekii Sakai, 1961.

The present species is similar to C. obolus Ortmann, 1892, in having the front very
distinctly bilobate and the carapace surface smoothly hemispherical without ridges whereas
in all other species of Cryptocnemus the front is variably produced as a single triangular or
rounded lobe, and the carapace usually bears a median ridge from between the orbits to
the centre of the carapace (faint ridge extending a short distance back in C. planus) with
sometimes additional posterolaterally radiating ridges from that point. While these differ-
ences also are possibly marked enough to warrant generic separation of C. obolus and
C. hemispheroides from other species of Cryptocnemus , examination of all species would be
desirable to determine other significant distinguishing features.

C. hemispheroides differs from C. obolus (fide, Ortmann, 1892, p. 576, pi. 26, fig. 12;
Yokoya, 1933, p. 117; Sakai, 1937, pp. 140-1; 1965, p. 45, pi. 18, fig. 2) in that the two
frontal lobes are less acutely projecting and have small subsidiary lobes on their median
margins ; the expanded carapace edge is not produced into as large a lobe laterally, and the
posterior half of this edge is not smoothly rounded but composed of five straight or even
slightly concave segments meeting to form distinct angles; the dorsal crest on the anterior
margin of the arm of the cheliped is much shorter, occupying only the distal half of the arm.

DISCUSSION

The 27 species of Dromiacea, Homolidea, Gymnopleura, Corystoidea and Oxystomata
thus far known from this area probably represent only a small fraction of the total fauna,
and only broad comparisons can be made with other areas at this time. Of these 27 species,
7 (26%) are not found outside Australia, 15 (56%) are shared with India and 18 (67%)
with Japan. In contrast, of the 18 species of these groups found in Moreton Bay (see Camp-
bell and Stephenson, 1970) 5 (28%) are not found outside Australia, 12 (67%) are shared
with India but only 4 (22 %) with Japan.

Only 4 (15%) of the 27 species found in the deeper water (>20fm) off southern
Queensland are also present in the predominantly shallow water (<20 fm) of Moreton Bay.
Although further collecting may alter these figures it would seem that faunal composition
is affected more markedly by a depth change of 20 fm than by spatial separation of some
6000 miles.

46

MEMOIRS OF THE QUEENSLAND MUSEUM

LITERATURE CITED

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1899. ‘An Account of the Deep-Sea Brachyura collected by the Royal Indian Marine Survey Ship
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Alcock, A. and Anderson, A. R., 1 894. Natural history notes from H.M. Indian Survey Steamer ‘Investiga-
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Barnard, K. H., 1950. Descriptive catalogue of South African decapod Crustacea (crabs and shrimps).
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Doflein, F., 1904. Brachyura. In ‘Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem
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Holthuis, L. B. and Gottlieb, E., 1956. Two interesting crabs (Crustacea Decapoda, Brachyura) from
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CRABS TRAWLED OFF SOUTHERN QUEENSLAND

47

Ihle, J. E. W., 1913. DieDecapoda Brachyura der Siboga-Expedition. I. Dromiacea. SibogaExped. Monogr.
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Laurie, R. D., 1906. Report on the Brachyura collected by Professor Herdman, at Ceylon, in 1902. In
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233-68.

McNeill, F. A., 1942. A crab wonder. Aust. Mus. Mag. 7 (12): 430.

1968. Crustacea, Decapoda and Stomatopoda. Sci. Rep. Gr. Barrier Reef Expect. 7 (1): 1-98, pis. 1, 2.

Man, J. G. de, 1907. On a collection of Crustacea, Decapoda and Stomatopoda chiefly from the Inland
Sea of Japan, with descriptions of new species. Trans. Linn. Soc. Lonct. ( Zoot .) (2) 9 (11): 387-454,
pis. 31-3.

Miers, E. J., 1877. Notes upon the oxystomatous Crustacea. Trans. Linn. Soc.Lond. ( Zoot .). (2) 1 (5): 235-49,
pis. 38-40.

1886. Report on the Brachyura collected by H.M.S. ‘Challenger’ during the years 1873-1876. In ‘Report
on the Scientific Results of the Voyage of H.M.S. “Challenger” during the years 1873-76 under
the command of Captain George S. Nares, R.N., F.R.S. and the Late Captain Frank Tourle
Thomson, R.N.’ Zoology, vol. 17, part 2, pp. i-1, 1-362, pis. 1-29.

Milne-Edwards, A., 1873. Description de quelques crustaces nouveaux ou peu connus provenant du musee
de M. C. Godeffroy. J. Mus. Godeffroy 1 (4): 77-88, pis. 12, 13.

Ortmann, A., 1892. Die Decapoden-Krebse des Strassburger Museums mit besonderer Berucksichtigung
der von Herrn Dr. Doderlein bei Japan and bei den Liu-Kiu-Inseln gesammelten und zur Zeit in
Strassburger Museum auf bewahrten Formen. V. Hippidea, Dromiidea und Oxystomata. Zoot. Jb.
Syst. 6: 532-88, pi. 26.

Paul’son, O., 1875. Niesslidovania rakoobraznich Krassnago Morias zamietkami otnossitelno rakoobraz-
nich drouguich morei, Tchasst I. Podophthalmata i Edriophthalmata (Cumacea). 143 pp., 21 pis.
(S. V. Kul’zhenko: Kiev). [Studies on Crustacea of the Red Sea with notes regarding other seas.
Part 1. Podophthalmata and Edriophthalmata (Cumacea).] (English translation, 1961, National
Science Foundation: Washington).

Rathbun, Mary J., 1906. The Brachyura and Macrura of the Hawaiian Islands. Bull. U.S. Fish. Comm.
23 (3): 827-930, pis. 1-24.

1910. The Danish Expedition to Siam, 1890-1907. 5. The Brachyura. K. danske Vidensk. Selsk. Skr •
(7): 4: 303-67.

1911. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the leadership of Mr.
J. Stanley Gardiner. No. XI. Marine Brachyura. Trans. Linn. Soc. Lond. {Zoot.) (2) 14 (2): 191-261,
pis. 15-20.

1923. Report on the crabs obtained by the F.I.S. “Endeavour” on the coasts of Queensland, New South
Wales, Victoria, South Australia and Tasmania. Biol. Res. “ Endeavour ” 5 (3): 95-156, pis. 16-42.

1932. Preliminary descriptions of new species of Japanese crabs. Proc. Biol. Soc. Washington 45: 28-38.

48

MEMOIRS OF THE QUEENSLAND MUSEUM

Sakai, T., 1936. Studies on the crabs of Japan. I. Dromiacea. Sci. Rep. Tokyo Bunrika Daigaku (B) 3 (suppl.
1): 1-66, pis. 1-9.

1937. Studies on the crabs of Japan. II. Oxystomata. Sci. Rep. Tokyo Bunrika Daigaku (B) 3 (suppl. 2):
67-192, pis. 10-19.

1961. New species of Japanese crabs from the collection of His Majesty the Emperor of Japan. Crusta-
ceana 8 (1): 97-106.

1965. ‘The Crabs of Sagami Bay collected by His Majesty the Emperor of Japan’, pp. i-xvi, 1-206, 1 26
(English), 1-92, 27-32 (Japanese), pis. 1-100. (Maruzen: Tokyo).

Serene, R., 1955. Sur quelques especes rares de brachyures (Leucosidae) du l’indopacifique. 2e Partie.
Treubia 22 : 137-218, pis. 6-11.

Sluiter, C. Ph., 1881. Bijdrage tot de kennis der Crustaceen-Fauna van Java’s Noordkust. Natuurk.
Tijdschr. Ned.-Ind. 40: 159-64, 1 pi.

Stebbing, T. R. R., 1920. South African Crustacea (Part X of S.A. Crustacea, for the Marine Investigations
in South Africa). Ann. S. Afr. Mus. 17 : 231-72, pis. 18-27.

1921. South African Crustacea (Part XI of S.A. Crustacea, for the Marine Investigations of South
Africa). Ann. S. Afr. Mus. 18 : 453-68, pis. 13-20.

Stephenson, W. and Cook, S., 1970. New records of portunids from southern Queensland. Mem. Qd Mus.
15 (4): 331-4.

Takeda, M. and Mijake, S., 1970. Crabs from the East China Sea. IV. Gymnopleura, Dromiacea and Oxys-
tomata. J. Fac. Agric. Kyushu Univ. 16 (3): 193-235, pi. 1.

Tyndale-Biscoe, Marina and George, R. W., 1962. The Oxystomata and Gymnopleura (Crustacea,
Brachyura) of Western Australia with descriptions of two new species from Western Australia and
one from India. J. Proc. R. Soc. West. Aust. 45 (3): 65-96, pis. 1-3.

Ward, M., 1933. New genera and species of marine Decapoda Brachyura. From the coasts of New South
Wales and Queensland. Aust. Zool. 7 (5): 377-94, pis. 21-3.

Whitelegge, T., 1900. Scientific results of the trawling expedition of H.M.C.S. “Thetis”, off the coast of
New South Wales, in February and March, 1 898 . Crustacea. Part I. Mem. A ust. Mus. 4 (2) : 1 35-99,
pis. 32-5.

Yokoya, Y., 1933. On the distribution of decapod crustaceans inhabiting the continental shelf around
Japan, chiefly based upon the materials collected by S. S. Soyo-Maru, during the year 1923-1930.
J. Coll. Agr. Tokyo Imp. Univ. 12 : 1-226.

MEMOIRS OF THE QUEENSLAND MUSEUM

Plate 2

Mursia australiensis, holotype.

A, dorsal view; B, chela, outer face; C, chela, inner face; D, ventral view.

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

Plate 2

MEMOIRS OF THE QUEENSLAND MUSEUM

Plate 3

A, Ebalia brevimana, holotype. 1, dorsal view; 2, male abdomen; 3, male abdomen, lateral

view; 4, male pleopod in situ.

B, Ar cania (?) heptacantha.

C, Cryptocnemus hemispheroides , holotype.

CRABS TRAWLED OFF SOUTHERN QUEENSLAND

Plate 3

I L! 1 1 il 1 1 111 IIML

Mem. Qd Mus . (1971) 16 ( 1 ): 49-67

AMPHIBIAN AND REPTILE TYPE-SPECIMENS
IN THE QUEENSLAND MUSEUM

Jeanette Covacevich
Queensland Museum

ABSTRACT

Types and probable types of 76 nominate species or subspecies in the Queens-
land Museum collection are listed, including 45 of the 79 described by C. W. De
Vis, between 1884 and 1911.

The amphibian and reptile collections of the Queensland Museum commenced in 1875.
They were initially documented by a complex and barely workable system of donor, pur-
chase, exchange and collection registers which applied to all specimens. In May 1911, a
new and simplified system was introduced and is still in use. Under this all amphibians,
reptiles and mammals were accessioned with the prefix ‘J’ in a single register. Type-
specimens received since the current register began have been registered with reasonable
efficiency, but mistakes have undoubtedly been made in handling the backlog of older type
specimens, some of which were unregistered until the compilation of this list.

In determining whether a particular specimen is or is not the type of a particular
nominate species group taxon the following criteria are available.

(1) Designation by the author, with reference to a museum registration number.

(2) Notation in the register indicating that the specimen is a ‘Type’ (or ‘co-type’,

‘syntype’, etc.).

(3) Notation on the label of the jar containing the specimen indicating that the

specimen is a type.

(4) Coincidence of data recorded with specimen with that given by original author

where date of accession indicates that specimen under consideration would

have been available to the author at that time.

(5) Coincidence of measurements and items of description of specific species.

50

MEMOIRS OF THE QUEENSLAND MUSEUM

Difficulties arise when evidence based on these criteria is conflicting or inconsistent.
Thus many specimens found labelled ‘type’, ‘co-type’, or ‘syntype’ have been included as
only ‘probable types’ because they either differ markedly from the type-description or are
accompanied by collection data conflicting with that published. Also listed are several
specimens which were not labelled as types but can reasonably be presumed ‘types’ or, at
least, ‘probable types’ following comparison of specimens and register entries with the type
descriptions. One specimen, J76 Limnodynastes olivaceus (see p. 51) found labelled ‘De Vis
type’ has not been included because of conflicting locality data and the confirmed existence
of the holotype of this species in the British Museum.

De Vis was Curator in charge of the Queensland Museum collections from 1882 until
1905, and as the bulk of his new descriptions were published during this time, it seems
reasonable that most of his type-specimens were deposited in this Museum. In one paper
(1888, pp. 811-26) he described 18 new species, 17 from Queensland and 1 without locality.
Types of 1 1 are in the collection, so it must be considered probable that all were lodged here
and that 7 have been lost. Some, like the type of l . olivaceus , may have been sent to the
British Museum but often many of the specimens from a particular paper are in the
Queensland Museum while others cannot be located and must be presumed lost. De Vis
never published register numbers for his frogs and reptiles, rarely stated how many were
used for his descriptions or where they were lodged, and must be held largely responsible
for the apparent loss of so many of his type specimens.

The syntypes of Oedura tryoni De Vis, 1884, were collected from Stanthorpe and would
almost certainly have been deposited in the Queensland Museum. These specimens could
not be located in a recent search and Bustard (1966, pp. 6-7, pi. 1), has designated as neotype
an Australian Museum specimen, R21601.

Type-specimens (including probable types) are listed by family in alphabetical order
under the name by which they were first described. All specimens bear ‘J’ register numbers
and, with the exception of the holotype of Devisia mythodes Ogilby (mounted specimen),
are stored in 75% alcohol with glycerine in screw-top glass jars. Type categories listed by
the International Commission on Zoological Nomenclature (1964, p. 77) are followed.
A question mark is used to distinguish specimens that are only ‘probable types’ from those
which are definitely type specimens. Current names are given where synonymies have been
published. All available collection data from tags with the specimens and from the register
is listed. Any discrepancy between this and published data is noted. Where a locality name
has been changed the present name is given in parenthesis beside the original. An asterisk
beside a date is to distinguish it as a definite date of collection in contrast to other dates
which may be of collection, donation, receipt or registration. Most specimens are in good
condition but faded with age. A short statement on the condition of each is included.
Notes on the types of five species which have been presumed to be in the Queensland
Museum collection and one from the Amateur Fisherman’s Association of Queensland
(A.F.A.Q.) Museum are listed separately.

AMPHIBIAN AND REPTILE TYPE-SPECIMENS

51

TYPE-SPECIMENS ERRONEOUSLY ASCRIBED TO THE
QUEENSLAND MUSEUM COLLECTIONS

Limnodynastes olivaceus De Vis. A specimen in the Queensland Museum collection
(J76, Herbert R.) is labelled ‘Type’. Both Parker (1940, p. 55) and Moore (1961, p. 347)
state that the holotype of this species is in the British Museum collections. Miss A. G. C.
Grandison, Curator of Herpetology at the British Museum writes ‘ ... we do have a speci-
men presented by H. Ling Roth, taken at Pt. Mackay, Queensland, which is labelled as the
type of Limnodynastes olivaceus. It is a female and bears the registration number 85.9.2.25
. . Considering this information and the fact that the holotype was collected at Port
Mackay according to De Vis (1885, p. 66) and not, as was J76, from the Herbert River, it
seems probable that this specimen, although agreeing reasonably well with the specimen
described by De Vis, has been incorrectly labelled as the type.

Ranaster convexiusculus Macleay, Hylarana nebulosa Macleay. Types of these species
were originally part of the Macleay Museum type collection but have erroneously been
presumed to be in the Queensland Museum collection by Goldman, Hill, and Stanbury
(1969, pp. 429-30) who quote Moore (1961, p. 354, p. 345). Moore apparently based his
statement on Fry (191b, pp. 46-50). A careful check of the collection has shown that these
specimens are not in the collection now and they have probably never been here.

Mixophyes iteratus Straughan, M. balbus Straughan. According to the original descrip-
tions (Straughan 1968, pp. 54-7) paratypes of these species were deposited in the Queensland
Museum. Two paratypes (unregistered, 1 male, 1 female) of M. iteratus , collected ‘Tweed
River, Mount Warning, N.S.W.’ are listed as being held here. Thirteen paratypes (un-
registered, 10 males, 3 females) of M. balbus collected ‘Point Lookout, New England National
Park, N.S.W., between 4,250 and 4,750 feet altitude’ are listed as being held by ‘Australian
Museum and Queensland Museum’.

The frog collection received from Dr Straughan has been registered, and Queensland
Museum holdings of these two species are —

M. iteratus — 2 unsexed specimens, J18851, Lynch’s Creek, Kyogle, N.S.W.;
1 unregistered specimen without data.

M. balbus — 10 unsexed specimens, unregistered, found with tadpoles in jar
labelled ‘New England sp. nov. M. balbus ’ but with no other data.

Dr H. G. Cogger of the Australian Museum writes * . . . The specimens we have from
(Dr Straughan) are as follows:

52

MEMOIRS OF THE QUEENSLAND MUSEUM

Mixophyes balbus

Holotype, R25922, Point Lookout, New England National Park, N.S.W.

Paratypes, six specimens, R25923-R25928, data as for Holotype.

Mixophyes Herat us

Holotype, R25929, Tweed River, Mount Warning National Park, N.S.W.

Paratype, one specimen only, R25930, data as for Holotype.

Hence we have only one of the three paratypes of Mixophyes Herat us and six
of the thirteen paratypes of Mixophyes balbus .’

Hyla luteiventris Ogilby. An attempt has been made to locate the holotype of this
species in the A.F.A.Q. Museum where, according to the description (see Ogilby, 1906,
pp. 31-2), the specimen was deposited. Fry (1912, pp. 97, 99) examined this specimen and
synonym ised H. luteiventris with H. gracilenta Peters, but there is no trace of it since then.
It was not transferred to the Australian Museum (Cogger, pers. comm.) and as it is no longer
in the A.F.A.Q. Museum and the register which might have contained some mention of it
has been discarded, it can only be concluded that the holotype has been lost in one of the
many reorganizations of that Museum.

TYPE SPECIMENS IN THE QUEENSLAND MUSEUM COLLECTIONS
AMPHIBIA
MICROHYL1DAE

Cophixalus biroi darlingtoni Loveridge, 1948
Bull. Mus. comp. Zoo/. Harv. 101 (2): 423-4

Paratype: J9612; Toromanbanua, 7,500 ft, Bismarck Range, Madang Division, Australian
New Guinea, coll. Dr P. J. Darlington, October 1944*. (Good).

LEPTODACTYLIDAE

Crinia dariingtoni Loveridge, 1933
Occ. Pap. Boston Soc. nat. Hist. 8: 57-8

Paratype: J5444; National Park, Macpherson Range, 3-4,000 ft, SE.Q., don. Harvard
Museum, 4.iv. 1932. (Good).

Crinia tinnula Straughan and Main, 1 967
Proc. roy. Soc. Qd 78 (2): 19-21, pi. 1, fig. 1

Holotype: J 13546; male, Rose Creek, Beerburrum, SE.Q., coll. A. R. Main, I. R. Straug-
han, 31.viii.1965*. (Good).

Paratypes: J 13547-53; 7 males, Rose Creek, Beerburrum, SE.Q., coll. A. R. Main, I. R.
Straughan, 31.viii.1965*. (Good).

AMPHIBIAN AND REPTILE TYPE-SPECIMENS

53

Heleioporus sudelli Lamb, 1911
Ann. Qd Mus. 10: 26-7

(=. Heleioporus pictus Peters after Fry, 1912, p. 106. Helioporus eyrei (Gray)
after Loveridge, 1935, pp. 15-16. ^Limnodynastes sp. after Parker, 1940, p. 42.)
Holotype: J78; Warwick, SE.Q., coll. Miss J. Sudell. (Faded).

Two specimens originally registered with this number; one sent to Australian Museum
29 February 1912. Measurements of J78 agree with those in description.

Limnodynastes marmoratus Lamb, 1911
Ann. Qd Mus. 10: 28

(= Limnodynastes fletcheri Boulenger after Fry, 1912, p. 106)

Holotype: J12597; Goondiwindi, SE.Q., coll. J. Lamb, November 1910*. (Faded).
Paratypes: J 12598—60 1 ; 4 specimens, Goondiwindi, SE.Q., coll. J. Lamb, November
1910*. (Faded).

These five specimens re-registered from J77 which originally applied to six specimens.
One of series sent to Fry at Australian Museum 29 February 1912 according to register.
Lamb does not mention how many specimens used in description. J 12597 agrees with meas-
urements given and, according to description, ‘Type in Queensland Museum’.

Mixophyes fasciolatus schevilli Loveridge, 1933
Occ. Pap. Boston Soc. nat. Hist. 8: 55-6

(= Mixophyes schevilli Loveridge after Straughan, 1968, pp. 57-8, pi. 2, fig. 2)
Paratype: J5443; Millaa Millaa, NE.Q., don. Harvard Museum (Harvard collection
number 18151), 4.iv.l932. (Good).

Taudactylus diurnus Straughan and Lee, 1966
Proc. roy. Soc. Qd 77 (6) : 63-6, pi. 4.

Holotype: J13398; male, Green’s Falls, Maiala National Park, Mt. Glorious, SE.Q., coll.
I. R. Straughan, 12.V.1965*. (Good).

Paratypes: J13399-411; 3 males, 4 females, 6 unsexed (3 juv.). Green’s Falls, Maiala
National Park, Mt Glorious, SE.Q., coll. I. R. Straughan, 12. v. 1965*. (7 good, 5 with
bellies slit, 1 with belly slit and skin removed).

HYLIDAE

Hyla becki Loveridge, 1945
Proc. biol. Soc. Wash. 58: 55-6.

Paratype: J9613; Mt. Wilhelm, 7,500-10,000 ft, Bismarck Range, Madang Division,
New Guinea, coll. Captain P. J. Darlington, October 1944*. (Good).

Hyla irrorata De Vis, 1884
Proc. roy. Soc. Qd 1: 128-9

Syntypes: J 12870-80; 11 specimens, Gympie, SE.Q., coll. H. F. Wallman, 1884*. (All faded
and brittle, 3 with broken limbs).

Not knowing of the existence of these syntypes Copland (1961, p. 261) designated
J9255 as the ‘neotype’ of H. irrorata. A case to have Copland’s designation invalidated is

54

MEMOIRS OF THE QUEENSLAND MUSEUM

currently being prepared for submission to the International Commission of Zoological
Nomenclature.

Hyla vinosa Lamb, 1911
Ann. Qd Mus. 10: 27

(= Hyla lesueurii vinosa Lamb after Fry, 1915, pp. 84-6, pi. 2)

Holotype: J74; Ithaca Creek, Brisbane, SE.Q., 4.viii.l908*. (Faded, pattern visible)
Moore (1961 , p. 345) remarks ‘Status uncertain ; possibly valid; possibly Hyla lesueuri'.

REPTILIA

CHELYDRIDAE

Devisia mythodes Ogilby, 1907
Proc. roy. Soc. Qd 19: 11-16

(=Chelydra serpentina Linnaeus after Loveridge and Shreve, 1947, pp. 120-3)
Holotype: J20207; Fly River, New Guinea, don. Sir W. Macgregor. (Mounted; tip of tail
broken but present; some claws missing).

GEKKONIDAE

Diplodactylus hiilii Longman, 1915
Mem. Qd Mus. 3: 32-3

(= Diplodactylus conspicillatus Lucas and Frost after Kluge, 1963, pp. 83-4)
Holotype: J1994; Port Darwin, N.T., 7.viii.l914. (Tail separate, but present).

Kluge (1967, p. 1045) lists holotype as ‘ . . . Q.M. J2017 (= J 14/ 1994) . . .’. Register
entries show holotype registered twice (J1994, J2017) — no specimen now numbered J2017.

Diplodactylus taenicauda De Vis, 1887
Proc. Linn. Soc. N.S.W. 1: 169-70

Syntype: J232; Chinchilla, SE.Q. (Faded, pattern still visible; fragile, shape distorted).

Apparently only specimen of type series in existence; not specimen measured for
description.

Diplodactylus williamsi Kluge, 1963
Proc. Linn. Soc. N.S.W. 88 (2): 230-4, pi. 14

Paratypes: J270; Warialda, N.S.W., don. Australian Museum, 29.viii.1912. (Good).
J2324; Mungindi, SE.Q., don. Mrs Haager, 4.vi.l915. (Faded; tail missing). J6136; Retro
Station, Capella, M.E.Q., don. P. C. Allan. (Tail regenerated; belly slit). J6139; Retro
Station, Capella, M.E.Q., don. P. C. Allan. (Good, very small). J8430-1 ; 2 specimens,
Murilla Station, near Dalby, SE.Q., don. W. Dunmall, 9.xii.l952. (Good).

Nephrurus levis De Vis, 1887
Proc. Linn. Soc. N.S.W. 1: 168-9

Holotype: J246; Chinchilla, SE.Q. (Faded; tail separate, but present).

Measurements of this specimen agree with those given in description. De Vis gives no
locality for his material.

AMPHIBIAN AND REPTILE TYPE-SPECIMENS

55

Oedura cincta De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 811-12

(=Oedura marmorata Gray after Cogger, 1957, pp. 172-4)

?Syntype: J226; no data. (Faded, pattern visible).

According to register this specimen is a ‘type’. It is not specimen measured for descrip-
tion. Type locality is Charleville, SW.Q.

Oedura coggeri Bustard, 1966
Bull. Brit. Mus. ( nat . Hist.) Zool. 14 (1): 9-13, pi. 3
Paratypes: J 1293-5; 3 specimens, Stannary Hills, NE.Q., don. T. L. Bancroft. (Good).
J9290; Mt Garnet, NE.Q. (Good).

Oedura monilis De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 811
Holotype: J228; no data. (Faded; tail separate, but present).

Fry (1915, pp. 86-7, pi. 3) redescribes and figures O. monilis. Bustard (1967, p. 312)
states ‘ . . . I propose to recognise two subspecies of this taxonomic species. Due to its
condition . . . impossible to assign the holotype to either’.

Oedura reticulata Bustard, 1969
W. Aust. Nat. 11 (4): 82-6, figs. 1-2
Paratypes: J 13858-9; 2 specimens, Coolgardie district, W.A. (Good).

Perochirus mestoni De Vis, 1890
Proc. Linn. Soc. N.S.W. 4: 1035-6

( =Gehyra variegata (Dumeril and Bibron) after Kluge, 1963, p. 84)
Holotype: J236; Bellenden Ker, NE.Q. (Faded; area immediately behind head damaged).

PYGOPODIDAE

Delma plebeia De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 825-6

(— Delma fraseri Gray var. plebeia De Vis after Kinghorn, 1926, pp. 52-3)
Syntypes: J254; Brisbane, SE.Q. (Faded). J 12768-70; 3 specimens, Brisbane or Gympie,
SE.Q. (Faded: two without tails).

Register entry for J254: ‘Type = selected by De Vis from 12/247’ [= J247]. J12768-70
re-registered from J247 which originally referred to ten specimens.

Delma tincta De Vis, 1 888
Proc. Linn. Soc. N.S.W. 2: 824-5
Syntype: J241 ; Normanton, NW.Q. (Faded; tip of tail missing).

Measurements of this specimen fit those given in description. Syntypes from Springsure
not located.

SCINCIDAE

Ablepharus timidus De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 824

56

MEMOIRS OF THE QUEENSLAND MUSEUM

Syntypes: J235; Charleville, SW.Q. (Faded). J13601 ; Charleville, SW.Q. (Faded; bifurcate
tail broken but present).

Ctenotus hilli Storr, 1 969
J. roy. Soc. W. Aust. 52 (4): 103

Paratypes: J2613-8; 6 specimens, Darwin, N.T., don. G. F. Hill. (Faded, pattern visible;
bellies slit ; two with tails separate but present).

Ctenotus pantherinus calx Storr, 1969
J. roy. Soc. W. A ust. 52 (4) : 99

Paratype: J13000; Roper River, N.T., don. W. Hosmer. (Belly slit).

Ctenotus saxatalis Storr, 1969
J. roy. Soc. W. Aust. 52 (4): 101

Paratype: j 13001; Mt. Doreen Station, N.T., don. W. Hosmer. (Good).

Egernia lauta De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 813-4

(= Tiliqua luctuosa (Peters) after Mitchell, 1950, pp. 299-300, fig. 8)
Holotype: J249; no data. (Good).

Type locality is ‘Queensland’.

Egernia whitei modesta Storr, 1968
J. roy. Soc. W. Aust. 51 (2): 55
Holotype: J464; Chinchilla, SE.Q., 12.xii. 1912. (Good).

Paratypes: J3825; Thomby Station, St George, SE.Q., don. Captain Wilkins. (Some head
scales missing). J 13207-1 3; 7 specimens, Chinchilla, SE.Q. (5 good, 1 with most of right
forelimb missing, 1 with most of tail missing). J 13366; Greymare, Thane, SE.Q. (Some
dorsal scales and tail missing).

Heteropus blackmanni De Vis, 1884
Proc. roy. Soc. Qd 1 : 168

(= Leiolopisma vivax (De Vis) after Mitchell, 1953, pp. 85-6)

?Syntypes: J7773 ; no data. (Faded; tail missing). J19968-90; 23 specimens, no data.
(Faded; many with damaged tails).

J7773 found in jar labelled ‘ . . . probably one of type series. . .’. J 19968-90 found in
jar labelled ‘Cotypes Heteropus blackmaniV. Type locality is ‘Port Curtis’.

Heteropus lateralis De Vis, 1884
Proc. roy. Soc. Qd 1 : 168

(= Leiolopisma pectoralis (De Vis) after Mitchell, 1953, pp. 86-8)

Holotype: J234; Pine River, SE.Q., don. K. Stokes, E. Humber. (Faded; tip of tail
missing).

Prior to Mitchell’s examination of this specimen (31 July 1951), no reference to type
status in register. Type locality is ‘Moreton Bay district’ which includes Pine River.

Heteropus pectoralis De Vis, 1884
Proc. roy. Soc. Qd 1: 169

( = Leiolopisma pectoralis (De Vis) after Mitchell, 1953, pp. 86-8)

AMPHIBIAN AND REPTILE TYPE-SPECIMENS

57

Holotype: J1414; Port Curtis, M.E.Q., coll. F. A. Blackman. (Tail missing).

Type locality is ‘Warro, Port Curtis’, f

Heteropus rostralis De Vis, 1884
Proc. roy. Soc. Qd 1: 171

(= Leiolopisma fuscum fuscum (Dumeril and Bibron) after Mitchell, 1953
pp. 77-8)

Holotype: J230; Cardwell, NE.Q. (Good).

Heteropus vertebralis De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 821

(= Leiolopisma vertebralis (De Vis) after Mitchell, 1953, pp. 80-1)

Lectotype: J248; Chinchilla, SE.Q. (Good).

?Paralectotypes : J 137 1 9—22 ; 4 specimens, Chinchilla, SE.Q., coll. K. Broadbent. (3 good;
1 tail missing).

Register entry for J248 mentions ‘4 specimens’. Five were present in 1966 when 4 were
re-registered (J 1 3719—22). Mitchell (1953, p. 81) refers to \ . . remaining four specimens of
the type series . . If original register entry for J248 was correct only 3 of 4 specimens
listed are paralectotypes.

Hinulia ambigua De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 817-18
Holotype: J242; Charleville, SW.Q. (Faded, banding visible).

Hinulia tigrina De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 817

Holotype: J245; Geraldton [= Innisfail, NE.Q.], don. Dr Bancroft. (Faded).

Leiolopisma triacantha Mitchell, 1 953
Rec. S. Anst. Mus. 11: 88-9

Paratype: J7788; Darwin, N.T., don. G. F. Hill, November 1915*. (Tail separate, but
present).

Lygosoma bancrofti Longman, 1916
Mem. Qd Mus. 5: 49

Holotype: J2560; Gyranda, Dawson River, M.E.Q., don. Dr Bancroft. (Faded).
Type locality is ‘Upper Dawson River district’.

tWarro (Warroo until 1885) Station, Port Curtis, a property that has not existed for many years, is
the type locality for several De Vis reptiles, some of which have apparently been lost. The map (fig. 1 , p. 58)
shows the area that was Warro when De Vis’ new species were collected from this locality.

Frederick Archibald Blackman first leased Warro in 1879 and increased its size slightly in 1886. Since
1896 this area has been divided into several other holdings. Port Curtis was one of the Lands Department
administrative districts. It seems reasonable to assume that ‘Port Curtis’ specimens, received from Blackman
and described by De Vis, also came from Warro or near it.

58

MEMOIRS OF THE QUEENSLAND MUSEUM

Fig. 1: Map showing boundaries of Warro Station, Port Curtis, 1879-1896

AMPHIBIAN AND REPTILE TYPE- SPECIMENS

59

Lygosoma darlingtoni Loveridge, 1933
Occ. Pap . Boston Soc. nat. Hist . 8: 98-9

Holotype: J5806; female, Millaa Millaa, Atherton Tableland, NE.Q., don. Harvard
Australian Expedition, coll. Dr P. J. Darlington, April 1932*. (Good).

Lygosoma (Hinulia) tryoni Longman, 1918
Mem. Qd Mus . 6: 38-9

Holotype: J3023; Springbrok [== Springbrook], SE.Q., coll. H. Tryon. (Good).
Paratype: J3024; Macpherson Range, 3,000 ft, SE.Q. (Good).

Type locality ‘Macpherson Range, 3,000 ft, SE.Q.’ includes Springbrook.

Mocoa spectabilis De Vis, 1888
Proc. Linn. Soc. N.S.W. 2 : 819-20

( Lygosoma ( Leiolopisma ) spectabile (De Vis) after Longman, 1918, p. 38)
Syntypes: J244; Gympie, SE.Q. (Faded). J255; Gympie, SE.Q. (Faded). J 19742-3; 2 speci-
mens, Gympie, SE.Q. (Faded; J19742 tail missing).

J 1 9742—3 re-registered from J255. Measurements of J244 fit those in description.

Ophioscincus frontalis De Vis, 1888
Proc. Linn. Soc. N.S.W. 2: 823-4

Syntypes: J243; Geraldton [— Innisfail, NE.Q.], don. Dr Bancroft. (Slightly faded).
J 11499; Geraldton [= Innisfail, NE.Q.], don. Dr Bancroft. (Good).

?Syntypes: J 19737-41 ; 5 specimens, no data. (J19740 good; others poor).

J1 9737—41 found unmarked in jar containing J 1 1499 and labelled ‘Co-types’.

Rhodona allanae Longman, 1937
Mem. Qd Mus. 11: 167-8

Holotype: J6180; Retro Station, Capella, M.E.Q., don. Mrs P. C. Allan. (Good).
Paratypes: J6040; Retro, Capella, M.E.Q., don. MrsP. C. Allan, 15.vii.1936. (Tail separate
but present). J6238; Retro, Capella, M.E.Q., don. Mrs P. C. Allan, 28.vi.1937. (Faded).
?Paratype: J6308; Retro, Capella, M.E.Q., don. Mrs P. C. Allan. (Incomplete, head
missing).

Longman (1937, p. 168) states ‘Described from three specimens . . Apart from holo-
type which is designated in description and was marked clearly in reference collection, three
specimens (J6040, J6238 and J6308) found in jar with tag marked ‘ Rhodona allanae Lgmn.
Paratypes’. J6040 is marked as paratype in register. J6238 is probably other paratype
because if Longman had used incomplete specimen (J6308) for his description he would
probably have mentioned it.

Silubosaurus zellingi De Vis, 1884
Proc. roy. Soc. Qd 1: 53-4

(= Egernia stokesii (Gray) after Longman, 1912, p. 25)

Holotype: J253; Barcoo, C.Q., don. C. W. de Burgh Birch. (Flattened dorsoventrally).

Sphenomorphus schevilli Loveridge, 1933
Occ. Pap. Boston Soc. nat. Hist. 8: 96-7

Holotype: J5805; male. Army Downs, 35 miles north of Richmond, NW.Q., found under
concretion containing Plesiosaur, coll. W. E. Schevill, July 1932*.

60

MEMOIRS OF THE QUEENSLAND MUSEUM

Tiliqua longicauda De Vis, 1 888
Proc. Linn. Soc. N.S.W. 2: 816
(= Tiliqua gerrardii (Gray) after Zietz, 1920, p. 206)

?Syntypes: J250; Burpengary, SE.Q., don. Dr Bancroft, 27.viii.1912. (Faded, pattern
visible). J1 186-7; 2 specimens, Rockhampton, M.E.Q., don. Mr Jaggard, 29.V.1913.
(Faded, pattern visible; J 1 187 belly slit).

Type locality is ‘Rockhampton, coll. Mr Jaggard; Johnson River, coll. Mr W. H.
Miskin’. As all information (except date which is probably of registration) for J 1 1 86— 7
agrees with description, these specimens (not J250) are probably syntypes. Approximate
measurements of J 1 187 agree with those given in description.

Tropidophorus queenslandiae De Vis, 1 890
Proc. Linn. Soc. N.S.W. 4: 1034-5

Syntypes: J233; Bellenden Ker, NE.Q., don. A. Meston. (Good). J252; Bellenden Ker,
NE.Q. (Tip of tail missing).

?Syntypes: J19744-51; 8 specimens. (J19745 good, remainder poor).

Register entry for J252 is ‘8 specimens Tropidophorus queenslandiae , De Vis . . . Type
collection . . Jar containing specimen tagged J252 also contained 8 untagged specimens
which have been re-registered (J 19744-51). Assuming register is correct, 7 of these 8 speci-
mens are syntypes from Bellenden Ker and one specimen without data is indistinguishable
from type series. De Vis also refers to specimens from Herberton which have not been
located.

AGAMIDAE

Calyptoprymnus verecundus De Vis, 1905
Ann. Qd Mus. 6: 46-7, pi. 15
Holotype: J462; Solomon Islands. (Some scales missing).

Type locality ‘ . . . uncertain, but believed to have been brought from one of the Solo-
mon Islands

Macrops nuchalis De Vis, 1884
Proc. roy. Soc. Qd 1 : 97-8

(= Amphibolous reticulatus (Gray) after Boulenger, 1885, p. 386)

Syntypes: J 1405-9; 5 specimens, Delta Station, Bogantungan, M.E.Q., coll. C. W. de
Burgh Birch. (J1405 good; remainder either with belly slit or gutted and shrunken.
?Syntype: J1410; ?Queensland. (Good).

All specimens found in jar labelled ‘type and cotypes’. Measurements of J1409 agree
with those in description.

Tympanocryptis maculosa Mitchell, 1948
Rec. S. Aust. Mus. 9: 78-80

Paratypes: J7420-1 ; female and male, surface of Lake Eyre North, S.A., coll. Dr C. T.
Madigan, August-December 1929*. (Good).

AMPHIBIAN AND REPTILE TYPE-SPECIMENS

61

VARANIDAE

Varanus punctatus Gray var. orientalis Fry, 1913
Rec. Aust. Mus . 10 (1): 18-20, figs. 7-10

(= Varanus ( Odatria ) tristis orientalis Fry after Mertens, 1958, p, 245)
?Paratype: J640; Eidsvold, Upper Burnett River, SE.Q., don. T. L. Bancroft. (Good).
Labelled cotype in register and on tag with specimen. Holotype in Australian Museum.

T YPH LOP1DAE

Typhlops diversus Waite, 1894
Proc. Linn. Soc. N.S.W. 9: 10-11, pi. 1, figs. 4-6
Holotype: J2943 (formerly D4432); Mowen, Central Railway, Q. (see Waite, 1918, p. 32,
for corrected spelling — Morven), don. F. W. Allpuss, l.vi.1887. (Shape distorted).

BOIDAE

Aspidifes collaris Longman, 1913
Mem. Qd Mus. 2: 40

Holotype: J944; Avondale Station, via Cunnamulla, SW.Q., don. E. T. Bignell. (Damaged
three to four inches behind head; some scales missing).

Liasis amethistinus kinghorni Stull, 1933
Occ. Pap. Mus. Zool. Univ. Mich. 267 : 3-4

Paratype: J5501 ; LakeBarrine, Atherton Tableland, NE.Q., 2,300 ft, coll. P. J. Darlington.
(Head and skin only).

COLUBR1DAE

Neospades kentii De Vis, 1889
Proc. roy. Soc. Qd 6: 238-9

(= Myron richardsonii Gray after Mack and Gunn, 1953, p. 58)
Holotype: J681; Cambridge Gulf, NW. Australia, don. W. Saville-Kent. (Faded).

Tropidechis dunensis De Vis, 1911
Ann. Qd Mus. 10: 21-2

(= Dasypeltis scabra (Linnaeus) after Cogger, 1966, pp. 893-4)
Holotype: J191 ; no data in register. (Faded, no colour or pattern).

Described as an Elapid. Type locality is ‘Darro, Darling Downs’.

EL API DAE

Brachysoma sutherlandi De Vis, 1884
Proc. roy. Soc. Qd 1: 139-40

(= Demansia nuchalis (Gunther) after Mack and Gunn, 1953, p. 60)
Holotype: J190; no data in register. (Faded, banding visible).

Type locality is ‘Carl Creek, Norman River’.

62

MEMOIRS OF THE QUEENSLAND MUSEUM

Cacophis warro De Vis, 1884

Proc. roy . Soc. Qd 1: 139.

( =Rhynchoelaps warro (De Vis) after Mack and Gunn, 1953, pp. 66-7)
Holotype: J188; Warro Station, Port Curtis, ME.Q., coll. F. A. Blackman. (Faded, no
pattern).

Diemenia carinata Longman, 1915

Mem. Qd Mas. 3: 31, pi. 14

(= Pseudonaja nuchalis nuchalis Gunther after Worrell, 1961, p. 20)
Holotype: J1508; Cane Grass Station, via Charleville, SW.Q., don. J. Oswald Paynter,
6.iii.l914. (Good).

Denisonia angulata De Vis, 1905

Ann. Qd Mus. 6:51

(= Hoplocephalus bitorquatus (Jan) after Mack and Gunn, 1953, p. 65)
Holotype: J194; no data. (Good).

Type locality is ‘Queensland’.

Denisonia bancrofti De Vis, 1911

Ann. Qd Mus. 10: 23-4

(= Furina diadema (Schlegel) after Mack and Gunn, 1953, pp. 59-60)
Syntypes: J195; Stannary Hills, NE.Q. (Faded, head pattern visible; belly slit; fragile).
J 12881 ; Stannary Hills, NE.Q. (Brittle, broken in several places).

Original register entry for J195 refers to two specimens, one of which has been re-
registered (J 12881). Mack and Gunn (1953, pp. 59-60) list J 195 as the holotype and do not
mention J12881. Measurements and scale counts as compared in table 1 below suggest
that De Vis’s measurements are a composite of both specimens.

TABLE 1

Measurements and Scale Counts of Syntypes of Denisonia bancrofti

J195

J1288I

De Vis’
description

Mid-body Scales

15

15

15

Ventrals

195

185

185

Subcaudals

33

34

33

Anal

divided

divided

entire

Total length

180 mm

305 mm

190 mm

Tail

21 mm

40 mm

40 mm

Denisonia fenestrata De Vis, 1905
Ann. Qd Mus. 6: 50

(— Glyphodon tristis (Gunther) after Mack and Gunn, 1953, p. 59)

AMPHIBIAN AND REPTILE TYPE-SPECIMENS

63

Syntype: J200; Queensland. (Good).

Description refers to two specimens, one apparently lost.

Denisonia frontalis Ogilby var. propinqua De Vis, 1905
Ann. Qd Mus. 6: 51

(= Denisonia suta (Peters) after Mack and Gunn, 1953, p. 64)

Holotype: J198; Queensland. (Faded; neck and mid-body damaged).

Denisonia nigra De Vis, 1905
Ann. Qd Mus. 6 : 50

(= Denisonia coronoides (Gunther) after Mack and Gunn, 1953, p. 63)
Holotype: J196; Tasmania. (Good).

Denisonia revelata De Vis, 1911
Ann. Qd Mus. 10: 22-3

(= Hoplocephalus bitorquatus (Jan) after Mack and Gunn, 1953, p. 66)
Holotype: J2957; Stannary Hills, NE.Q., don. Dr Bancroft. (Faded, head pattern visible;
some scales loose).

Furina multifasciata Longman, 1915
Mem. Qd Mus. 3: 30

(= Vermicella annulata multifasciata (Longman) after Storr, 1967, pp. 80—1)
Holotype: J2019; Port Darwin, N.T. don. G. F. Hill. (Good).

Furina robusta De Vis, 1905
Ann. Qd Mus. 6 : 51-2

(= Vermicella bertholdi bertholdi (Jan) after Storr 1967, pp. 82-3)

Holotype: J205; no data. (Good).

Type locality is ‘Coolgardie, W.A.’.

Hoplocephalus ornatus De Vis, 1884
Proc. roy. Soc. Qd 1 : 1 00

(= Denisonia maculata (Steindachner) after Mack and Gunn, 1953, pp. 64-5)
Holotype: J199; Surat, SE.Q.; wooden tag in jar — ‘D. ornat . . . De Vis Warro (Type)’.
(Faded, some of pattern visible).

Register entry for J 1 99 refers to four specimens, three of which cannot be located.
Waite and Longman (1920, p. 179) refer to only one specimen as the type of H. ornatus.
Type locality is ‘near Surat’.

Hoplocephalus vestigiatus De Vis, 1884
Proc. roy. Soc. Qd 1: 138-9

(= Demanisa olivacea (Gray) after Mack and Gunn, 1953, p. 61)

Holotype: J206; no data. (Faded; neck damaged; shape distorted; most of tail missing).

Three untagged specimens found in jar labelled ‘J206 Type’ and register entry for this
number refers to three specimens. De Vis refers to a single specimen ‘in damaged condition’
and he did not give subcaudal scale count, apparently because tail was incomplete. Only
one of the three specimens was damaged and Mack and Gunn (1953, p. 61) regarded this
as the holotype. Two undamaged specimens re-registered (J 19849-50). Type locality is
‘uncertain’.

64

MEMOIRS OF THE QUEENSLAND MUSEUM

Micropechis crucifer De Vis, 1905
Ann. Qd Mus. 6: 52
Holotype: J197; no data. (Good).

Two specimens (one tagged J197, one untagged), accompanied by paper label ‘ Micro-
pechis crucifera De Vis Type’ found in jar marked ‘Type’. Register entry for J197 refers to
one specimen only. Description is of one specimen. Measurements and scale count given
by De Vis fit those of J197 reasonably well and do not fit the second specimen which has
been re-registered (J13685).

Type locality is ‘uncertain, believed to be the Solomon Islands’.

Pseudechis guttatta De Vis, 1905
Ann. Qd Mus. 6: 49-50

(= Pseudechis colletti guttatus De Vis after Worrell, 1961, p. 21)

Holotype: J 1 89 ; Cecil Plains, SE.Q. (Gutted).

Pseudechis mortonensis De Vis, 1911
Ann. Qd Mus. 10: 24

(= Pseudechis guttatus De Vis after Mack and Gunn, 1953, pp. 61-2)
?Holotype: J207; Brisbane suburbs, SE.Q. (Belly slit).

Mack and Gunn (1953, pp. 61-2) remark that this specimen was ‘labelled doubtfully
as the type . .

Pseudechis wilesmithii De Vis, 1911
Ann. Qd Mus. 10: 24-5

(= Oxyuranus scutellatus (Peters) after Mack and Gunn, 1953, pp. 62-3)
Holotype: J201 ; Walsh River, Cape York, don. Dr T. L. Bancroft. (Head and skin only;
head badly damaged, apparently when venom glands removed; body scales in good condi-
tion; tip of tail missing).

Pseudelaps bancrofd De Vis, 1911
Ann. Qd Mus. 10: 25

(= Demansia nuchalis (Gunther) after Mack and Gunn, 1953, p. 60)
Holotype: J 1 87 ; Stannary Hills, Atherton Tableland, NE.Q., don. Dr T. L. Bancroft.
(Faded, no pattern; body damaged approximately three inches behind head; some scales
slipping).

Rhynchelaps latizonatus De Vis, 1905
Ann. Qd Mus. 6: 49

(= Vermicella annulata (Gray) after Mack and Gunn, 1953, pp. 68-9)
Holotype: J 1 92 ; no data in register but tag in jar — ‘Herberton’. (Faded, banding visible).

Type locality is ‘Queensland’.

HYDROP1IDAE

Distira nasalis De Vis, 1905
Ann. Qd Mus. 6 : 48

(= Hydrophis major (Shaw) after Mack and Gunn, 1953, pp. 58-9)

Holotype: J203; Queensland coast. (Faded, dorsal pattern visible).

AMPHIBIAN AND REPTILE TYPE-SPECIMENS

65

Platurus frontalis De Vis, 1905
Ann. Qd Mus. 6: 48

(= Laticauda colubrina (Schneider) after Longman, 1918, p. 41)
Holotype: J202; New Guinea. (Good).

ACKNOWLEDGMENTS

Miss A. G. C. Grandison, of the British Museum, and Dr H. G. Cogger, of the Aus-
tralian Museum have provided me with information on specimens in their care.

Information on several of the type localities was made available by Mr W. H. Muir-
head, of the Queensland Place Names Board, and Miss S. Geddes of Queensland State
Archives. The map, prepared for publication by Miss M. McKenzie, could not have been
drawn without the co-operation of officers of the Land Administration Commission and
Survey Office, Brisbane. Mr B. Campbell of the Queensland Museum provided helpful
advice throughout the compilation of this list.

LITERATURE CITED

Boulenger, G. A., 1885. ‘Catalogue of the lizards in the British Museum (Natural History).’ Vol. 1. (British
Museum: London).

Bustard, H. R., 1966. The Oedura tryoni complex: east Australian rock-dwelling geckoes (Reptilia : Gek-
konidae) Bull. Brit. Mus. ( nat . Hist.)Zool. 14 (1): 1-14, pis. 1-3.

1967. The status of the gekkonid lizard name Oedura ocellata Boulenger, 1885. Herpetologica 23 (4):
312-3.

1969. Oedura reiiculata, a new velvet gecko from south-west Western Australia. W. Aust. Nat. 11 (4):
82-6, figs. 1-2.

Cogger, H. G., 1957. Investigations in the gekkonid genus Oedura Gray. Proc. Linn. Soc. N.S. W. 82 : 167-79,
pis. 7-8.

1966. The status of the “Elapid” snake Tropidechis dunensis De Vis. Copeia 4 : 893-4.

Copland, S. J., 1962. Rediscovery of a little known Victorian frog. Proc. Linn. Soc. N.S. W. 86: 258-61 .

De Vis, C. W., 1884a. On new Australian lizards. Proc. roy. Soc. Qd 1 : 53-6.

1884b. On new species of Australian lizards. Proc. roy. Soc. Qd 1 : 97-100.

1884c. On a new species of Hoplocephalus. Proc. roy. Soc. Qd 1 : 100, pi. 15.

1884d. On new species of Hyla. Proc. roy. Soc. Qd 1 : 128-30.

1884e. Descriptions of new snakes with a synopsis of the genus Hoplocephalus. Proc. roy. Soc. Qd 1 :
138-40.

1884f. A conspect of the Genus Heteropus. Proc. roy. Soc. Qd 1 : 166-73.

1885. On some new batrachians from Queensland. Proc. Linn. Soc. N.S.W. 9 : 65-7.

1887. On certain geckoes in the Queensland Museum. Proc. Linn. Soc. N.S. IV. 1 : 168-70.

1888. A contribution to the herpetology of Queensland. Proc. Linn. Soc. N.S. W. 2: 81 1-26.

66

MEMOIRS OF THE QUEENSLAND MUSEUM

1889. Description of two new vertebrates in Mr Saville-Kent’s collection. Proc. roy. Soc. Qd 6: 237-9.
J890. Descriptions of two lizards of genera new to Australian herpetology. Proc. Linn. Soc. N.S.W. 4:
1034—6.

1905. Reptilia. Ann. Qd Mus. 6: 46-52, pi. 15.

1911. Description of snakes apparently new. Ann. Qd Mus. 10: 21-5.

Fry, D. B., 1912. Description of Austrochaperina a new genus of Engystomatidae from north Australia.
Rec. Aust. Mus. 9: 87-106, pis. 8-9.

1913a. On a Varanus and a frog from Burnett River, Queensland, and a revision of the variations in
Limnodynastes dorsalis. Gray. Rec. Aust. Mus. 10 (1): 17-34, pis. 1-3, figs. 7-13.

1913b. A re-examination of Macleay’sNew Guinea and Queensland frog types. Mem. Qd Mus. 2:46-50.
1915. Herpetological notes. Proc. roy. Soc. Qd 27: 60-88, pis. 1-4.

Goldman, J., Hill, L. and Stanbury, P. J., 1969. Type specimens in the Macleay Museum, University of
Sydney (Amphibians and Reptiles). Proc. Linn. Soc. N.S.W. 93 (3): 427-38.

International Commission on Zoological Nomenclature, 1964. ‘International Code of Zoological
Nomenclature adopted by the XVI International Congress of Zoology.’ (International Trust for
Zoological Nomenclature: London).

Kinghorn, J. R., 1926. A brief review of the family Pygopodidae. Rec. Aust. Mus. 15 : 40-64.

Kluge, A. G., 1963a. The systematic status of certain Australian and New Guinean gekkonid lizards.
Mem. Qd Mus. 14 (3): 77-86.

1963b. A new species of gekkonid lizard, genus Diplodactylus Gray, from eastern Australia. Proc. Linn.
Soc. N.S.W. 88 (2): 230-4, pi. 14.

1967. Systematics, phylogeny, and zoogeography of the lizard genus Diplodactylus Gray (Gekkonidae).
Aust.J. Zool. 15: 1007-108, pis. 1-19.

Lamb, J., 1911. Description of three new batrachians from southern Queensland. Ann. Qd Mus. 10: 26-8,

Longman, H. A., 1912. Herpetological notes. Mem. Qd Mus. 1 : 23-5.

1913. Herpetological notes, Mem . Qd Mus. 2: 39-42.

1915. Reptiles from Queensland and the Northern Territory. Mem. Qd Mus. 3: 30-4, pis. 14-15.

1916. Snakes and lizards from Queensland and the Northern Territory. Mem. Qd Mus. 5: 46-51, pi. 6.
1918. Notes on some Queensland and Papuan reptiles. Mem. Qd Mus. 6: 37^44, pis. 11-15.

1937. Herpetological notes. Mem. Qd Mus. 11: 165-8, pi. 14.

Loveridge, A., 1933a. Four new crinine frogs from Australia. Occ. Pap. Boston Soc. nat. Hist. 8: 55-60.
1933b. New scincid lizards of the genera Sphenomorphus, Rhodona, and Lygosoma from Australia.
Occ. Pap. Boston Soc. nat. Hist. 8: 95-100.

1935. Australian Amphibia in the Museum of Comparative Zoology, Cambridge, Massachusetts.
Bull. Mus. Comp. Zool. Harv. 78 (1): 3-60, pi. 1.

1945. New tree frogs of the genera Hyla and Nyctimystes from New Guinea. Proc. biol. Soc. Wash. 58:
53-8.

1948. New Guinean reptiles and amphibians in the Museum of Comparative Zoology and United States
National Museum. Bull. Mus. comp. Zool. Harv. 101 (2): 306-430.

Loveridge, A. and Shreve, B., 1947. The “New Guinea” Snapping Turtle ( Chelydra serpentina). Copeial :
120-3.

Mack, G. and Gunn, S. B., 1953. De Vis’ types of Australian snakes. Mem. Qd Mus. 13 : 58-70, figs. 1-3.

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67

Mertens, R., 1958. Bemerkungen fiber die Warane Australiens. Senckenbergiana biol. 39 (5-6): 229-64,
pis. 24-31.

Mitchell, F. J., 1948. A revision of the lacertilian genus Tympanocryptis . Rec. S. Aust. Mus. 9: 57-86, pis.
4-6, figs. 1-10.

1950. The scindid genera Egernia and Tiliqua (Lacretilia). Rec. S. Aust. Mus. 9; 275-308, pi. 23.

1953. A brief revision of the four-fingered members of the genus Leiolopisma (Lacertilia). Rec. S. Aust.
Mus. 11 : 75-90, figs. 1-4.

Moore, J. A., 1961. The frogs of eastern New South Wales. Bull. Amer. Mus. nat. Hist. 121: 153-387, pis.
27-46.

Ogilby, J. D., 1906. A new tree frog from Brisbane. Proc. roy. Soc. Qd 20: 31-2.

1907. Catalogue of the emydosaurian and testudinian reptiles of New Guinea. Proc. roy. Soc. Qd 19:
1-31.

Parker, H. W., 1940. The Australian frogs of the family Leptodactylidae. Novit. zool. 42 (1): 1-106, pi. 1.

Storr, G. M., 1967. The genus Vermicella (Serpentes, Elapidae) in Western Australia and the Northern
Territory. J. roy. Soc. W. Aust. 50 (3): 80-92.

1968. Revision of the Egernia whitei species-group (Lacertilia, Scincidae). J. roy. Soc. W. Aust. 51 (2):
51-62.

1969. The genus Ctenotus (Lacertilia, Scincidae) in the Northern Territory. J. roy. Soc. W. Aust. 52 (4) :
97-108.

Straughan, I. R., 1968. A taxonomic review of the genus Mixophyes (Anura, Leptodactylidae). Proc. Linn.
Soc. N.S.W. 93 (l) no. 416: 52-9, pis. 1-2.

Straughan, I. R. and Lee, A. K., 1966. A new genus and species of leptodactylid frog from Queensland.
Proc. roy. Soc. Qd 77 (6) : 63-6, pi. 4.

Straughan, I. R. and Main, A. R., 1967. Speciation and polymorphism in the genus Crinia Tschudi
(Anura, Leptodactylidae) in Queensland. Proc. roy. Soc. Qd 78 (2): 11-28, pis. 1-6.

Stull, O. G., 1933. Two new subspecies of the family Boidae. Occ. Pap. Mus. Zool. Univ. Mich. 267: 1-4.

Waite, E. R., 1894. Notes on Australian Typhlopidae. Proc. Linn. Soc. N.S.W. 9: 9-14, pi. 1, figs. 1-9.

1918. Review of the Australian Blind Snakes (family Typhlopidae). Rec. S. Aust. Mus. 1 : 1-34, pi. 1,
charts 1-9.

Waite, E. R. and Longman, H. A., 1920. Descriptions of little-known Australian Snakes. Rec. S. Aust.
Mus. 1: 174-80, pi. 27.

Worrell, E., 1961. Herpetological name changes. W. Aust. Nat. 8: 18-27.

Zietz, F. R., 1920. Catalogue of Australian lizards. Rec. S. Aust. Mus. 1: 181-228.

Mem . QdMus. (1971) 16 ( 1 ): 69-102

A REVIEW OF THE MEGASCOLECOID EARTHWORM GENERA
(OLIGOCHAETA) OF AUSTRALIA.

PART III— THE SUBFAMILY MEGASCOLECINAE

B. G. M. Jamieson

Zoology Department, University of Queensland

ABSTRACT

This paper forms the final part of a review in which the earthworm family
Megascolecidae has been redefined and its constituent genera have been arranged
in new groupings. The subfamily Ocnerodrilinae and its two tribes the Ocnero-
drilini and Malabarini represent no departure from the previous classification
of Gates beyond their reduction from familial and subfamilial ranks, respectively,
but the constitution of the remaining subfamilies here recognized, the Acantho-
drilinae and Megascolecinae, differs significantly from that proposed by other
workers. The subfamily Acanthodrilinae, which was treated in part II, was
amended to include holonephric and meronephric forms in which the prostates
varied in number but were never restricted to a pair near or combined with male
pores on XVIII. The subfamily Megascolecinae, with which this paper deals,
is reserved for worms with the pores of the single pair of prostates close to or
combined with the male pores on XVIII and for those with the acanthodrilin
arrangement (or an approximation to this) which have exonephric stomate
nephridia median to micromeronephridia in posterior segments. The latter
condition of the excretory system is diagnostic of the Tribe Dichogastrini. The
Dichogastrini corresponds with the Octochaetinae (-idae) of some previous
workers after transfer of Octochaetus and some other genera to the Acantho-
drilinae as the Tribe Octochaetini. Tribes of the Megascolecinae recognized in
addition to the Dichogastrini are the Perionychini, which possess holonephridia,
and the Megascoiecini which have various non-dichogastrin arrangements of
meronephridia. The Megascolecinae, their tribes, and all constituent Australian
genera are defined and check lists of the species of the latter genera are provided.

In part I of this review (Jamieson, 1971a) a new classification of the megascolecoid
earthworms of the world was advanced in synoptic form. In the present work fullered fini-
tions of suprageneric taxa recognized within the subfamily Megascolecinae will be given,

70

MEMOIRS OF THE QUEENSLAND MUSEUM

and additional evidence for recognizing the proposed groupings will be advanced. The
Australian genera will be defined in preparation for their revision and checklists of Aus-
tralian species will be given. This paper completes a review of the 18 genera and listing of
the 247 species comprising the known megascolecoid fauna indigenous to Australia.

SYSTEMATICS

Subfamily MEGASCOLECINAE

Megascolecidae in which the prostates have a single, central lumen or a central lumen
receiving lateral canals, and are externally tubular or (racemose prostates) the lumen is
branched from its junction with the duct. Purely holonephric or meronephric, or with
meronephridia in a varying number of segments anterior to holonephridia. If stomate
nephridia are present median to micromeronephridia, no holonephridia are present. Male
pores on XVIII or its homeotic equivalent; prostates a single pair with the pores on the same
segment as the male pores, with which they are almost always united; or male terminalia
acanthodrilin, with prostate pores 2 pairs, on XVII and XIX and male pores intermediate
on XVIII; such acanthodrilin worms differing from the Acanthodrilinae in having stomate
nephridia median to micromeronephridia in posterior segments and no holonephridia.
Homeotic forms ( Tonoscolex ) with combined male and prostatic pores on XVII, differing
from Acanthodrilinae with similarly located male terminalia in possessing racemose
prostates. Calciferous glands, if present, not arranged as in the Ocnerodrilinae ; intestine
commencing in or behind XIV ; last hearts rarely in XI.

Tribes: Dichogastrini ; Perionychini ; and Megascolecini.

Distribution: Palaearctic: China, Korea; Japan. Nearctic: N. America; Queen
Charlotte I. Neotropical: Mexico; Central and Tropical South America; West Indies;
Cuba. Ethiopian: Tropical Africa. Oriental: India, Pakistan and northwards beyond
Nepal; Ceylon; Andaman I.; Assam; Annam; Burma; Java; Sumatra; Philippines;
Molluccas. Australian: Australia and Tasmania; Norfolk Island; New Guinea. New
Caledonia. New' Zealand and neighbouring islands including Chatham Island. Sub-
antarctic islands : Auckland I. ; Snares I. ; Stewart I. Fiji. Endemicity over much of the
Pacific, from the Mariannas, Bismarck Archipelago and the Solomons eastwards remains
to be established. Some species peregrine, sometimes widely.

Type genus: Megascolex Templeton, 1844.

Tribe DICHOGASTRINI

Megascolecinae which are purely meronephric; with micromeronephridia anteriorly,
often with bucco-pharyngeal nephridia; posteriorly with astomate micromeronephridia

MEGASCOLECOID EARTHWORMS

71

together with, on each side in each segment, a median nephridium with one or more
preseptal funnels, which frequently is enlarged as a megameronephridium. Prostates tubular
or less commonly racemose or intermediate, one pair, their pores on XVII, XVIII or XIX,
united with or rarely near the male pores; or two pairs, their pores on XVII and XIX,
with the male pores intermediate on XVIII or sometimes nearer to or united with the
anterior prostatic pores ; exceptionally ( Hoplochaetella ) with intestinal enteronephry of the
megameronephridia and with two pairs of prostatic pores combined with two pairs of male
pores, on XVII and XIX or 17/18 and 18/19.

Distribution: Palaearctic: China (?). Nearctic: N. America. Neotropical:
Mexico; Central and Tropical S. Africa; West Indies; Cuba. Ethiopian: Tropical Africa.
Oriental: India, Pakistan and northwards beyond Nepal; Burma; Ceylon. Australia.
New Zealand. Fiji. Some species peregrine, sometimes widely, in warmer regions.

Type-genus: Dichogaster Beddard, 1889b.

Genera: (see Jamieson, 1971a).

Australian Genera: Digaster Perrier, 1872 (part, including the type-species?);
Didymogaster Fletcher, 1887a; Megascolides (part) McCoy, 1878; Notoscolex (part, includ-
ing the type-species?) Fletcher, 1887a, and Spenceriella ( S . gigantea only?) Michaelsen,
1907a.

Remarks : The author, in preparing a list of those genera of the Octochaetidae s. Gates
which have both micromeronephridia and, posteriorly, megameronephridia or at least
stomate meronephridia, found that precisely those genera ( Deinodrilus , Leucodrilus, Hoplo-
chaetina and Octochaetus ) which Lee suspected of having their closest affinities with genera
placed by Gates (1959) in the Acanthodrilidae stood apart in lacking these features, having
neither megameronephridia nor median stomate nephridia though purely meronephric.
The other genera of Gates’s Octochaetidae were purely meronephric and in the hindbody
had median stomate nephridia, often enlarged as ‘meganephridia’ (megameronephridia) in
addition to closed ‘micronephridia’, with the exception of the two Indian genera Octochae-
tona and Octochaetoides which were significantly products of division of Octochaetus. The
genera with median stomate nephridia comprise the Dichogastrini of the present work.

The homogeneity of the Dichogastrini is supported by Sims’s computer analysis and
by virtue of their being the core of Gates’s Octochaetidae. Although the author recognized
the existence of the tribe independently, its validity and delineation on the basis of the type
of excretory system is supported by earlier work of Gates (1940b) who dispensed with the
‘obsolescent’ name Octochetinae and substituted the name ‘ Hoplochaetella group’ which
comprised Dichogaster (from which Millsonia and Benhamia have since been resurrected),

72

MEMOIRS OF THE QUEENSLAND MUSEUM

Hoplochaetella, Ramiella , Eutyphoeus, Scolioscolides, Barogaster, Eudichogctster, Lenno-
gaster, Pellogaster, Rillogaster, Priodochaeta and provisionally also Priodoscolex.

References attesting the coexistence of median stomate nephridia with micromerone-
phridia, with the dubious exceptions of Priodoscolex and Trigaster, for all genera of the
Dichogastrini are set out below. In the examples bearing an asterisk, the median stomate
nephridium is known to be enlarged as a megameronephridium. Bibliographic references
in parentheses denote the source of information and only coincidentally refer to authors
of taxa.

Bahlia : Nephridia are undescribed but the genus is said to be similar to Eutyphoeus
q.v. (Gates, 1945). Barogaster : type-species, B. barodensis* (Gates, 1939b, p. 158). Ben-
hamia : type-species, B. rosea * (Omodeo, 1958, p. 39); B. liberiensis* (Omodeo, 1958, p. 45).
Calebiella : type-species, C. parva* (Gates, 1945, p. 244). Celeriella : in the type-species,
C. duodecimals, some meronephridia (four in a segment?) are enlarged almost to the
appearance of holonephridia according to Michaelsen (1907a) but in C. quadripapillata a
single median nephridium on each side, in the hindbody, has a preseptal funnel (Gates,
1958, p. 614). Dichogaster : type-species, D. damonis * (Beddard, 1889a, p. 258); D. bolaui*
(Gates, 1958, p. 619); D. titillata* (personal observations of a type-specimen). Didymogaster
sylvaticus* (Horan, personal communication). Digaster* (part(?), seep. 75); Eudichogaster*
(Gates, 1939b, p. 160). Eutrigaster : in the type-account (E. oraedivitis Cognetti, 1904)
nephridia are merely described as diffuse but Omodeo (1955) considers it to belong to the
viridis- group of Dichogaster. Eutyphoeus : E. foveatus *, E. nicholsoni* and E. waltoni*
(Bahl, 1942, p. 440). Hoplochaetella : H. khandalensis* (Bahl, 1947, p. 122); this genus is
exceptional in that the megameronephridia, instead of being exonephric, open into a pair of
longitudinal excretory canals which run along the inner surface of the body wall, one on
each side of the middorsal line, from segment XX to the anal end, each canal opening
separately at the place where the outer body wall passes into the wall of the rectum, a first
step in the direction of the enteronephric condition (see also Gates, 1940b). Lennogaster :
type-species, L. yeicus, median stomate nephridium thickened but not enlarged (Gates,
1939b, p. 183). Megascolides : type-species, M. australis * (Spencer, 1888, p. 22; Bahl, 1947,
p. 121); M. raglani* and M. urewerae* have ‘meganephridia’ in addition to ‘micronephridia’
in the posterior segments (Lee, 1959) as do some Indian species (Stephenson, 1923);
megameronephridia or median stomate nephridia are not described by Lee for other New
Zealand species of this genus but no indication of examination of the posterior end is given.
Millsonia*: entire genus (always?) (Omodeo, 1955); details for M. inermis * (Omodeo,
1955, p. 222); M. anomala* shows an interesting approach to the condition in Hoplochae-
tella khandalensis : the large megameronephridia of each side (present in XXI posteriorly)
are connected to a longitudinal duct which runs on each side of the nerve cord. This duct
opens to the exterior regularly in each intersegment, medianly by a minute pore whereas
in Hoplochaetella there are no pores in the course of the duct (Omodeo, 1955, p. 220).
Neogaster\ type-species, N. americanus, with two pairs of nephridia in each segment (mid-
body only investigated), the ventral nephridia alone having funnels (Pickford, 1937, p. 608).

MEGASCOLECOID EARTHWORMS

73

Notoscolex (part, including the type-species ?) : Fletcher’s account of the type-species
indicates merely that it has parietal (exonephric ?) meronephridia. Both Notoscolex ulla-
dullae* and N. attenuatus * (Australia) have tufts which in the former are probably and in
the latter are certainly pharyngeal. They also have scattered parietal micromeronephridia
and, coexisting with those in each of the hindermost segments, a (stomate?) megamero-
nephridium on each side (Boardman, 1931). The Australian species Notoscolex (= Mega-
scolides ) albertsi* Cognetti, 1910, has enlarged (stomate?) ventral nephridia from segment
XXX posteriorly. Omodeona: in the type-species, O. proboscoides *, there is in each of the
posterior segments, on each side, a large megameronephridium, shortly lateral of the nerve
cord and below the gut. It has a large preseptal funnel and a wide duct which enters the
parietes shortly in front of seta c. Lateral to the megameronephridium there are, on each
side of the body, three or four very small micromeronephridia (personal observations of a
syntype).

Pellogaster : type-species, P. bengalensis* , (Gates, 1939b, pp. 200, 203). Priodochaeta* :
‘meganephridia’ with funnels extending to the posterior end ; posterior extent of ‘micro-
nephridia’ uncertain (Gates, 1940a). Priodoscolex : Gates (1940b) was uncertain whether
this genus should be included in his ‘ Hoplochaetella group’. It appears from his account of
the genus that each of the posterior meronephridia, rather than merely the medianmost
one, is stomate. The description is as follows: Excretory organs: two pairs of closed, exo-
nephric ( ?) micronephridia (or clusters of micronephridia) on the parietes of III (?), one pair
of large clusters on the parietes of IV (?), two pairs of large clusters close to the ventral
parietes in IV and V, paired clusters of micronephridia on the anterior faces of the septa
in VI-IX (and X-XI ?) and XII-XIII ; from XIV posteriorly a transversely placed, preseptal
band of closed, exonephric micronephridia on each side, the band of two rows; in the
posteriormost segments a single transversely placed row of 10-12 small, exonephric micro-
nephridia on each side with a corresponding row of preseptal funnels.

Ramiella : (Gates, 1958, p. 610); R. parva and R. pachpaharensis (Bahl, 1942, p. 441 ;
contrary to Stephenson who found no septal connections or funnels). The small number
of meronephridia, the absence of pharyngeal and of other enteronephry (Bahl, 1942, p. 441)
and the absence or rudimentary condition of calciferous glands in Ramiella appear to be
primitive features and suggest that with the partly holonephric Howascolex (Indian section),
Ramiella is a little-changed representative of the ancestral stock of the Indian and other
genera of the Dichogastrinae. Neogaster americanus , like the type-species, Ramiella bis-
hambari, has only 2 pairs of meronephridia per segment (including the median stomate
nephridia) but has well developed calciferous glands.

Ramiellona: like Ramiella, this genus lacks enteronephric nephridia. There are several
ranks of meronephridia on each side, the median nephridium on each side in posterior
segments having a preseptal funnel, vide R. guatemala, R. balantina, R. strigosa and R.
mexicana , in all of which the median stomate nephridium is not enlarged (Gates, 1962c,
pp. 198, 203, 207, 211) and R. irpex , in which the median stomate nephridium is described

74

MEMOIRS OF THE QUEENSLAND MUSEUM

as a meganephridium. (Pickford, 1937, p. 607). Rillogaster* : (Gates, 1939b, p. 207); Bahl
(1942, p. 446) drew attention to the similarity of the excretory systems of Scolioscolides,
Barogaster , Eudichogaster, Lennogaster, Pellogaster and Rillogaster.

Scolioscolides : type-species, S. bergtheili* (Gates, 1937, p. 309; Bahl, 1942, p. 446).
Spenceriella (part, excluding the type): S. gigantea* (Lee, 1959, p. 344).

Trigaster : the condition in this genus is not entirely certain. Gates (1962b) states that
the nephridia of T. cavernicola form at least six longitudinal ranks posteriorly; those of the
median rank are larger than the others but funnels are not mentioned and he queries that
they may be composite. Wegeneriona : Pickford (1937, p. 608) who placed W. michaelseni
in Howascolex, states that this species has two or three pairs of nephridia in each segment
of the middle region of the body which she examined, and that the ventralmost nephridium
appeared to have a septal attachment though she was unable to state definitely that a septal
funnel was present.

Median stomate nephridia also occur in some genera of Gates’s Megascolecidae, and
the condition presumably indicates phylogenetic relationship with the Dichogastrini, but
in the ‘megascolecid’ genera the stomate nephridia discharge into the intestine, with the
notable exception of some species of Notoscolex and of Didymogaster sylvatica. In these
species they remain exonephric and in view of the reduction in significance of the racemose
condition of prostates, demonstrated in the present work there seem no barriers to placing
them in the Dichogastrini. This step resolves yet another anomaly to which Sims (1966)
drew attention, clustering of Notoscolex with the octochaetine genera.

The position of Hoplochaetella deserves special mention. It alone of the Dichogastrini
has intestinal enteronephry of the posterior stomate megameronephridia. This might appear
to necessitate placing it in the Megascolecini but the close affinity to octochaetine genera
and isolation from ‘megascolecid’ genera demonstrated by Sims (1966) suggests that there
is more justification in grouping it in the Dichogastrini. Its highest coefficient of similarity
(16.3%) was with the dichogastrin genus Lennogaster in Sim’s analysis, the low figure
indicating its relatively isolated position. Its two pairs of openings of the male ducts is
unique in the megascolecoid earthworms and exceedingly rare in megadrile oligochaetes.

Genus Digaster Perrier, 1872, emend. Jamieson, 1963

Terrestrial worms ranging from slender forms little more than an inch long to inch-wide
specimens over 5 feet in length. Segments less than 100 to over 300. Pigmentation of the
body wall present or absent. Prostomium very variable. Setae 4 pairs per segment, rather
widely to closely paired. Clitellum annular or saddle-shaped, embracing part of the region
between intersegments 11/12 and 19/20. A pair of pores, each of the united prostatic duct,

M EG ASCOLECOID EARTHWORMS

75

and sperm duct of its side, present in XVIII. Accessory puberty papillae frequently present
in the vicinity of the male pore segment or in the forebody. Female pores 1 pair, or single
and median, presetal in XIV. Spermathecal pores 2 pairs, in 7/8 and 8/9 with sometimes
(intraspecific variation in D. longmani ) a third pair in 6/7. Position of the first dorsal pore
variable, usually in 5/6 or 11/12.

Two gizzards, in V and VI, or in VI and VII (perhaps in V and VII in the type-species) ;
or 3 gizzards, in V, VI and VII; oesophagus usually highly vascularized and showing partial
or complete development of calciferous glands ; intestine commencing in XVII and XVIII,
exceptionally in XVI. Dorsal blood vessel single; last hearts in XII or less commonly in
XIII. Meronephric; some species with tufted enteronephric and/or exonephric nephridia
formed by apposition of ducts of micromeronephridia; some species with megamero-
nephridia in addition to micromeronephridia posteriorly. Holandric or metandric ; (always ?)
lacking testis-sacs. Prostates racemose with branched ducts within the gland. Sperm ducts
(always?) uniting entally with the prostate ducts. Spermathecae 2 pairs, in VIII and IX, or
3 pairs, in VII, VIII and IX, each (always ?) with 1 or more diverticula which are never
long tubes.

Distribution: Australia: New South Wales, Queensland and Victoria.
Type-species: Digaster lumbricoides Perrier, 1872.

Species (All Australian) :

1. Digaster anomala Jamieson, 1970b

2. Digaster armifera Fletcher, 1887a

3. Digaster bradburyi Jamieson, 1970b

4. Digaster brunneus Spencer, 1900

5. Perissogaster excavata Fletcher, 1888a

6. Digaster gayndahensis Spencer, 1900

7. Digaster lamingtonensis Michaelsen, 1916

8. Digaster longmani Boardman, 1932

9. Digaster lumbricoides Perrier, 1872

10. Digaster minor Spencer, 1900

1 1 . Perissogaster nemoralis Fletcher, 1 889a

12. Digaster perrieri Fletcher, 1889a

13. Perrisogaster queenslandica Fletcher,
1889a

Remarks: The trigastric genus Perissogaster Fletcher, 1887a, was included in Digaster
(Jamieson, 1963) because of the demonstration of intraspecific variation from two to three
gizzards in D. perrieri and because of general similarity of the two genera. The author
(Jamieson, 1970b) demonstrated the dichogastrin condition of posterior nephridia in
Digaster anomala (a species closely resembling the type-species, D. lumbricoides), in D.
bradburyi and in D. armifera and previously demonstrated megameronephridia occurring
alongside exonephric micromeronephridia, although funnels were not demonstrated for the
megameronephridia, in D. lamingtonensis, D. longmani and Perissogaster queenslandica
(vide Jamieson, 1963). On the other hand the latter paper showed the type-species of
Perissogaster, P. excavata, and also P. nemoralis, to lack stomate nephridia having only

76

MEMOIRS OF THE QUEENSLAND MUSEUM

dense astomate (exonephric ?) micromeronephridia in posterior segments. It thus appears
that the latter two species should be placed in a re-established Perissogaster, a genus which,
from its excretory system, appears to belong in the Megascolecini. Resurrection of Perisso-
gaster will be deferred pending further revision of the Digaster-Perissogaster complex.

Genus Didymogaster Fletcher, 1887

Setae 8 per segment, rather widely paired, all ventral, none penial. Clitellum occupying
XI 11 -XVIII. Prostates 1 pair, with small lobules, their pores (combined with the male
pores?) anterior on XVIII. Genital markings present. Female pores paired, median to a
lines, in XIV. Spermathecal pores conspicuous, 3 pairs intrasegmental in IX, X and XI.
First dorsal pores in 4/5.

Gizzards 2, moderately developed in VI and VII. Calciferous glands absent. Intestine
commencing in XVII. Dorsal blood vessel single; or double only intrasegmentally. Mero-
nephric with stomate exonephric megameronephridia median to micromeronephridia
posteriorly; tufted nephridia absent. Holandric; paired testis-sacs present, confluent
medianly and intersegmentally. Seminal vesicles in IX and XII. Ovaries and funnels in
XIII. Spermathecae in VII, VIII and IX, 2 segments in front of their pores; each with a
small diverticulum.

Distribution: Australia: New South Wales. New Zealand (?).

Type-species: Didymogaster sylvaticus Fletcher, 1887a.

Other Species: None known.

Remarks: Lee (1959) rightly considered the New Zealand record to be dubious. The
type-species was redescribed by Stephenson (1933) and distinction from Digaster was
upheld by Jamieson (1963). Stephenson did not observe stomate nephridia but these have
been demonstrated in this laboratory and in a configuration which indicates that Didymo-
gaster must be transferred to the Dichogastrini from the Megascolecidae s. Gates though
remaining isolated in the extremely unusual location of the spermathecal pores.

This unusual worm is common in the Audley National Park and in adjacent regions
of New South Wales where Livistona palms occur.

Genus Megascolides McCoy, 1878

Setae 8 per segment. Clitellum developed over at least four segments between XII
and XIX, usually annular. Male pores 1 pair, on XVIII, combined with the pores of a pair

MEGASCOLECOID EARTHWORMS

77

of tubular prostates, not always on papillae. Female pores on XIV ; paired, ventrolateral, or
unpaired, median ventral. Spermathecal pores 1-5 pairs, anterior to 9/10. Gizzard single,
in V, or VI or VII ; calciferous glands present or (typically) absent. Dorsal vessel usually
unpaired, occasionally paired; 2-5 pairs of hearts, the last in XII or XIII. Meronephridia
numerous in each segment; usually in lateral bands, occasionally concentrated in ventro-
lateral tufts. The median nephridium on each side in each posterior segment typically
possessing a preseptal funnel. Holandric; testis-sacs absent. Seminal vesicles in any or all
of IX, X, XI, XII. Penial setae present or absent. Spermathecae diverticulate.

Distribution; Australia including Tasmania. New Zealand (restricted to northern
North Island). North America.

Type-species: Megascolides australis McCoy, 1878.

Australian Species:

1 . Megascolides australis McCoy, 1878
(synonym Notoscolex gippslandicus
Fletcher, 1888b)

2. Megascolides diaphanus Spencer, 1900

3. Megascolides nokanenaensis Michaelsen,
1907b

4. Cryptodrilus polynephricus Spencer, 1895

5. Cryptodrilus tenuis Fletcher, 1889a

Remarks : The genus is a congeries which requires revision. Separation from Spencer-
iella solely by possessing eight rather than many setae per segment is unsatisfactory. The
type-species differs further from that of Spencer iella in lacking calciferous glands and in
having the last hearts one segment further posteriorly, in XIII, and it remains to be seen
whether additional and more significant features separate the two genera. Megascolides
was distinguished from Notoscolex by Michaelsen (1907b) by the unbranched condition of
the central canal of the prostates. Earlier (Michaelsen, 1900) he had distinguished it from
Notoscolex by possesion of a pair of ‘meganephridia’ in addition to ‘diffuse’ nephridia in
the hindbody and ignored the condition of the prostates which were stated to be tubular
or racemose in each genus, though the existence of racemose prostates in Megascolides was
questioned. The nephridia of the type-species of Notoscolex are described as small tufts of
glandular tubes attached to the coelomic wall and most conspicuous in some anterior
segments (Fletcher, 1887a) but there is no certainty that the Megascolides-condition of the
nephridia is not found posteriorly, as it is in some Notoscolex species and revision of the
two genera should be made paru passu. The type-species of the Notoscolex differs from
Megascolides australis in possessing calciferous glands but the significance of this difference,
at least in Australian earthworms, is uncertain as glands may be present or absent in closely
similar species of Megascolex (personal observations).

Apart from M. australis (the type-species) only M. nokanenaensis qualifies for inclusion
in Megascolides , and in the Dichogastrini, in having megameronephridia in addition to

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MEMOIRS OF THE QUEENSLAND MUSEUM

micromeronephridia in posterior segments. Of the remaining species, M. diaphanus,
Cryptodrilus tenuis and C . polynephricus have, respectively, two, three and five (stomate?)
nephridia on each side in each segment and their affinities are uncertain. They probably
have affinities with the Perionychini.

Twenty-two New Zealand species, nearly half of the generic total, are described by
Lee (1959). Very few of them are known to have median stomate nephridia and their
affinities appear to lie with the Megascolecini as is suggested, though not unequivocally,
by the dendrogram given by Lee (in press). It is therefore noteworthy that Gates (1940a)
divided the Indian species of Megascolides among the genera Scolioscolides , and Barogaster
(Dichogastrini s. mihi ) and Travoscolides (Megascolecini s. mihi ) and considered affinities
of the Assamese species M. antrophyes Stephenson, 1924, to lie with the Tonoscolex-
Nel/oscolex group (also Megascolecini s. mihi).

N. American species are described by Smith (1937) and Macnab and McKey-Fender
(1948).

Genus Notoscolex Fletcher, 1887a

Setae 8 per segment; prostates 1 pair, with branched system of ducts, externally race-
mose or sometimes appearing tubular, their pores (always?) combined with the male pores,
on XVIII. Female pores in XIV. Spermathecal pores 1-3 pairs, the last in furrow 8/9 (in
certain abnormal species in 7/8).

Gizzard in V or VI. Calciferous glands lacking or one to at least six pairs in the region
of XI-XVII. Intestinal origin in one of XV-XIX. Micromeronephridia only present (species
to be transferred to the Megascolecini), or (Dichogastrini) these associated with mega-
meronephridia.

Distribution: Australia. New Zealand (1 species). South India and Ceylon.

Type-species: Notoscolex camdenensis Fletcher, 1887a. (New South Wales).

Australian Species:

1. Megascolides albertsi Cognetti, 1910

2. Notoscolex attenuatus Boardman, 1931

3. Notoscolex brancasteriensis Michaelsen,

1910b

4. Notoscolex camdenensis Fletcher, 1887a

5. Megascolides earner oni Spencer, 1892a

6. Cryptodrilus campestris Spencer, 1895

7. Cryptodrilus dubius Spencer, 1892a

8. Cryptodrilus fastigatus Fletcher, 1889a

9. Notoscolex grandis Fletcher, 1887a

10. Notoscolex hortensis Michaelsen, 1907b

11. Megascolides hulmei Spencer, 1892

MEGASCOLECOID EARTHWORMS

79

12. Megascolides illawarrae Fletcher, 1889a

13. Cryptodrilus illawarrae Fletcher, 1889a

(homonym)

14. Megascolides insignis Spencer, 1892a

15. Cryptodrilus irregularis Spencer, 1895

16. Notoscolex jenolanensis Michaelsen, 1907c

17. Notoscolex leai Michaelsen, 1910b

18. Notoscolex leios Jackson, 1931

19. Notoscolex maecenatis Michaelsen, 1907b

20. Cryptodrilus mediocris Fletcher, 1889a

21. Notoscolex modestus Michaelsen, 1907b

22. Cryptodrilus mudgeanus Fletcher, 1889a

23. Megascolides obscurus Spencer, 1892a

24. Cryptodrilus officeri Spencer, 1895

25. Notoscolex prestonianus Michaelsen,

1907a

26. Megascolides (?) ( Notoscolex ) pygmaeus

Fletcher, 1889a

27. Cryptodrilus queenslandica Spencer, 1900

28. Notoscolex rubescens Michaelsen, 1907b

29. Cryptodrilus rusticus Fletcher, 1887a

30. Crytodrilus saccarius Fletcher, 1887b

31. Megascolides simsoni Spencer, 1895

32. Cryptodrilus simulans Fletcher, 1889b

33. Cryptodrilus singulars Fletcher, 1889a

34. Megascolides sinuosus Spencer, 1 892a

35. Notoscolex ( Trinephrus ) suctorius
Michaelsen, 1907b

36. Notoscolex ulladullae Boardman, 1931

37. Megascolides victoriensis Spencer, 1892a

38. Cryptodrilus wellingtonensis Spencer,
1895

Remarks: The above definition is based on Stephenson (1930) and a discussion of the
genus by Gates (1960), who removed N. pumila (Stephenson, 1931) as the type of Lenno-
scolex. Gates rightly states that Notoscolex is a congeries with no geographical justification
and that it is not based on the over-all similarity which is to be expected in species so closely
related as to belong in one genus. No attempt can be made here to revise this large genus.

The genus was placed in the Megascolecidae 5. strict, by Gates (1959) and was retained
in the Megascolecinae s. strict, by Sims (1966) despite the fact that the example he considered
( N . lunatus Gates, 1929) showed its closest affinities to lie with Eudichogaster (Octochaetidae
s. Gates; Dichogastrini s. mihi). As some Australian species have megameronephridia
median to micromeronephridia in posterior segments they must, despite their racemose
prostates, be referred to the Dichogastrini, a transfer supported by Sims’s computer ana-
lysis. The condition of the nephridia in the hind body in the type-species is unknown but its
similarity to the dichogastrin Notoscolex ulladullae suggests that the two species are strictly
congeneric.

Tribe PERlONYCHINi

Male and prostatic pores coincident, or {Diplot r etna part, New Caledonia) near to-
gether, on XVIII; sometimes with a single median combined male and prostatic pore.
Prostates one pair, tubular to racemose. Purely holonephric, or with meronephridia in a
varying number of segments anterior to holonephridia; never (?) with intestinal entero-
nephry.

Distribution: Nearctic: N. America; Queen Charlotte I. Neotropical: Guatemala;
Northern S. America. Oriental: India; Ceylon; Burma. Australia (including Tasmania).

80

MEMOIRS OF THE QUEENSLAND MUSEUM

New Zealand (and Chatham I.). New Caledonia. Subantarctic Islands : Auckland I . ;
Snares I. ; Stewart I.

Pontodrilus circummundane on sea shores in the warmer parts of the world, including
the Palaearctic.

Type-genus: Perionyx Perrier, 1872. (India).

Genera: (see Jamieson, 1971a).

Australian Genera: Diporochaeta Beddard, 1890; Fletcherodrilus Michaelsen, 1891;
Heteroporodrilus Jamieson, 1970a; Plutellus Perrier, 1873; Pontodrilus Perrier, 1874;
Pseudoperichaeta Jamieson, 1970a and Woodwardiella s. strict.

Remarks : Perionyx and Woodwardiella were the only genera of Gates’s Megascolecidae
which possessed holonephridia. Perionyx Perrier, 1872, restricted by Gates (1960) to
oriental species, includes some species in which there are several holonephridia (?) per
segment which are not, in Gates’s view, to be considered meronephridia as all are stomate.
The affinity of these requires further investigation. Where there is a pair of holonephridia,
the pores are in a single series on each side or alternate and nephridial vesicles (bladders)
are sometimes present. The type-species, P. excavatus , lacks vesicles and has nephropores
in a rather irregular, approximately midlateral rank on each side.

Comarodrilus Stephenson, 1915 (Cochin, India) is here placed in the Perionychini as
it has a pair of ‘meganephridia’ (presumably holonephridia, a term not employed at that
date) in each segment behind XII. In II to XII it possesses parietal and septal micromero-
nephridia. The nephridia have not been reinvestigated since Stephenson’s original descrip-
tion of the type-, and only, species, C. gravelyi , and absence of meronephridia behind XII
requires confirmation.

With regard to Diporochaeta , the nephridia of Diporochaeta davallia were described by
Jamieson (1970a). It is found, like the closely similar Pseudoperichaeta smithi, to have
anterior meronephric tufted nephridia with composite ducts but, whereas those of smithi
are limited to a single segment, those of D. davallia extend from segment II for about one
third of the length of the intestine; stomate holonephridia are present further posteriorly.
Lee (1959) records only holonephridia (with large ovoidal vesicles) for the type-species,
D. intermedia (Beddard, 1888) which is doubtfully congeneric with D. davallia.

Fletcherodrilus Michaelsen, 1891, here separated from Plutellus with which it was
synonymized by Michaelsen, 1910a, p. 22, has holonephridia only. The form and arrange-
ment of the nephridia and features such as the single median male pore indicate that the

M EG ASCOLECOID EARTHWORMS

81

type-species, F. unicus, is not congeneric with the type-species of Plutellus, P. heteroporus.
In a specimen of F. unicus from the Numinbah Valley, Queensland, examined by the writer,
the first nephridial bladder opens to the exterior anteriorly in III. In III-VIII, the terminal
bladder is a bilobed sac, with ental and ectal chamber, the terminal tubule of the nephridium
entering the ental chamber. In segment IX the ectal chamber is beginning to elongate as a
diverticulum or caecum. By XV the diverticulum reaches its maximum size, this being almost
twice the longitudinal extent of the segment. In XX posteriorly the diverticulum is an only
slightly elongate sac and is joined ectally by the widened, bladderlike end of the terminal
tubule. The nephropores form a single series on each side, inclines. The prostrate glands are
thick, tortuous, short tubes with a wide central lumen. The nephridia of Heteroporodrilus
are stomate holonephridia the postseptal portions of which commence in II and are equipped
with very large subspherical vesicles. The pores show a peculiar alternation which is remark-
ably constant in the ten species of the genus. Woodwardiella , the genus in which Hetero-
porodrilus was previously included, appears from an examination of the type-species
which, however, requires confirmation, to have tufted (meronephric ?) nephridia in some
anterior segments in addition to avesiculate stomate holonephridia and the nephropores
are apparently in a single series on each side. A very close relationship between Heteroporo-
drilus and Plutellus manifest us has been demonstrated (Jamieson, 1970a). The latter species
has subspherical vesicles as in Heteroporodrilus but the alternation of pores differs in detail
from that in the latter genus and agrees exactly with that described by Perrier (1873) for
the type-species of Plutellus , P. heteroporus . Despite a difference in the arrangement of
calciferous glands, the last glands being in XIII in manifestus and XII in heteroporus , the
two species are evidently congeneric, very closely related, and constitute Plutellus s. strict.
in the present work. Inclusion of Heteroporodrilus and Plutellus in the same tribe is therefore
justified.

Pontodrilus is totally holonephric and its nephridia commence in the region of the
clitellum. Nephropores are in a single row on each side (see p. 89). In no species have
nephridial vesicles been reported.

Pseudoperichaeta has a pair of anterior meronephric tufts with composite ducts follow-
ed in the succeeding segments by stomate, avesiculate holonephridia.

A series thus exists in the Perionychini, though it is not suggested that it is an actual
evolutionary lineage, in the direction of increasing development of meronephry. It com-
mences with total holonephry, as in Perionyx (most species), Plutellus, Fletcherodrilus,
Heteroporodrilus, and supposedly the type-species of Diporochaeta, and proceeds to modi-
fication of the first pair of nephridia as exonephric (astomate ?) meronephric tufted nephridia
in Pseudoperichaeta, to several pairs of (meronephric?) tufts in Woodwardiella s. strict and
finally to development of astomate tufted nephridia through many anterior segments in
Diporochaeta davallia. This entire series occurs in Australian species. Australia thus
harbours the most primitive known megascolecines. Although Pontodrilus (circummundane)
is totally holonephric, absence of preclitellar nephridia could conceivably have been

82

MEMOIRS OF THE QUEENSLAND MUSEUM

preceded by the development of meronephry in that region. This seems unlikely,
however, in view of apparent affinities with Plutellus and the demonstration by Hague
(1923) of anterior holonephridia in juveniles of Sparganophilus (Glossoscolecidae)
which similarly lacks preclitellar nephridia in the adult and is similarly amphibious.
If Stephenson’s account of Comarodrilus is correct, this Indian genus stands apart
from the Australian genera which also have partial meronephry in lacking tufted
nephridia.

It may be necessary, when knowledge of the Megascolecinae is more complete, to sub-
divide the Perionychini into further tribes as association on the presence of holonephridia
may represent a grade rather than a clade. However, as the holonephric condition is
evidently primitive relative to the meronephric condition, it is reasonable to conclude that
the genera of the Perionychini are cladistically more closely related one to the other than
they are to exclusively meronephric species. We have seen that some genera are morpho-
logically or phenetically closely linked and that other genera appear to be closely similar.
Thus Heteroporodrilus and Plutellus s. strict, are almost congeneric; Pseudoperichaeta and
at least one species of Diporochaeta and apparently Woodwardiella s. strict., are mutually
close. These two groups are sufficiently similar for constituent genera to have been confused
in the ‘classical’ work of Michaelsen and Stephenson. Pontodrilus is generally considered
to have affinities with Plutellus s. lat. Fletcherodrilus is near enough to Plutellus to have been
synonymized with it by Michaelsen while showing a number of remarkable similarities with
Perionyx (flattened form of the body ; an extreme condition of the tendency of male pores
to approach the ventral midline; reddish pigmentation; irregularity of posterior setae as a
possible prelude to the perichaetine condition in the latter genus). Perionyx in turn is closely
similar (Sims, 1966) to Diporochaeta. Knowledge of Plutellus s. lat., Diporochaeta, Perionyx
and Woodwardiella and of most genera of the Megascolecini is very rudimentary, however,
and changes in the internal classification of the Megascolecinae must be expected.

Genus Diporochaeta Beddard, 1890, emend. Michaelsen, 1900

Setae, at least in the mid and hind body, numerous (more than 8) per segment Clitellum
developed over at least 3 segments, between XIII and XVII; annular for most or all of its
length. Male pores 1 pair, on XVIII, combined with the pores of the single pair of prostates,
ventrolateral, not always on papillae. Female pores 1 pair, on XIV, ventrolateral. Sperma-
thecal pores paired, or unpaired and midventral in 2-5 intersegments, anterior to 9/10
Nephropores in a single sometimes irregular series on each side. Calciferous glands present
or absent. Gizzard sometimes absent; if present, single, in V, VI or VII. Dorsal blood vessel
unpaired ; hearts 2, 3 or 4 pairs, the last in XI, XII or XIII. Holandric or metandric; testis-
sacs absent. Prostates externally tubular or racemose or intermediate but (always ?) with a
central lumen which may or may not have side branches. Penial setae and copulatory
muscles usually absent. Spermathecae with or without diverticula. Possessing holonephridia
and with or without anterior tufted exonephric nephridia with composite ducts. Holo-
nephridia with (e.g. type-species) or without terminal vesicles.

MEGASCOLECOID EARTHWORMS

83

Distribution: Australia: Victoria, Tasmania, Queensland. New Zealand, including
Stewart, Chatham, Auckland and Snares Islands. South India.

Type-species: Perichaeta intermedia Beddard, 1888. (New Zealand).

Australian Species :

1. Perichaeta alsophila Spencer, 1892b

2. Diporochaeta apiocystis Stephenson, 1933

3. Diporochaeta arnoldi Spencer, 1900

4. Perionyx ( Diporochaeta ) athertonensis

Michaelsen, 1916

5. Perichaeta barronensis Fletcher, 1887b

6. Perichaeta bakeri Fletcher, 1888b

7. Perichaeta canaliculata Fletcher, 1888a

8. Perichaeta copelandi Spencer, 1892b

9. Diporochaeta davallia Spencer, 1900

10. Perichaeta dendyi Spencer, 1892b

11. Perichaeta dicksonia Spencer, 1892b

12. Perichaeta dilwynnia Spencer, 1895

13. Perichaeta dubia Spencer, 1892b

14. Perionyx {Diporochaeta) erici Michael-

sen, 1916

15. Diporochaeta euzona Spencer, 1900

16. Diporochaeta faucium Michaelsen, 1907b

17. Diporochaeta frosti Spencer, 1900

18. Diporochaeta grandis Spencer, 1900

19. Perichaeta irregularis Spencer, 1895

20. Perionyx lacustris Stephenson, 1924

21. Diporochaeta lindti Spencer, 1900

22. Perichaeta lochensis Spencer, 1892b

23. Diporochaeta manni Spencer, 1900

24. Diporochaeta mediocincta Spencer, 1900

25. Perichaeta moroea Spencer, 1895

26. Diporochaeta nemoralis Spencer, 1900

27. Perichaeta obscura Spencer, 1 892b

28. Perionyx ( Diporochaeta ) phalacrus

Michaelsen, 1916

29. Megascolex pritchardi Spencer, 1900

30. Diporochaeta richardi Spencer, 1900

31. Perichaeta richea Spencer, 1895

32. Perichaeta scolecoidea Spencer, 1895

33. Diporochaeta sedecimalis Michaelsen,

1907c

34. Perionyx ( Diporochaeta ) sigillatus

Michaelsen, 1916

35. Diporochaeta spenceri Michaelsen, 1907a

36. Pericheata tanjilensis Spencer, 1892b

37. Diporochaeta telopea Spencer, 1900

38. Perichaeta (?) terraereginae Fletcher,

1889b

39. Perichaeta walhallae Spencer, 1892b

40. Perichaeta yarrensis Spencer, 1892b

Remarks : The most recent account of Diporochaeta is that of Lee (1959) who described
the ten New Zealand species, bringing the total for the genus to approximately 63 species.
D. pellucida (Bourne, 1894) was transferred by Gates (1940a) to a new genus, Priodochaeta.
Michaelsen (1900) established Diporochaeta in its present form but later (1916, 1924a)
combined it with Perionyx, at the latter date as a subgenus. Stephenson (1923) separated
the two genera, while suggesting a close relationship. Gates (1959) placed Diporochaeta in
the Acanthodrilidae and Perionyx in the Megascolecidae but the two genera are here placed
in the same tribe of the subfamily Megascolecinae for reasons given on p. 82.

Morphological heterogeneity in the 40 Australian species is sufficient to suggest that
revision of the genus will necessitate establishment of several new genera. Jamieson (1970a)
reported the existence of anterior exonephric tufted nephridia in D. davallia although the

84

MEMOIRS OF THE QUEENSLAND MUSEUM

genus is defined by Stephenson (1930) as ‘purely meganephridial’, the sole supposed distinc-
tion from Spencer iel la . Transfer of D. davallia to Spenceriella is, however, insupportable as
the latter genus is meronephric throughout the body. Some species currently placed in
Plutellus also have anterior tufts and their distinction from species of Diporochaeta which
have the same condition solely because their setal arrangement is lumbricine is questionably
valid. Revision of Diporochaeta requires consideration of such species of Plutellus and both
genera are concomitantly being reviewed in this laboratory. It appears that Perionychella
must be resurrected for the type-species Perichaeta dendyi.

Genus Fletcherodrilus Michaelsen, 1891, emend.

Medium to large terrestrial worms (85-325 mm) with less than 160 segments. With
strong purplish parietal pigmentation. Prostomium slightly epilobous. Body not canaliculate
First dorsal pore 4/5 or 5/6. Setae in 8 longitudinal rows throughout or cd irregular posterior-
ly; some posterior segments occasionally with one or two supernumerary setae. Ventral
setal couples ( ab ) widely paired; dorsal couples ( cd ) distant, further apart than the two
couples of a side ( cd>bc ); dorsal median intersetal distance {dd) equal to half of the
circumference; dd>4cd. Penial setae absent. Nephropores in (/lines throughout. Clitellum
annular, occupying 4 to 5 segments beginning on XIII or XIV. The combined opening of
the male and prostatic ducts unpaired, midventral on a conical penis-like papilla. Accessory
genital markings absent. Female pores anteromedial to setae a of XIV, inconspicuous.
Spermathecal pores unpaired, midventral, 4 or 5, commencing at 4/5 or 5/6.

Some preclitellar septa thickened. Gizzard well developed, in VI but the post-gizzard
oesophagus may not commence until anterior VIII or IX. Calciferous glands lateral, sessile
pouches in XIII, XIV and/or XV with laminate internal folds but no partitions. Intestine
commencing in XVIII; typhlosole and caeca absent. Dorsal blood vessel continued onto the
pharynx. Dorsoventral commissural vessels in VII -XII ; those of X-XII forming latero-
oesophageal hearts which are unbranched ventrally; those of VII to IX slender, with dorsal
but no supra-oesophageal connectives and giving off parietal branches ventrally. Supra-
oesophageal vessel in IX (and further forward ?)-XII. Subneural vessel absent. Nephridia
stomate holonephridia; post-septal bodies commencing in II; each with a fairly large
elongate bladder which, after the first few, bears a lateral diverticulum. Testes and funnels
free, in X and XI ; seminal vesicles 2 to 4 pairs the last on the anterior face of XII. Ovaries
and funnel in XIII ; ovisacs absent. Prostates thickly tubular with large central lumen ; vasa
deferentia joining their ducts at mid-length. Spermathecae unpaired, each discharging
anteriorly in its segment ; duct shorter than the ampulla and bearing, ectally, two (some-
times one) digitiform diverticula.

Distribution: Widespread from the Richmond River, in New South Wales, to the
Cape York Peninsula, in Queensland.

Type-species: Cryptodrilus (?) unicus Fletcher, 1889a.

MEGASCOLECOID EARTHWORMS

85

Species (All Australian) :

1. Plutellus affinis Stephenson, 1933 3 Cryptodrilus fasciatus Fletcher, 1889b

2. Cryptodrilus (?) unicus Fletcher, 1889a

Remarks: Fletcherodrilus was placed by Michaelsen (1910a) in the synonymy of
Plutellus because he had found an Indian species, supposedly belonging to the latter genus,
in which the spermathecal pores were unpaired. The genus is here re-established as the
type-species is clearly not congeneric with the type-species of Plutellus ( P . heteroporus) or
its congener P. manifest us. The generic account has been augmented from the author’s
study of material of F. unicus from the Numinbah Valley and of F.facsiatus from Binna
Burra, Queensland. Highly distinctive features of Fletcherodrilus are the straight series of
nephropores, the presence of diverticula of the nephridial vesicles; the indefinite, lateral
calciferous glands and the large ratio dd:u.

Four species of the genus have been described three of which ( F . unicus (Fletcher,
1889a); F. purpureus (Michaelsen, 1889) and F. fasciatus (Fletcher, 1889b)) were first
assigned to Cryptodrilus. Michaelsen (1891) considered these to be ‘varieties’ with a new
variety, pelewensis, of one species F. unicus. Later (1900) he placed purpureus in the synony-
my of i F. unicus typicus ’ and pelewensis in F. unicus fasciatus. The author agrees that
purpureus is a synonym of unicus but considers that fasciatus should be reinstated as a
distinct species as the material from Binna Burra, examined in the present study differs
from unicus in having the gizzard more posteriorly situated and no calciferous glands in XV,
two differences noted also by Fletcher in the type-description of fasciatus. Michaelsen’s var.
pelewensis , with gizzard in VI and calciferous glands in XIII-XV presumably belongs in
unicus. It is hoped to settle the question of specific status for fasciatus in a separate publica-
tion based on additional material which has been acquired since this paper was commenced.

The third valid species, Plutellus affinis Stephenson, 1933, differs from the others in
having a single diverticulum only to each spermatheca.

Genus Heteroporodrilus Jamieson, 1970a

Moderate-sized to large terrestrial worms (52-580 mm long) with less than 300 and
usually less than 200 segments. With or without brownish pigmentation. Prostomium
variable from prolobous to tanylobous. Body with or without a narrow dorsal groove.
First dorsal pore in 5/6 or 6/7 (variable intraspecifically) or rarely in 7/8 or 8/9 (?), excep-
tionally with a rudimentary pore at 4/5. Setae in 8 regular longitudinal lines, commencing
with II. Ventral setal couples widely paired to distant (aa equal to or <2 . 5 ab ); setae of the
dorsal couples ( cd ) widely separated, always further apart than those of each ventral couple
and usually a greater interval, rarely slightly smaller, than the interval between the couples
of a side ( cd>ab and > or rarely slightly <bc); dorsal median intersetai distance {dd)< 0 . 3

86

MEMOIRS OF THE QUEENSLAND MUSEUM

of the circumference. Setae b and often setae a absent in XVIII; b rarely replaced by penial
setae. Nephropores conspicuous at the anterior borders of their segments, in d lines in II-IV
or V, in c lines for a few segments, then alternating in successive segments between d and c
(or exceptionally mid be) until in X, or less commonly within a segment or two of this,
alternation between b and d commences and is continued to the posterior extremity. Clitel-
lum annular, well developed on XIV-XVI, frequently extending through whole or part of
XIII and XVII, rarely impinging on XVIII. A pair of combined male and prostatic pores
on XVIII, in b lines or less commonly in ab. At maturity, accessory genital markings always
present in the vicinity of the male pores and frequently in the fore-body, sometimes forming
continuous longitudinal series; integumentary only, not represented by intracoelomic
glands. Female pores a pair anteromedial to setae a of XIV, inconspicuous, though some-
times in a common glandular field. Spermathecal pores 2-4 pairs in or very slightly lateral
to b lines, the last in intersegment 8/9.

Some preclitellar septa strongly thickened, gizzard vestigial to strong in V. Well
developed paired sessile ventrolateral extramural calciferous glands, 3 to 5 pairs, the last
always in XIII ; the two members of each pair almost contiguous midventrally ; the lumen of
each gland divided by numerous transverse lamellae. Intestine commencing in XV or,
rarely, XVI ; typhlosole and caeca absent. Dorsal blood vessel (always ?) continued onto the
pharynx. Dorsoventral commissural vessels in VI (sometimes V?) to XIII; those of X, or
(< oxleyensis ) XI, to XIII with connectives from a pair of efferent vessels which arise from the
oesophagus or from its calciferous glands in each of these segments ; the efferent (calciferous)
vessels discharging thence into a paired or single supra-oesophageal vessel. Commissurals
(hearts) of X-XIII differing from those more anteriorly situated in lacking ventral branches.
A paired or single suboesophageal vessel between the calciferous vessels continuous to the
anterior end of the body as a pair of ventrolatero-oesophageal vessels running median to
the hearts. Nephridia stomate holonephridia, their postseptal portions commencing in II;
the duct of each with a very large subspherical, thin walled muscular vesicle which lacks
diverticula; nephropore with a small sphincter. Testes and funnels in X and XI, free or in
unpaired pericardiac testis-sacs; seminal vesicles in IX and XII, rarely (mediterreus) in XI
and XII. Ovaries and funnels in XIII; ovisacs absent; septal pouches exceptionally present
in XIII. Prostates one pair, racemose, sometimes (intraspecific variation) externally ap-
proaching a tubular form; vasa deferentia joining their short muscular ducts entally to
ectally. Spermathecae discharing anteriorly in their segments; each with a distinct duct,
shorter than or slightly longer than the ampulla, bearing one, or less commonly, two or
more diverticula which sometimes are branched.

Distribution: Australia: In the basins of the Murray-Darling (in the vicinity of the
Gwydir, Barwon, Castlereagh, Lachlan and Darling Rivers); of the Wimmera River
(southern Victoria); and of some Eastern coastal rivers (Brisbane R., Paramatta R. and
headwaters of Mary R. and Albert R.).

Type-species: Cryptodrilus tryoni Fletcher, 1889b.

MEGASCOLECOID EARTHWORMS

87

Species (All Australian) :

1. Woodwardiella ashworti Stephenson, 1933

2. Cryptodrilus canaliculatus Fletcher, 1889a

3. Cryptodrilus cooraniensis Spencer, 1900

4. Woodwardiella dioecia Stephenson, 1933

5. Cryptodrilus fletcheri Beddard, 1887

6. Heteroporodri/us lamingtonensis Jamieson,

1970a

7. Cryptodrilus mediterreus Fletcher, 1888b

8. Cryptodrilus oxleyensis Fletcher, 1889a

9. Cryptodrilus shephardi Spencer, 1900

10. Cryptodrilus sloanei Fletcher, 1889a

11. Cryptodrilus tryoni Fletcher, 1889b

(synonym Woodwardiella youngi
Boardman, 1932)

Remarks : This is undoubtedly a natural assemblage closely related to Plutellus s. strict.
(see Jamieson, 1970a).

Genus Plutellus Perrier, 1873

Seatae 8 per segment. Male pores united with or near the pores of the single pair of
tubular prostates on XVIII. Holonephric ; nephridia present in front of the clitellum. Female
pores a pair, or sometimes single, on XIV. Spermathecal pores paired or single, in one
to five intersegments, the last in 8/9.

Gizzard well developed to vestigial, in V-VII ; sometimes in two segments, V-VI or
VI-VII. Calciferous glands absent or 2-5 pairs present in the region of X-XVI. Intestinal
origin in XIV-XIX; typhlosole present or absent. Hearts in X-XI, XII or XIII. Holandric
or meroandric; testis-sacs present or absent. Ovaries a single pair, in XIII. Penial setae
present or absent. Spermathecae with one or more diverticula.

Distribution: Australia, including Tasmania; New Caledonia; New Zealand; Auck-
land Is.; Queen Charlotte I.; the Pacific coastal strip of the United States; Guatemala;
northern South America; India; Ceylon; Burma.

Type-species: Plutellus heteroporus Perrier, 1873. (Pennsylvania (?)).

Australian Species:

1. Plutellus asymmetric us Michaelsen, 1907b

2. Megascolid.es attenuatus Spencer, 1892a

3. Megascolides bassanus Spencer, 1895

4. Plutellus blackwoodianus Michaelsen,

1907b

5. Plutellus Candidas Jackson, 1931

6. Plutellus carneus Michaelsen, 1907b

7. Plutellus dalgarangae Jackson, 1931

8. Plutellus decatheca Michaelsen, 1910b

9. Cryptodrilus ellisi Spencer, 1895

10. Megascolides eucalypti Spencer, 1900

11. Cryptodrilus frenchi Spencer, 1892a

12. Cryptodrilus gippslandicus Spencer, 1892a

13. Cryptodrilus hobartensis Spencer, 1895

14. Megascolides incertus Spencer, 1892a

15. Cryptodrilus insularis Spencer, 1895

(homonym, see Pontodrilus )

16. Cryptodrilus intermedins Spencer, 1892a

88

MEMOIRS OF THE QUEENSLAND MUSEUM

17. Plutellus levis Michaelsen, 1907b

18. Cryptodrilus lucasi Spencer, 1892a

18a. Cryptodrilus macedonensis Spencer, 1892a

19. Cryptodrilus manifestus Fletcher, 1889a

20. Megascolides manni Spencer, 1892a

21. Plutellus mendilai Michaelsen, 1907b

22. Cryptodrilus minor Spencer, 1892a

23. Plutellus murrayensis Michaelsen, 1907b

24. Cryptodrilus narrensis Spencer, 1892a

25. Megascolides roseus Spencer, 1892a

26. Cryptodrilus rubens Fletcher, 1888a

27. Plutellus schumanni Michaelsen, 1907b

28. Cryptodrilus semicinct us Fletcher, 1889b

29. Megascolides steeli Spencer, 1900

30. Plutellus strelitzi Michaelsen, 1907b

31. Cryptodrilus tanjilensis Spencer, 1892a

32. Notoscolex tasmanianus Fletcher, 1888b

33. Plutellus termitophilus Michaelsen, 1907b

34. Megascolides tisdalli Spencer, 1900

35. Notoscolex tuberculatus Fletcher, 1888b

36. Plutellus uncinatus Stephenson, 1933

37. Plutellus varicystis Jackson, 1931

38. Cryptodrilus victoriae Spencer, 1892a

39. Megascolides volvens Spencer, 1900

40. Megascolides warragulensis Spencer, 1 900

41. Plutellus wellingtonensis Michaelsen,

1907b

42. Plutellus whistler i Michaelsen, 1935b

43. Cryptodrilus willsiensis Spencer, 1892a

44. Plutellus wood war di Michaelsen, 1907b

Remarks: The above description considerably augments that of Stephenson (1930)
but serves to demonstrate a morphological heterogeneity which is inacceptable, whether on
phylogenetic or phenetic grounds, within a single genus. The characters which all species
of Plutellus supposedly have in common are italicised at the beginning of the description
but it should be noted that not even these are applicable to all species currently included in
Plutellus. Thus Gates (1962c, p. 187) mentions species in which the prostates open to the
exterior in XIX or XX, and P. macedonensis (Spencer, 1892a) has anterior tufted micro-
meronephridia (Horan, personal communication).

The Australian members of the genus are at present under revision in this laboratory
and it is hoped that subdivision among discrete genera of approximately 105 species making
up the global record will soon be practicable. Gates (1961) has already suggested that the
oriental species, which differ from the type-species in lacking calciferous glands, should be
placed in a separate new genus, though he has postponed this step until better character-
ization of the type-species becomes possible.

If, as in Heteroporodrilus , the nephropore arrangement seen in Plutellus heteroporus
and P. manifestus is taken as a generic character, it appears that none of the Australian
species other than manifestus can be included in Plutellus. Oriental Plutelli, all of which
lack calciferous glands, must presumably be excluded. Whether the American Plutelli
include species which are congeneric with the supposedly Pennsylvanian heteroporus is not
known.

The supposedly Pennsylvanian origin of P. heteroporus has been queried by Gates
(1961, p. 428). In view of the close similarity of manifestus (from New South Wales) it now
seems likely that the type-species is Australian. Two lots of specimens were mixed in the

MEGASCOLECOID EARTHWORMS

89

bottle in the Paris museum containing the types and Perrier described Australian specimens
of other genera in the same collections.

Genus Pontodrilus Perrier, 1874

Small to moderately large littoral (mainly marine) worms. Pigmentless. Prostomium
epilobous, rarely prolobous. Dorsal pores absent. Setae 8 per segment, commencing on II,
in regular longitudinal rows or posteriorly irregular. Nephropores inconspicuous, in b lines
(sometimes in a or c lines?) throughout the body. Clitellum annular, on XII, XIII-XVII,
XVIII. Combined openings of a pair of tubular prostates and male ducts on XVIII, in ab
or b \ a male duct joining each prostate gland at its junction with its duct, where the lumina
unite, or running in the gland to open into the ental end of the latter. Accessory genital
markings present behind the male pores. Female pores a pair on XIV, anterior to setae a.
Spermathecal pores 2 pairs, in 7/8 and 8/9, in b.

Gizzard rudimentary or absent, in front of the testis-segments. Intestine commencing
in XIV ( ?) to XX ; calciferous glands, intestinal caeca and typhlosole absent. Dorsal vessel
single, complete; dorso-ventral commissural vessels in V-XIII; those of X-XIII (always?)
with supra-oesophageal connectives; paired extraoesophageal vessels passing onto the
oesophagus in XII XIII and uniting midventrally in XIV-XV. Supra-oesophageal vessel
typically (always?) present. Subneural vessel absent. Nephridia all avesiculate holo-
nephridia; absent from the preclitellar region. Testes and funnels in X and XI; seminal
vesicles in XI and XII. Penial setae absent (unless lacustris be included). Ovaries in XIII.
Spermathecae diverticulate.

Distribution: Circummundane on seashores in the tropics and warmer temperate
regions (extending into the freshwater littoral in Lake Chilka, India). The lacustrine P.
lacustris (New Zealand) and the terrestrial P. agnesae (India) are excluded from the above
definition.

Type-species: Lumbricus litoralis Grube, 1855. (Southern France).

Australian Species:

1. Pontodrilus litoralis (Grube, 1855) 2. Pontodrilus bermudensis Beddard, 1891

Remarks: The above description augments that of Stephenson (1930).

The following four species can unequivocally be assigned to Pontodrilus’.

(1) P. litoralis (Grube, 1855) (syn. : P. marionis Perrier, 1874; P. crosslandi Beddard,
1905; P. albanyensis Michaelsen, 1907b, Jackson, 1931 (SW. Australia)).

90

MEMOIRS OF THE QUEENSLAND MUSEUM

(2) P. bermudensis Beddard, 1891 (recorded from the Great Barrier Reef by Stephen-

son, 1931) (syn. : Pontoscolex arenicola Schmarda, 1861 (part); Cryptodrilus
insularis Rosa, 1891 ; Pontodrilus arenae Michaelsen, 1892; Pontodrilus hesperi-
dum Beddard, 1894; Pontodrilus michaelseni Eisen, 1895 \ Pontodrilus ephippiger
Rosa, 1898 (recorded from SW. Australia by Michaelsen, 1907b); Pontodrilus
laccadivensis Beddard, 1903).

(3) P. matsushimensis lizuka, 1898 (syn.: Pontodrilus chathamianus Benham, 1901).

(4) P. gracilis Gates, 1943.

Two further species are only doubtfully assignable to Pontodrilus, namely the terrestrial
P. agnesae Stephenson, 1915, (India) and the lacustrine Plutellus lacustris Benham, 1903,
(New Zealand) which was referred to Pontodrilus by Michaelsen, 1907b. Michaelsen (1922,
1928a) included Pontodrilus as a sub genus in Plutellus but Stephenson (1930) retained the
two genera. Although Pontodrilus may have affinities with some species currently included
in Plutellus, accordance with a restricted Plutellus (p. 88) is small and separation of the
two genera is here endorsed.

Pontodrilus lacustris shows the following departures from the four Pontodrilus species
from which the generic definition above has been drawn: it has four pairs of spermathecal
pores; the last hearts are in XII ; seminal vesicles are in IX and XI and penial setae are
present. Each of these features is known to vary intragenerically in other genera but the
combination sets lacustris apart from the four marine littoral species as does its habitat
deep in a glacial lake. On the other hand, Lee (1959) states that 'Plutellus' parvus Lee, 1959,
is more similar to lacustris than it is to the two other New Zealand species assigned to
Plutellus. Several features, including presence of calciferous glands in XIII separate parvus
from lacustris , however, and while parvus could justifiably be made the type of a new genus,
it is questionable that lacustris could be placed in the latter. Pontodrilus lacustris must
therefore be regarded, until Pontodrilus and Plutellus are revised, as a species incertae sedis.
The above conclusions are endorsed by the computer dendrogram given by Lee (in press).
In it P. lacustris clusters with the core of Rhododrilus while Pontodrilus matsushimensis ,
a true Pontodrilus, clusters with Megascolecini s. strict., Plutellus parvus clusters with an-
other group of Rhododrilus species which have a high but apparently not generic affinity
with the typical cluster. The Rhododrilus affinities of Pontodrilus lacustris and Plutellus
parvus may suggest that there are exceptional members of the Acanthodrilini in which
association of male and prostatic pores has occurred on XVIII rather than, as in Rhodo-
drilus, on XVI 1 but the present definition of the Acanthodrilini will be retained pending
further evidence.

Pontodrilus agnesae is too poorly known to be placeable with certainty either in Ponto-
drilus or in the Indian section of Plutellus s. lat. and must, again, be regarded as incertae
sedis at least until the type-series in the British Museum is re-examined.

MEGASCOLECOID EARTHWORMS

91

Genus Pseudoperichaeta Jamieson, 1970a

Typically moderate-sized terrestrial worms (<150 mm) with less than 200 segments;
tanylobous; dorsal pores commencing in or anterior to 4/5. Setae 8 per segment; some
rows irregular posteriorly ; the ventral couples widely paired, the dorsal setal couples closely
paired but translocated far dorsally so that the dorsal median intersetal distance is less than
cd. Nephropores in a single series which is slightly oblique relative to the setal lines, on each
side. Clitellum annular, occupying approximately 4 whole segments and anterior to a pair of
combined male and prostatic pores which are situated on XVIII in ab. Accessory genital
markings present in the forebody and in the vicinity of the male pores. Female pores antero-
medial to setae a of XIV. Spermathecal pores 4 pairs, the last in 8/9.

Some anterior septa moderately thickened. Gizzard in V. Extramural calciferous
glands absent but oesophagus vascularised and swollen in the region of XIII-XVI. Intestine
commencing in XIX. Dorsoventral commissural vessels in V-XJ1 ; hearts in X-XII, latero-
oesophageal. Nephridia holonephridia (stomate and avesiculate) with the exception of the
first pair which are tufted meronephridia, each tuft having a composite duct. Testes and
funnels in X and XI, free; seminal vesicles in IX and XII. Prostates racemose. Ovaries and
funnels in XIII. Spermathecae diverticulate.

Distribution: The Yarra Basin, Victoria.

Type-species: Cryptodrilus smithi Fletcher 1889b. (Synonym Megascolides punctatus
Spencer, 1900).

Other Species: None known.

Remarks: The genus shows affinities with Woodwardiella s. strict, and with Diporo-
chaeta davallia (see Jamieson, 1970a).

Genus Woodwardiella Stephenson, 1925, emend. Jamieson, 1970a

Small to moderate-sized terrestrial worms (20-80 mm) with less than 150 segments.
With or without parietal pigmentation. Prostomium variable from proepilobous to tany-
lobous. Body rarely ( tesselatus ) canaliculate dorsally. First dorsal pore in (3/4) 5/6 or 6/7,
exceptionally ( callichaeta ) postclitellar. Setae in 8 longitudinal rows throughout or {tessel-
atus) some rows irregular; ventral setal couples widely paired; setae of the dorsal couples
(cd) widely separated, little if at all closer together than the two couples of a side; dorsal
intersetal distance <0 . 3 of the circumference. Penial setae present in XVIII or exceptionally
{tesselatus) absent. Nephropores in a single series on each side. Clitellum annular on (XIII)
XIV -XVII (XVIII). A pair of combined male and prostatic pores on XVIII in a or ab.

92

MEMOIRS OF THE QUEENSLAND MUSEUM

Accessory genital markings always present at maturity within the region of segments X-
XIV and sometimes in the vicinity of the male pores. Female pores anteromedial to setae a
of XIV, inconspicuous. Spermathecal pores 2 pairs, in a or ab lines, in intersegmental fur-
rows 7/8 and 8/9.

Some preclitellar septa moderately thickened. Gizzard well developed, in V. Extra-
mural calciferous glands absent. Intestine (always?) commencing in XVII or XVIII; typh-
losole and caeca unknown. Last hearts in XII, commissurals at least sometimes beginning
in VI. Supra-oesophageal vessel present (?). Nephridia holonephridia becoming tufted but
apparently remaining exonephric in III and IV ( callichaeta ); vesicles unrecorded. Testes
and funnels in X and XI ; seminal vesicles in IX and XII or in XII only. Ovaries and funnels
in XIII; ovisacs unknown. Prostates racemose; vasa deferentia joining the prostate ducts
entally. Spermathecae each with a single diverticulum ; the duct shorter than the ampulla.

Distribution: Southwestern Australia (Swan and Serpentine River Basins); Tasmania
(Mt Olympus and, mortoni, Dee Bridge); Victoria (Healesville by the Yarra), if W. healesi
be included.

Type-species: Woodwardia callichaeta Michaelsen, 1907b.

Species (All Australian) :

1. Woodwardia affinis Michaelsen, 1907b

(synonym Woodwardiella magna
Jackson, 1931)

2. Woodwardia callichaeta Michaelsen, 1907b

Species inquirenda

6. Woodwardia healesi Michaelsen, 1923 7. Cryptodrilus mortoni Spencer, 1895

3. Woodwardia libferti Michaelsen, 1907b

4. Woodwardia molaeleonis Michaelsen,

1907b

5. Cryptodrilus tesselatus Spencer, 1895

Remarks: Species 1-4 occur in southwestern Australia and are undoubtedly closely
related. The Tasmanian W. tesselatus appears to have close affinities with the latter species
but knowledge of all species is insufficient for conclusive demonstration that tessalatus is
congeneric. To avoid erection of inadequately characterized genera, Jamieson (1970a)
included the Victorian W. healesi and Tasmanian W. mortoni as species inquirenda.

Tribe MEGASCOLEC1NI

Male and prostatic pores coincident on XVI 1 1 (rarely XVII) ; prostates one pair, race-
mose or (less commonly) tubular. Purely meronephric; median stomate nephridia, if
present, opening into the intestine.

MEGASCOLECOID EARTHWORMS

93

Distribution: Palaearctic: China; Korea; Japan. Oriental: India; Ceylon; Anda-
man Is.; Assam; Burma; Annam; Java; Sumatra; Philippines; Moluccas. Australian:
Australia and Tasmania; Norfolk Island; New Guinea. New Caledonia. New Zealand.
Endemicity from the Mariannas, Bismarck Archipelago, the Solomons, and eastwards
across the Pacific Ocean remains to be established. Some species (Pheretima, Lampito )
widely peregrine.

Type-species: Megascolex Templeton, 1844.

Genera: (see Jamieson, 1971a).

Australian Genera: Digaster Perrier, 1872 (part only, excluding the type-species?);
Megascolex Templeton, 1 844 ; Notoscolex Fletcher, 1 887a (part, excluding the type-species ?) ;
Pheretima Kinberg, 1866.

Remarks: The tribe Megascolecini has been delimited so as to include only non-dicho-
gastrin genera of the Megascolecinae which totally lack holonephridia, that is, in which
total meronephry has supervened. Again although close mutual similarity of some genera
can be discerned, the tribe possibly represents a grade (acquisition of total meronephry)
rather than a single clade or monophyletic group and it is noteworthy that from Sims’s
computer analysis some genera appear to be morphologically closer to Perionychini or
Dichogastrini than they are to other genera of the Megascolecini. Nearest neighbours of
genera considered in Sims’s analysis as shown in the matrix of coefficients of similarity were
as follows (D=Dichogastrini, P Perionychini, M Megascolecini): Pheretima-Lampito
(both M); Didymogaster (D)-Heteroporodrilus (P); Lampito ( M)-Heteroporodrilus (P);
Plutellus s. lat. ( P)-Acanthodrilus (but the type-species of Plutellus is unquestionably close
to Heteroporodrilus). Unfortunately the type-genus Megascolex was not considered and it
appears that it was rarely feasible to use type-species for the various genera. Of the twelve
genera of the Megascolecini, six form a group (a Pheretima-group) which is unified by a
combination of characteristics of the excretory system. The 6 genera are: Megascolex (a
campestris-g roup of species, excluding the type-species); Pheretima (East Asia, including
China and Japan; Indonesia; Australia and widely spread); Lampito (India); Nellogaster
(Ceylon, S. India?); Plionogaster (?) (Philippines and Moluccas); and, hitherto placed in
the Octochaetidae, Travoscolides Gates, 1940a (India). In all of these there are tufts of
astomate micromeronephridia (with either composite or common ducts) opening into the
foregut (‘pharyngeal nephridia’) together with stomate meronephridia opening into the
intestine and consisting of one to many pairs per segment, the intestinal enteronephric
nephridia of separate segments being interconnected by one or two longitudinal excretory
ducts. In addition a pair to many exonephric micromeronephridia, which in Travoscolides
and some species of Megascolex (e.g. M. cochinensis) form tufts, are always present anterior-
ly and in all but Travoscolides extending into the intestinal region (Bahl, 1926b, 1946, 1947;
Gates, 1940a, 1943, 1945; Vidya Vati, 1947). Inclusion of Plionogaster in the group is

94

MEMOIRS OF THE QUEENSLAND MUSEUM

questionable as pharyngeal nephridia have not been demonstrated. Despite the unity of the
Pheretima- group and its distinction in these nephridial characters from the remainder of
the genera here included in the Megascolecini, it does not at present seem advisable to
separate it taxonomically from the residue.

With regard to the remaining genera, Gates (1939a) suggests that Tonoscolex has arisen
directly from Nelloscolex by constriction olf of extramural calciferous pouches, and Bahl
(1947) places Tonoscolex in a morphological sequence culminating with Travoscolides.
Thus inclusion of Tonoscolex and Nelloscolex in the group of genera here termed the
Megascolecini has considerable justification. Megascolex caeruleus (the type-species) and
M. templetonianus which lack intestinal enteronephry may be integrated with such a se-
quence as primitive (simplified?) forms. If M. caeruleus were placed in a separate tribe the
tribe containing Pheretima would require renaming. Of twenty four species of Megascolex
which have been closely examined, 19 species have pharyngeal nephridia and intestinal
enteronephry, while 5 species, including the type-species, are purely exonephric in the
intestinal region but have pharyngeal enteronephry (Bahl, 1942, 1946, 1947; Yidya Vati,
1947). This suggests a basis for restriction of this very large genus to species lacking in-
testinal enteronephry. The reader is referred to Bahl (1947) for details of the series of
progressively more highly developed enteronephry from Megascolex sarasinorum etc.,
through Tonoscolex, Megascolex cochinensis, Megascolex camp ester to Pheretima, in turn,
and thence to Travoscolides. In Travoscolides the two longitudinal supraintestinal canals of
Pheretima are represented by a single canal which is no longer supraintestinal but has
become embedded in the typhlosole. The enteronephric nephridia commence in segment
XV, number 20-24 per segment, have preseptal funnels and, in contrast to Pheretima , are
not accompanied by exonephric nephridia. A pair of pharyngeal tufts, each with a common,
not composite, duct is situated in segment V, and in each of eight segments behind this
is a pair of similar but exonephric tufts. Several species of Megascolex, including M.
cochinensis, have a similar arrangement of one pair of pharyngeal tufts followed by 8 or 9
pairs of exonephric tufts though with composite ducts.

The excretory system of Lennoscolex (type-species Woodwardiella pumila Stephenson,
1931) is meronephric and lacks nephrostomes. Behind the clitellum there is a pair of clusters
of 4-6 small exonephric nephridia in each segment (Gates, 1960). This small South Indian
genus stands apart from the Pheretima- group.

Genus Megascolex Templeton, 1844

Setae, at least in the middle and hinder parts of the body numerous (more than 8) in
each segment. Prostates 1 pair, racemose, their pores on XVI II or, exceptionally, an adjacent
segment; sometimes associated with penial setae. Female pores paired or exceptionally un-
paired. Spermathecal pores usually 1-5 pairs, between IV and IX (the exceptions are the
few cases where the pores are fused in the midline, or where they are numerous on each side
in each segment). One gizzard in V, VI, or VII. Calciferous glands present or absent.

MEGASCOLECOID EARTHWORMS

95

Meronephric, with or without enteronephric nephridia. Holandric, rarely metandric;
testis-sacs present or absent. Ovaries in XIII. Spermathecae with or without free diverticula.

Distribution: Ceylon and India. Australia. New Caledonia. New Zealand. Norfolk
Island. Annam.

Type-species: Megascolex caeruleus Templeton, 1844.

Australian Species:

1 . Megascolex albanyensis Michaelsen,

1907b

2. Megascolex albidus Jackson, 1931

3. Megascolex andersoni Spencer, 1900

4. Perichaeta attenuata Fletcher, 1889a

5. Perichaeta australia Fletcher, 1887a

6. Trichaeta australis Spencer, 1900

(homonym)

7. Perichaeta austrina Fletcher, 1887b

8. Megascolex bistichus Michaelsen, 1907b

9. Megascolex colliensis Michaelsen, 1 907b

10. Perichaeta coxi Fletcher, 1887a

11. Megascolex crateroides Boardman, 1943

12. Perichaeta dorsalis Fletcher, 1888b

13. Perichaeta enormis Fletcher, 1889a

14. Perichaeta exigua Fletcher, 1888a

1 5. Megascolex fardyi Spencer, 1900

16. Megascolex fecunda Fletcher, 1888a

17. Perichaeta fielderi Spencer, 1892b

18. Megascolex fletcheri Michaelsen, 1907

19. Megascolex fletcheri Shannon, 1920
(homonym)

20. Perichaeta frenchi Spencer, 1 892b

21. Perichaeta frosti Spencer, 1892b

22. Megascolex fuscus Michaelsen, 1916

23. Megascolex galei Michaelsen, 1907b

24. Perichaeta goonmurk Spencer, 1892b

25. Perichaeta gracilis Fletcher, 1887b

26. Perichaeta halli Spencer, 1 892b

27. Perichaeta hamiltoni Fletcher, 1888a

28. Megascolex harveyensis Michaelsen,

1907b

29. Megascolex heterochaetus Michaelsen,

1916

30. Perichaeta hoggi Spencer, 1 892b

31. Megascolex jenolanensis Boardman, 1943

32. Megascolex illidgei Spencer, 1900

33. Megascolex imparicystis Michaelsen,

1907b

34. Perichaeta indissimilis Fletcher, 1889a

35. Megascolex inermis Stephenson, 1933

36. Megascolex larpentensis Spencer, 1900

37. Perichaeta lateralis Spencer, 1892b

38. Megascolex lob ul at us Spencer, 1900

39. Megascolex longicystis TN icholls and

Jackson, 1926

40. Perichaeta macleayi Fletcher, 1889a

41 . Perichaeta macquariensis Fletcher,

1889b

42. Megascolex mediaeviae Michaelsen,

1907b

43. Megascolex minor Spencer, 1900

44. Megascolex mjobergi Michaelsen, 1916

45. Megascolex monostichus Michaelsen,

1907b

46. Megascolex montanus Spencer, 1900

47. Perichaeta monticola Fletcher, 1888a

48. Megascolex newcombei Beddard, 1887

49. Megascolex parvus Michaelsen, 1916

50. Megascolex purpurascens Michaelsen,

1907b

51. Perichaeta raymondiana Fletcher, 1888a

52. Megascolex rodwayi Stephenson, 1931

53. Perichaeta rubra Spencer, 1892b

54. Perichaeta steeli Spencer, 1892b

55. Perichaeta stirlingi Fletcher, 1888a

56. Megascolex swarbricki Nicholls and
Jackson, 1926

96

MEMOIRS OF THE QUEENSLAND MUSEUM

57. Perichaeta sylvatica Spencer, 1892

58. Megascolex syndetoporus Jackson, 1931

59. Perichaeta tasmanica Spencer, 1895

60. Perichaeta tenax Fletcher, 1887b

61. Megascolex terangiensis Spencer, 1900

62. Megascolex torbayensis Michaelsen,

1907b

63. Megascolex whistleri Michaelsen, 1907b

64. Megascolex wiburdi Boardman, 1943

65. Perichaeta wilsoniana Fletcher, 1888a

66. Megascolex zietzi Michaelsen, 1907c

Remarks: An attempt is made above to augment the brief description given by
Stephenson (1930) but this very large genus remains one of the major problems in oligo-
chaete taxonomy. It is clearly a polyphyletic congeries in need of extensive revision. The
type species is from Ceylon. It has 2 pairs of spermathecal pores, in 7/8 and 8/9; the gizzard
in V; no calciferous glands; is holandric, though with seminal vesicles in XII only; no
penial setae; and each spermatheca has a very small diverticulum concealed in the duct.

Gates (1938) reinstated the genus Lampito which Michaelsen (1900) and Stephenson
(1930) considered inseparable from Megascolex.

Indian species are described by Stephenson (1923, 1924, 1925a, b) and Aiyer (1929);
from Ceylon by Michaelsen (1900, 1909a, 1910a) and Gates (1941); from Assam by
Stephenson ( 1922); from Australia by Spencer (1900), Michaelsen (1907b, c; 1916), Shannon
(1920), Nicholls and Jackson (1926), Jackson (1931), Stephenson (1933), Boardman (1943);
and from New Zealand by Lee (1959).

Some species, for instance the Eastern Australian Megascolex dorsalis (Fletcher, 1888b),
are very closely similar to the genus Pheretima.

Genus Pheretima Kinberg, 1866

Digestive system without supraintestinal and calciferous glands (calciferous tissue in
low ridges that are not lamelliform in region of XIII?) but with gizzard that develops in
VIII. Vascular system with unpaired dorsal, ventral and supraoesophageal trunks, a sub-
neural adherent to the parietes, paired extra-oesophageal trunks median to hearts, (latero- ?)
oesophageal hearts in some of segments X-X1IL Excretory system meronephric; paired
clusters of astomate nephridia in LV-VI with ducts opening into the pharynx; astomate
exonephric, very small, V-shaped parietal nephridia numerous in each segment back from
III; larger stomate nephridia with funnels in the same segment as body of tubule, on both
sides of septa from XVI posteriorly, joining postseptal canals that pass to longitudinal
supraintestinal excretory ducts opening at frequent intervals into gut. Setae numerous, in a
circle at the equator of each segment from II posteriorly. Ovaries fan-shaped and with
several egg-strings. Male pores postclitellar. (Female always intraclitellar ?). Testes and

MEGASCOLECOID EARTHWORMS

97

male funnels enclosed in testis-sacs; seminal vesicles postseptal. Spermathecae diverticulate
and pregonadial. Prostates racemose, of mesoblastic origin, with ducts joined entally by
vasa deferentia.

Distribution: Andaman Islands. The mainland of Asia from the Chindwin-Irrawaddy
axis of Burma east through Yunnan and Szechuan provinces of China, including Korea and
Japan, thence south into New Guinea, Java and Sumatra. Endemicity from the Marianas,
Bismarck Archipelago, the Solomons, and New Caledonia eastwards remains to be
established. Peregrine species throughout the world but none yet recorded from Alaska,
Greenland, the Sahara, or Arabia.

Type-species: Pheretima montana Kinberg, 1866.

Australian Species:

1. Perichaeta queenslcmdica Fletcher, 1887b

Remarks: The writer is indebted to Professor G. E. Gates for the above, hitherto
unpublished generic description.

Several peregrine species have been found in Australia and Sims and Easton are
currently studying these as part of a taxonometric revision of the genus. Pheretima queens-
landica (Fletcher, 1887b) is questionably indigenous in Australia. In the author’s experience
introduced species of Pheretima are now the most commonly found of all earthworm
species where ground has been cultivated or natural vegetation removed.

Pheretima is distinguished from Megascolex by the location of the gizzard. It is expected
that the revision by Sims and Easton will provide more substantial differences. The
Australian Megascolex dorsalis despite its more anterior gizzard appears to the author to
have closer affinities with Pheretima than with the type-species of Megascolex. Pheretima is
the largest genus in the Oligochaeta, with some 800 recorded species, a number which will be
very greatly reduced by synonymy in the above-mentioned revision. The anatomy of
Pheretima posthuma, including that of the excretory system, has been described in great
detail by Bahl (1926a). Gates (1960) after many years of study of a large number of species
found no departure in the excretory system from the posthuma- type but it remains to be
seen whether the whole genus is uniform in this respect.

Michaelsen (1928b, 1934) recognized six subgenera as follows (slightly modified):

Archipheretima: No distinct creeping-sole on the ventral surface of the body. Clitel-
lum occupying more than 3 segments. Intestine without caeca. Holandric. Sperma-
thecae not multiple. Borneo, Philippines.

98

MEMOIRS OF THE QUEENSLAND MUSEUM

Pheretima : No distinct creeping-sole. Clitellum restricted to X1V-XVL. Intestine
usually with caeca. Holandric with or without copulatory bursae. Spermathecae
only exceptionally multiple. Southeast Asiatic and Malayan regions.

Metapheretima: No distinct creeping-sole. Clitellum restricted to XIY-XVI. Metan-
dric. No copulatory bursae. New Guinea, New Britain, North Queensland.

Parapheretima : No distinct creeping-sole. Clitellum restricted to XIV-XVI. Intestine
with or without caeca. Holandric. Copulatory bursae present, into which open
glands with muscular walls. New Guinea.

Planapheretima : A distinctly differentiated creeping-sole on the ventral surface
marked by an extremely dense crowding of setae. Clitellum occupying more than
three segments. Intestine without caeca. Holandric. No copulatory bursae. Sperma-
thecae not multiple. Borneo.

Polypheretima : No distinct creeping-sole. Clitellum restricted to the three segments
XIV-XVI. Intestine without caeca. Holandric. Copulatory bursae present or absent.
Spermathecae multiple, in pairs of transverse groups of 2 or more. Caroline Islands,
Sangir Islands (introduced?). Celebes. Borneo. Malay Peninsula. Lombok. Aru
Islands (introduced?). New Guinea.

The validity of these subgenera seems somewhat questionable.

Genus Spenceriella Michaelsen, 1907b

Prostomium zygolobous, prolobous or epilobous. Setae more than 8 per segment
throughout the body or in the postclitellar region; where multiplied, 12-72 per segment.
Penial setae absent. Clitellum annular or saddle-shaped, occupying 3-6 segments in XIII-
XVIII. Genital markings present. Prostates 1 pair, tubular though often with lateral
canaliculi from the central lumen, their pores combined with the male pores on XVIII.
Female pore unpaired, midventral, or paired, in XIV. Spermathecal pores 1-3 pairs, in
6/7, 7/8—8 /9, or 7/8 only. Dorsal pores (always?) present; location of the first pore variable.

Gizzard large, in V. Discrete (but not stalked?) calciferous glands 4 pairs, in X-XIII,
or forming a single or somewhat intersegmentally constricted dilatation on each side in 2-3
of segments XI -XIV ; always present in XIII. Intestine commencing in XVI or (maplestoni)
XVII. Hearts 3-7 pairs, the last in XII or XIII. Supra-oesophageal vessel (always?) present.
Meronephridia in broad or narrow lateral parietal bands; pharyngeal nephridia present in
Australian species only. Holandric or metandric; testis-sacs absent. Seminal vesicles IX
and XII, XI and XII or XII only. Ovaries and funnels in XIII. Spermathecae each with
(i notabilis ) a long tubular diverticulum or with 1 or 2 small diverticula.

MEGASCOLECOID EARTHWORMS

99

Distribution: Australia: Victoria. New Zealand: northern districts and neighbouring
coastal islands.

Type-species: Diporochaeta notabilis Spencer, 1900.

Australian Species :

1. Diporochaeta maplestoni Spencer, 1900 2. Diporochaeta notabilis Spencer, 1 900

Remarks: Michaelsen (1907b) erected Spenceriella for four species formerly placed in
Diporochaeta but distinguishable by their meronephry, viz. D. notabilis and D. maplestoni
(Australia), both of Spencer, 1900, and D, gigantea and D. shakespeari (New Zealand),
both of Benham, 1906. His tentative inclusion of Perichaeta lateralis Spencer, 1892b,
appears unwarranted as it has ‘leaf-shaped’ prostates which, though branching of the ducts
is not recorded, seem to necessitate inclusion in Megascolex as at present defined. Subse-
quently (1907b) he added an Indian species, S . duodecimalis. S. shakespeari was shown by
Lee (1962) to be a synonym of S. (= Megascolex) antarctica Baird, 1871. Michaelsen (1916)
transferred the New Zealand species to Megascolex because of lateral branching of the cen-
tral prostatic lumen in S. shakespeari. Stephenson (1923) accepted this change but later
(1930) again included the New Zealand species. Gates (1958) erected Celeriella for the
Indian species, a genus retained in the present work. Lee (1959) was unaware of Gates’s
paper and retained the Indian species in Spenceriella while adding two further New Zealand
species, S. argillae and S. pallida. His definition of the genus is partly erroneous with regard
to the Australian species and a new definition of the genus has therefore been given above.

S. gigantea (Benham, 1906) from New Zealand, is known to have median megamero-
nephridia posteriorly and thus qualifies for inclusion in the Dichogastrini. It may be place-
able in a perichaetine section of Megascolides.

Even after exclusion of S. gigantea , Spenceriella shows more heterogeneity than seems
acceptable in a single genus and clearly requires revision. The type-species is unique in the
diverticulate form of its calciferous glands and both Australian species differ from the
New Zealand species in their shorter clitellum, significantly smaller numbers of setae per
segment (maximally 14 and 24) and, if Spencer’s account be correct, in possessing entero-
nephric pharyngeal nephridia.

Affinities with Diporochaeta and Megascolides are discussed under those genera.

ACKNOWLEDGMENTS

The author thanks Mr R. W. Sims of the Annelida Section, British Museum, Dr B. J.
Smith of the National Museum of Victoria, Miss Elizabeth Pope of the Australian Museum,

100

MEMOIRS OF THE QUEENSLAND MUSEUM

and Dr D. D. Ryde of the Western Australian Museum for providing the opportunity to
examine material which has been the subject of this paper. Mr Sims and Professor R. O.
Brinkhurst, University of Toronto, are especially to be thanked for providing laboratory
and other facilities, and Dr Patricia Mather (Kott) for her helpful criticism. The work forms
part of a project carried out during tenure of Canadian National Research Council and
Australian Research Committee Grants.

LITERATURE CITED*

Aiyer, K. S. P., 1929. An account of the Oligochaeta of Travancore. Rec. Indian Mus. 31 : 13-76.

Bahl, K. N., 1926a. Pheretima (The common Indian earthworm). Indian Zoological Memoirs of Indian
Animal Types.

1926b. Enteronephric system in Woodwardia and the nephridia of Lampito dubius. Quart. J. tnicr. Sci.
70: 113.

1942. Part 1. Nephridia of the Octochaetinae. Quart. J. micr. Sci. 83: 423.

Baird, W., 1871. Megascolex antarctica, an earthworm from New Zealand. Proc. Linn. Soc. Lond. 11 : 96.

Beddard, F. E., 1888. Remarks upon a species of Coccidium infesting Perichaeta. Ann. Mag. nat. Hist. (6)
2: 434-9.

1891 . Abstract of some investigations into the structure of the Oligochaeta. Ann. Mag. nat. Hist. (6) 7 :
88-96.

1894. Some new or little known Oligochaeta. Proc. R. Phys. Soc. Edinb. 12 : 3 1—45.

1903. The earthworms of the Maidive and Laccadive Islands. In J. S. Gardiner (Ed.) ‘Fauna and
Geography of the Maidive and Laccadive Archipelagoes.' Vol. 1, pp. 374-5.

1905. On a new species of worm of the genus Pontodrilus from the shores of the Red Sea. Proc. zool.
Soc. Lond. 1905 : 558-63.

Benham, W. B., 1901. On some earthworms from the islands around New Zealand. Trans. N.Z. Inst. 33:
129-44.

1903. On some new species of some aquatic Oligochaeta from New Zealand. Proc. zool. Soc. Lond.
1903 : 202-32.

1906. Earthworms from Little Barrier Island. Trans. N.Z. Inst. 38: 248-56.

Bourne, A. G., 1894. On certain points in the development and anatomy of some earthworms. Quart. J.
Micr. Sci. 36 :

Cognetti de Martiis, L., 1910. Nuove specie dei generi Megacsolides e Pheretima. Ann. Mus. Stor. nat.
Giacomo Doria (3) 4 : 327-34.

Eisen, G., 1895. Pacific Coast Oligochaeta. I. Mem. Calif. Acad. Sci. 2: 63-122.

^References not cited here are given in Jamieson, 1971a and b.

MEGASCOLECOTD EARTHWORMS

101

Gates, G. E., 1929. A summary of the earthworm fauna of Burma with descriptions of fourteen new species.
Proc. U.S. Nat. Mas. 75 (10): 1-41 .

1940a. Indian earthworms. V1II-XI. Rec. Indian Mas. 42: 1 15-44.

1940b. Indian earthworms. XII. The genus Hoplochaetella. Rec. Indian Mus. 42: 199-252.

1941. On some earthworms from Ceylon. Spolia zeylan. Colombo 23 (1): 27-60.

1943. On some American and Oriental earthworms. Ohio J . Sci. 43 (3): 99-116.

1961. On some Burmese and Indian earthworms of the family Acanthodrilidae. Ann. Mag. nat. Hist.
(13)4: 417-29.

Grube, E., 1855. Beschreibungen neuer oder wenig bekannter Anneliden. Arch. Naturgesch. 21 (1): 81-136.

Hague, F. S., 1923. Studies on Sparaganophilus eiseni Smith. Trans. Amer. micr. Soc. 42: 1-42.

Iizuka, A., 1898. On a new species of littoral Oligochaeta ( Pontodrilus matsushimemsis). Annot. zool. jap. 2:
21-7.

Jamieson, B. G. M., 1970a. A review of the Australian earthworm genus Woodwardiella with descriptions
of two new genera (Megascolecidae : Oligochaeta). J. Zool., Lond. 162: 99-144.

1970b. Two new sympatric species of the earthworm genus Digaster (Megascolecidae : Oligochaeta)
from Queensland. Proc. roy. Soc. Qd 82 (3): 35-46.

1 971a. A review of the megascolecoid earthworm genera (Oligochaeta) of Australia. Part I — Reclassifica-
tion and checklist of the megascolecoid genera of the world. Proc. roy. Soc. Qd (In press).

1971b. A review of the megascolecoid earthworm genera (Oligochaeta) of Australia. Part II — The
subfamilies Ocnerodrilinae and Acanthodrilinae. Proc. roy. Soc. Qd (In press).

Macnab, J. A. and McKey-Fender, D., 1948. North American Plutellus and Megascolides with synonymical
notes (Annelida, Oligochaeta). Amer. Midi. Nat. 39 (1): 160-4.

Michaelsen, W., 1909a. The Oligochaeta of India, Nepaul, Ceylon, Burma and the Andaman Islands.
Mem. Indian Mus. 1 (3): 103-253.

1909b. On a new Megascolex from Ceylon. Spolia zeylan. 6: 96-101.

1910a. Die Oligochaten Fauna der vorderindisch-ceylonischen Region. Abh. naturw. Ver. Hamburg
19 (5): 1-108.

1910b. Oligochaten von verschiedenen Gebieten. Mitt, naturh. Mus. Hamb. 27: 47-169.

1924. Oligochaten von Neuseeland und den Auckland-Campbell-Inseln, nebst einigen anderen
Pacifischen Formen. Vidensk. Medd. dansk naturh. Foren. Kbh. 75: 197-240.

1928a. Clitellata. In Kukenthal and Krumbach, ‘Handbuch der Zoologie’, Vol. 2, part 8, pp. 1-112
(Berlin).

1928b. Die Oligochaten Borneos. Ark. Zool. 20 (3): 1-60.

1934. Oligochaeta from Sarawak. Quart. J. micr. Sci. 77: 1-48.

Nicholls, G. E. and Jackson, A. A., 1926. Some new species of Megascolex from South-Western Australia.
J. roy. Soc. W. Aust. 12 (16): 141-7.

Omodeo, P., 1955. Eudrilinae e Octochaetinae della Costa d’Avorio (Oligochaeta). Mem. Mus. civ. Stor.
nat. Verona. 4: 213-29.

Rosa, D., 1891. Die exotische Terricolen des k.k.naturhistorischen Hofmuseums. Ann. naturh. {Mus.)
Hofmus., Wien 6 : 379-406.

1898. On some new earthworms in the British Museum. Ann. Mag. nat. Hist. (7) 2: 277-90.

102

MEMOIRS OF THE QUEENSLAND MUSEUM

Schmarda, L. K., 1861. ‘Neue wirbellose Thiere gesammelt auf einer Reise um die Erde’, Vol. 1, part 2
(Leipzig).

Smith, F., 1937. New North American species of earthworms of the family Megascolecidae. Proc. US. Nat.
Mus. 84 (3009): 157-81.

Stephenson, J., 1922. Some earthworms from Kashmir, Bombay and other parts of India. Rec. Indian Mus.
24 : 427-43.

1923. Oligochaeta. In ‘The Fauna of British India’. (Taylor and Francis: London).

1925. Oligochaeta from various regions, including those collected by the Mount Everest Expedition,
1924. Proc. zool. Soc . Lond. 1925 : 879-907.

1931. Oligochaeta from Burma, Kenya and other parts of the world. Proc. zool. Soc. Lond. 1931 : 33-92.

Vidya Vati, 1947. The enteronephric system in species of Lampito and Megascolex. Rec. Indian Mus. 45 :
97-109.

Mem. Qd M us. (1971) 16 ( 1 ): 103-140

REVISION OF THE CALL10NYM1D FISHES REFERABLE TO
THE GENUS CALLIONYMUS FROM AUSTRALIAN WATERS

Clifford Ray Johnson*

Department of Zoology, University of Queensland

ABSTRACT

The genus Callionymus Linnaeus from Australian waters is revised. C. ocelli-
gena McCulloch is considered a synonym of C. calcaratus Macleay, C. limiceps
sublaevis McCulloch has been raised to specific level, C. macdonaldi Ogilby has
been redescribed, C. kaianus moretonensis has been described as a new subspecies,
and C. punctatus Langsdorff has been added to the known callionymids of
Australia.

The systematic treatment of callionymids in general has been poor due to the lack of
appreciation of their sexual dimorphism and the scarcity of specimens in museum collec-
tions. A "world wide revision is needed and regional revisions such as this may provide a
basis for more extensive treatment.

Lack of knowledge concerning sexual dimorphism has resulted in several species being
erected erroneously in Australian waters. Also many attempts, particularly by earlier
Australian workers, to subdivide the genus Callionymus has further complicated the situa-
tion, with almost every Australian species being typed as a new genus at one time or another.
These attempts have met with little success and are not generally accepted. As more taxo-
nomic work is done on this group, some Australian species may be synonymized with other
Indo-Pacific species. The possibility of additional species being added to the known Aus-
tralian callionymid fauna is ever present as exploratory sampling is lacking along much of
the coastline and few localities have been sampled deeper than 180 m.

McCulloch (1929) listed 14 species within the genus Callionymus from Australia. One
of these species has since been placed in the genus Synchiropus (Schultz et al., 1960). Mees
(1963) synonymised two species and added several others to the known callionymid fauna
of Western Australia.

* Present address: Department of Zoology, University of New England.

.104

MEMOIRS OF THE QUEENSLAND MUSEUM

In this work, it has been possible to synonymise two species, raise one subspecies to
the specific level with further information on biology and distribution, redescribe one
species, describe one new subspecies, and add two species to the callionymid fauna of
Australia. Corrections have been made of erroneous reports of species upon re-examination
of misidentified museum material.

Material examined in this study was collected from Moreton Bay with an 8 fathom
Siebenhauser otter trawl (1 mesh, measured stretched) operated aboard the R/V Wanderer
II, and at various locations offshore along the Queensland coast with standard commercial
prawning gear operated aboard several vessels, mostly the L. F. B. Bossanova and the
L. F. B. Jodi. Some material was also beach seined. Fathoms have been used rather than
meters when reporting depth distribution to be consistent with earlier records. Additional
material was examined from the collections of various museums. List of abbreviations of
museums and collections are as follows:

AM Australian Museum, Sydney

BM British Museum (Natural History), London

CAS California Academy of Sciences, San Francisco
QM Queensland Museum, Brisbane

SAM South Australian Museum, Adelaide

USNM U.S. National Museum of the Smithsonian Institution, Washington
P Western Australian Museum, Perth

KU Kyoto University, Maizuru, Japan

F Dept. Agriculture, Stock and Fisheries, Research and Survey Station,

Kanudi, Papua

FMNH Field Museum of Natural History, Chicago
SU Stanford University, Stanford

All lengths are standard length (SL) and all measurements are in mm. All measurements
are direct measurements. Most of the measurements used are self explanatory. The following
definitions have been used for some that are not self evident or characteristic of calliony-
mids: head length is measured from the tip of the snout to the gill opening; head width is
measured at the base of the preopercular spines; snout length is measured from the tip
of the snout to nostrils; postorbital length of head is measured from the rear edge of eye
to posterior edge of gill openings; length of caudal peduncle is measured from the base
of the last anal ray to the midbase of the caudal fin; body depth is greatest body depth.
Head width should be considered a crude measurement which is dependent upon the
condition of the branchial chamber at the time of preservation.

Key to Species and Subspecies of Callionymus found in Australian Waters

1 .

Preopercular spine stout, recurved, bent or angular with recurved tip and
spines along inner margin

2

CALLiONYMID FISHES FROM AUSTRALIAN WATERS

105

Preopercular spine slender (possibly stout), spear-like, straight or very slightly
curved with serrations or teeth along inner margin; basal antrorse spine

present 10

2(1). Antrorse spine present on base of preopercular spine 4

Basal antrorse spine absent 3

3 (2). DIV, 1\\ anal 6|; preopercular spine curved upwards, with 2 hooks

C. papilio

DIV, 8|; anal 1\\ preopercular spine curved upwards, with 2 hooks

C. calauropomus

DIV, 8 \ or 9^; anal 1\\ preopercular spine curved upwards, with 3 hooks
C. phasis

4 (2). DIV, 8^; anal 1\ \ preopercular spine with 3 hooks C. calliste

DIV, 9 1 (rarely 8| or 10|); anal 9\ (rarely 7$, 8^ or 10|); preopercular
spine with 2-5 hooks 5

5 (4). Preopercular spine with 2 hooks 6

Preopercular spine with 3-5 hooks 8

6 (5). Preopercular spine slightly curved upwards with a barbed tip; first dorsal

with a large black blotch between 2nd and 4th spines, first dorsal spine
filamentous C. kaianus moretonensis sub sp. nov.

Preopercular spine curved inwards 7

7 (6). Upper surface of cranium and supraorbital region rugose C. limiceps

Upper surface of cranium and supraorbital region smooth and covered by

integument C. sublaevis

8 (5). Raised erests and circular pits along each side of the occiput; first dorsal in

males grey, in females black with anterior spines white to grey

C. macdonaldi

No raised crests or circular pits along each side of the occiput; first dorsal
with dark blotch 9

9 (8). Preopercular spine broad; blotch on dorsal large, light in males and dark on

a white background in females C. calcaratus

Preopercular spine long, approximately the length of eye; dorsal blotch of
concentric shape in males, in females the first dorsal is dark grey to dusky
C. punctatus

106

MEMOIRS OF THE QUEENSLAND MUSEUM

10 (1). DIV, 8|; anal 1\\ preopercular spine more or less straight with 5-12 small

serrations 11

DIV, 9^; anal 9| or 8|; preopercular spine with 6-16 coarse spines or barb-
like serrations 12

1 1 (10). Snout slightly shorter than eye ; only 1 st pectoral ray simple C. rameus

Snout larger than eye ; 1 st and 2nd pectoral rays simple , C. goodladi

12 (10). First dorsal on line with gill openings; 1st and 2nd pectoral rays always

simple C. grossi

First dorsal not on line with gill openings; sometimes only 1st pectoral ray
simple 13

13 (12). Caudal short, does not equal body less head C. belcheri

Caudal elongate, equals body less head 14

14(13). Two bony bucklers, one on each side of occiput, with smooth ridges or
tubercles radiating from their centres; median area behind eyes covered
with smooth integument C. japonicus japonicus

Two bony bucklers, one on each side of occiput, with rugose ridges radiating
from their centres; median area behind eyes covered in bony rugosities
C. japonicus scaber

Callionymus japonicus japonicus Houttuyn

Callionymus japonicus Houttuyn, 1782, p. 311.

Callionymus reevesi Richardson, 1844, p. 60.

Callionymus longicaudatus Temminck and Schlegel, 1845, p. 151.

Callionymus Belcheri Bleeker, 1879, p. 85 [not C. belcheri Richardson, 1844]

Calliurichthys japonicus: Jordan and Fowler, 1903, pp. 942-3. McCulloch, 1929, p. 338.

Callionymus numeri Tanaka, 1917, p. 12.

Callionymus, Calliurichthys japonicus [$fc]: McCulloch, 1926, pp. 196-7.

[?] Callionymus ajfinis Ogilby, 1910b, p. 134 [not C. affinis Regan, 1908]

Callionymus affinis: McCulloch, 1929, p. 339.

Material Examined: 2 specimens, 106 and 175 mm SL (Research and Survey Kanudi, unregistered
and F01705 respectively), from waters N. of Cape York Peninsula off Yule T. and Bramble Cay, Gulf of
Papua.

Description: Dorsal IV, 9|; anal 8^; pectoral iil6ii or il6iii; ventral 1,5; caudal
ii6ii or i6iii; body compressed in front and elongate; snout pointed; preopercular spine
strong and straight with 7-16 barb-like serrations along inner margin; caudal elongate,

CALUONYMID FISHES FROM AUSTRALIAN WATERS

107

equals body less head; 1st or 2nd or both dorsal spines filamentous in males; occiptal
region with a pair of elevated crests (bony bucklers) separated by smooth integument of
head.

Head length 4.19 to 4.58; head width 5.60 to 6.25; length of pectoral fin 5.22 to
6.04; body depth 11 .00 to 11 . 14; all in standard length. Eye 3 .06 to 3 .74; snout 2 .87 to
2 .99; pectoral fin length 1 .24 to 1.32; all in head length. Eye 1 .02 to 1 .30 in snout.

Colour in Alcohol: As described by Jordan and Fowler (1903).

Sexual Dimorphism: Marked sexual dimorphism exists in colouration and fin shape.
In the male the anterior dorsal spines are produced into long filaments and the median
caudal rays are produced. Snout length is also greater in the male. All the dorsal spines are
short in the female. The median caudal rays are produced in the female but are usually
shorter than the body while those of the male are characteristically longer. The snout is
blunt in female specimens. Colour differences between sexes are quite evident with a deep
brown to black blotch on the chest and the branchiostegals greyish brown to black in the
male. The female undersurface is pure white and the chest blotch is absent.

Remarks: Since no specimen of C. affinis was available for examination, I can not be
sure that it would be placed under C.j. japonicus. McCulloch (1926) synonymized C. affinis
with C.j. japonicus from an examination of the only existing specimen (now appears to have
been lost) and since he described C.j . scaber it seems unlikely that it would be placed under
this subspecies. Other subspecies may be erected when an examination is completed on a
world wide basis.

Distribution: Previously from south of New Guinea, station 188 (Gunther, 1880)
and off Cape Moreton, SE. Queensland (Ogilby, 1910b; McCulloch, 1929).

Callionymus japonicus scaber McCulloch

Callionymus japonicus scaber McCulloch, 1926, p. 197.

Callionymus longicaudat us Waite, 1898, p. 60. McCulloch, 1923, p. 8.

Material Examined: 85 specimens, 91-193 mm SL, coastal south Queensland (20 miles SE. of Double
Island Point, 32-3 fm ; E. of Noosa, 41-46 . 5 fm ; and off Caloundra, 41-46 . 5 fm). One specimen, off Caloun-
dra, QM 18209.

Description: Dorsal IV, 9|; anal 8|; pectoral il6ii, iil6i, iii 1 6i, i 1 7i or iil7i; ventral I,
5; caudal i7ii; ridges or bucklers (elevated crests) are very rough and the area behind the
eyes is covered with similar bony rugosities; preopercular spine with 9-12 serrations along
inner margin; otherwise same as C. j. japonicus.

108

MEMOIRS OF THE QUEENSLAND MUSEUM

Head length 4.24 to 4.81 ; length of pectoral fin 5.00 to 5.77; head width 4.36 to
5.58; depth of body 8 . 47 to 1 3 . 30 ; all in standard length. Eye length 3 . 00 to 3.60; snout
2 . 73 to 3 . 46 ; pectoral fin l . 1 8 to 1 . 20 ; all in head length. Eye 1 . 00 to 1 . 1 1 in snout.

Colour in Alcohol: These specimens agree with the colour description of Jordan
and Fowler (1903) except that the following slight differences were noted. Remarks in
parentheses refer to Jordan and Fowler’s description. Light grey above (deep rich brown)
with numerous rounded spots of grey and brown (pale brown) margined with dark brown;
ventral surface white to yellowish (pure white), males have a dark brown to black (rich
brown) blotch on the chest, branchiostegals greyish in males and white in females.

Sexual Dimorphism: As for C. j. japonic us.

Remarks: Slight differences were noted in the proportions of the head length and of
the eye in snout from the material reported upon by Jordan and Fowler (1903) but these
were within the ranges given in de Beaufort and Chapman (1951) for C. j. japonic us. The
differences in head length could be due to the method of taking measurements as the criteria
for measurements are not mentioned by any of the above workers. Munro, 1967, and de
Beaufort and Chapman, 1951, both report 7 to 8 serrations along the inner margin of the
preopercular spine in C.j. japonicus. My material ( C.j . scaber) ranged from 9 to 12 (usually
10) serrations.

Distribution: Previously from Lord Howe I., off New South Wales (type locality)
(McCulloch, 1926), and now SE. Queensland.

Callionymus kaianus moretonensis subsp. nov.

(Figs. 1-5)

Holotype: Adult female, 158.3 mm SL, 7 miles E. of Cape Moreton, southeast Queensland (approx-
imately 27°02'S lat. 153°36'E long.) 68-72 fm, collected by L.F.B. Bossanova at night on 4 August 1969,
AM 115608.00.

Paratypes: Five, 95.2-149.0 mm SL, AM 115608-002-6; four, 130.9-164.9 mm SL, CAS 24764-67;
all collected with the holotype. Two, 30 miles E. of Mooloolaba, 60-70 fm, 12-14 August 1967, R. Elks,
QM 19156-7. One, southern Queensland, Department of Fisheries, QM 13428.

In addition, 894 specimens (74-162 mm SL) used in biological studies were recorded from southern
Queensland off shore (7 miles E. of Cape Moreton 65-80 fm; 18 miles E. of Caloundra, 56-60 fm; ENE. of
Noosa, 54-8 fm; 62-8 fm; E. of Mooloolaba, 59-67 fm). A few of these specimens will be deposited in the
Queensland Museum and the United States National Museum.

C ALLION YM ID FISHES FROM AUSTRALIAN WATERS

109

Description: Spinous dorsal fin IV; second dorsal elements 9| (last ray branched at
base); anal fin elements 9 \ (last ray branched at base); pectoral rays il7ii (19 specimens),
il8ii (20 specimens); ventral rays 1,5; caudal rays i7ii.

Measurements (mm) of holotype: Head length 42.4, greatest body depth 20.8, least
depth of caudal peduncle 6 . 9, snout 9.3, bony interorbital 1 . 0, eye 13.1, postorbital length

TABLE 1

Measurements in Percent of Standard Length or Head Length (*) of the Holotype
and Nine Paratypes of Callionymus kaianus moretonensis

Sex

Catalogue no.

$

AM

115608

-001

holo-

type

3

AM

115608

-002

3

AM

115608

-003

3

AM

115608

-004

$

AM

115608

-005

3

AM

115608

-006

3

CAS

24764

3

CAS

24765

9

CAS

24766

9

CAS

24767

Standard length(mm)

158.3

124.6

149.0

140.5

95.2

128.6

130.9

134.0

141.7

164.9

Head length

26.7

29.6

26.5

27.9

31.6

27.6

28.9

28.9

28.1

27.6

Greatest body depth

13.1

12.2

9.5

11.3

12.8

11.5

10.5

10.2

12.0

13.2

Least depth of

caudal peduncle

4.3

3.0

3.6

3.8

3.5

2.8

4.1

4.0

3.2

3.9

Snout*

21.9

17.3

17.2

17.8

17.2

16.0

18.2

15.2

15.7

18.4

Bony interorbital*

2.3

2.1

3.7

2 0

1.6

2.2

2.1

1.5

2.0

3.5

Eye*

30.8

30.3

35.4

35.8

35.2

34.2

32.1

33.5

32.3

29.1

Postorbital length

of head

10.9

11.8

10.8

11.5

13.9

10.3

11.2

10.5

11.2

12.6

Length of caudal

peduncle

16.8

21.5

18.2

18.6

17.8

18.4

19.4

20.9

18.4

18.2

Snout tip to origin

of first dorsal

26.8

26.9

25.9

27.3

28.8

26.3

26.7

26.1

26.9

25.6

Snout tip to anal

origin

42.5

43.5

45.8

45.7

47.4

45.9

46.0

45.4

48.2

44.2

Length of first

dorsal spine

25.9

22.9

24.0

20.7

31.5

22.6

19.9

21.1

21.0

22.4

Longest soft dorsal

ray

17.9

20.7

18.1

18.6

21.1

18.5

19.0

17.8

19.8

20.4

Longest pectoral ray

17.6

19.5

17.4

17.5

21.1

18.8

18.2

18.2

20.3

20.0

Longest pelvic ray

23.6

23.6

21.2

20.2

24.2

22.7

23.5

24.0

23.4

23.2

Longest caudal fin

ray

41.6

54.7

43.2

46.1

49.6

48.6

50.1

48.8

50.3

43.3

Tip of snout to rear

edge of maxillary*

28.8

22.7

25.0

26.2

21.9

23.5

25.0

20.7

21.0

24.5

Length of preoper-

cular spine

6.5

7.2

7.5

7.5

8.2

8.3

8.6

6.9

7.6

6.6

110

MEMOIRS OF THE QUEENSLAND MUSEUM

of head 17.3, length of caudal peduncle 26 . 6, snout tip to origin of first dorsal 42 . 5, snout
tip to anal origin 67.3, length of first dorsal spine 41.1, longest soft dorsal ray 28.4,
longest pectoral ray 27.9, longest pelvic ray 37 . 5, longest caudal fin ray 66.0, tip of snout
to rear edge of maxillary 11 .4, length of preopercular spine 10.3. Additional measurements
are given in table 1.

Body elongate, compressed in front with greatest depth near 2nd and 3rd spine of first
dorsal; body broader than deep tapering posteriorly. Head compressed, greatest depth
about one-half width at preopercular spine; snout rounded, compressed above, greatest
depth about one-half width ; eyes close together, directed upward, 29.1 to 35 .8% of head
length; mouth small, inferior, jaws unequal, maxillary reaching nostril; dentition villiform,
in approximately 8 bands upper jaw and 6 bands lower; preoperculum with a strong spine
with a slightly recurved barbed tip with 2 barbs near tip and a strong basal antrorse spine
(fig. 2). Gill openings small, slit-like, on upper surface of body approximately midway
between origin of first dorsal fin and preopercular spine base and as far apart as outer
margins of eyes.

Spinous dorsal inserted approximately midway, but nearer posterior margin of eye
than origin of soft dorsal, spines long, the first produced into a long filament, decreasing in
length posteriad; origin of soft dorsal just anterior to a vertical line through anal papilla;
anal originating under 3rd dorsal ray, lower than soft dorsal, the last ray branched and
produced; pectorals between origin of 3rd spinous dorsal and 4th dorsal ray, broad, with
median rays longest; pelvics large, broad and longer than pectorals, originating before the
gill openings and extending to about mid pectoral length and joined to base of pectoral by a
broad membrane; caudal long, graduated above and below the median rays which are
produced; caudal peduncle long and compressed, depth less than the eye.

Colour in Alcohol: Background colouration dorsal greyish; ventral yellowish white,
head and body marbled dorsally with brown markings, sides of body with 2 to 3 irregular
brown blotches; dark brown blotch at pectoral base; first dorsal dusky to greyish with a
large black irregular occellated spot between 2nd and 4th spine; second dorsal with numer-
ous brown spots regularly dispersed between the rays (2 on each membrane between 1st
and 3rd ray, 3 on rest of membrane except 1 or 2 between last branched ray). Anal blackish
distally, clear basally; caudal with blackish streak along ventral rays otherwise irregularly
marked with brown spots; pectorals clear; pelvics dusky to yellowish basally, upper rays
with brown bands. Life colouration not known.

Sexual Dimorphism: Little sexual dimorphism exists in this subspecies. No colour
differences could be detected between sexes. The median caudal rays are produced and are
longer in males. The first dorsal spine is slightly longer in males being produced in both
males and females (fig. 3). Considerable sexual dimorphism is present in the length of the
anal papilla as would be expected (fig. 4). The lack of sexual dimorphism in the other fins
is here demonstrated by the example of the last anal ray in fig. 5. Ochiai et al. (1955) reported

Preopercular spine of C. k. moretonensis. A, spine as it appears partly covered by integument; B, spine with integument removed.
Sexual dimorphism of the first dorsal spine in C. k. moretonensis. Dark circles males (n=102), open circles females (n=104).

112

MEMOIRS OF THE QUEENSLAND MUSEUM

greater sexual dimorphism in the length of the first dorsal spine in C. k. kaianus from
Japanese waters than was found in this subspecies.

Remarks: Juveniles of 80 to 100 mm have only two spots on each membrane of the
second dorsal and have the tip of the preopercular spine not barbed or only slightly barbed.
At around 110 to 112 mm the barbed tip becomes more pronounced. Ochiai et al. (1955)
reported the lack of barbed preopercular spine tips in young C. k. kaianus.

5-

4-

£

£

< 3-

_i

Cl

<

Q_

<

-2L

< 2 -

1 - •

o

Fig. 4: Sexual dimorphism of the anal papilla in C. k. moretonensis. Dark circles males (n=130), open
circles females (n=85).

CALLIONYMID FISHES FROM AUSTRALIAN WATERS

113

E

20

<

cc

<

<

t—
tO
<
I

IQ-

80 ' 100 ' 120 ' 140

STANDARD LENGTH mm

Fig. 5: Sexual dimorphism of the last anal ray in C. k. moretonensis . Dark circles males (n=81), open
circles females (n=79).

Dr Alwyne Wheeler kindly examined the type specimen of C. k. kaianus (BM 1879.5.
14.565) and sent me data and photographs for comparison with C. k. moretonensis , one
specimen of C. k. kaianus was examined from Japan (KU 23274).

C. kaianus moretonensis can be readily distinguished from C. k. kaianus by the differ-
ence in body markings ; regular pattern of spots on second dorsal versus irregular blotches
in C. k. kaianus ; the more anterior position of the lunate spot on spinous dorsal; the black
band always present on the distal portion of the anal fin ; and by a more slender preopercular
spine.

Etymology : The subspecific name moretonensis is taken from the type locality, Cape
Moreton.

Distribution : Known only from southern Queensland offshore.

Callionymus punctatus Langsdorff
(Fig. 6)

Callionymus punctatus Richardson, 1846, p. 210.

Callionymus richardsoni Bleeker, 1854, p. 414.

Callionymus curvicornis Gunther, 1861, p. 145.

Callionymus valenciennesi Schlegel, 1845, p. 153.

114

MEMOIRS OF THE QUEENSLAND MUSEUM

Material Examined: 432 specimens, 57-105 mm SL, Moreton Bay (Shark Spit, Moreton I., 16 fm;
1 mile E. of Otter Rock, near Redclifife, 3.5 fm; 4 miles E. of Redcliffe, 5 fm; 4 5 miles E. of Queens Beach,
5-6 fm ; 6 miles E. of Scarborough, 8-9 fm ; 1 . 5 to 2 miles E. of Bishop I., 2 . 5-3 fm ; E. of St Helena I ., 2 . 5-5
fm; E. of St Helena I. and Mud Islands, 4 fm; 7.5 miles W. of Tangalooma, Moreton 1., 5 fm) and Gulf
of Carpentaria 16°40'S, 139°42'E (one specimen, AM 115557-217). Museum material examined consisted
of 34 specimens: USNM 177165, 71076(12); CAS 24868(2); KU 22093-94; SU 7758(3), 7948(6), 7759(3),
7854(3); FMNH 44956; AM 115557-217.

Description: Dorsal IV, 9| (8^ or 10£ rarely); anal 8| (7|, 9|, or 10| rarely); pectoral
il3i2i (1 specimen), il4ii, il5i, or i!5ii; ventral 1,5; caudal i7ii, ii7ii or ii7iii; preopercular
spine nearly as large as eye with 3 or 4 spines besides recurved tip and an antrorse spine
at base; snout pointed, larger than eye; last soft dorsal and anal rays extend beyond caudal
base; dorsal spines not filamentous; caudal long with median rays longest; first dorsal in
male grey with irregular dusky blotches and a darker blotch (black in some specimens)
between 3rd and 4th spine; first dorsal in female dark grey to dusky.

Head length 3.58 to 4.00; greatest body depth 9 .00 to 10.66; postorbital length of
head 10.13 to 11 .40; length of caudal peduncle 5.79 to 6.50; snout tip to origin of first
dorsal 2 .90 to 3.14; snout tip to anal origin 1 .91 to 2 .00; longest soft dorsal ray 3 .94 to
5.58; longest pectoral ray 3 . 94 to 4 . 34; longest pelvic ray 4 04 to 4 . 79 ; longest caudal fin
ray 2 . 63 to 3 . 1 2 ; all in standard length. Least depth of caudal peduncle 4 . 80 to 5 . 48 ; snout
3.08 to 4.00; eye 3 .04 to 3.43; length of first dorsal spine 1 .53 to 2.44; tip of snout to
rear edge of maxillary 3 . 1 6 to 3 . 87 ; length of preopercular spine 3 . 40 to 4 . 1 4 ; all in head
length. Bony interorbital 8.75 to 12.43 in eye.

Colour in Alcohol: As described by Ochiai et ah (1955). Background colouration of
dorsal surface light brown, ventral surface yellowish to white; numerous light to dark
brown streaks along the side of the body in males, may be faint or absent in females ; first
dorsal in males grey with irregular dusky to black blotches and a darker blotch (black in
some specimens) between 3rd and 4th spines, fin edged in black; first dorsal in females
dusky to black with anterior spines grey in Australian material, differing from Japanese
specimens in lacking the occellated black blotch on the membrane between the 3rd and 4th

Fig. 6: Some examples of variation in right and left preopercular spines of Callionymus punctatus.

CALLIONYMID FISHES FROM AUSTRALIAN WATERS

115

spines; anal with a dark distal streak in males and white in females; caudal rays mottled
with dark brown spots; ventral dusky distally and basally lighter (much lighter in females);
upper pectoral rays spotted with brown, lower rays white.

Sexual Dimorphism: Besides the colour differences mentioned above, the length of the
last dorsal rays, anal papilla, and caudal fin rays are greater in males. Spinous dorsal spines
appear to be slightly larger in males. Some male specimens exhibit the female first dorsal
colour pattern of the described material from Japan with the occellated black blotch
between the 3rd and 4th spines. This has also been discussed by Ochiai et al. (1955) and
Jordan and Hubbs (1925).

Remarks : This is the first report of this species from Australian waters and it appears
to be very abundant wherever it occurs. Considerable variation occurs in the preopercular
spine in this species (fig. 6).

Distribution : Moreton Bay and Gulf of Carpentaria.

Callionymus phasis Gunther

Callionymus phasis Gunther, 1880, p. 28. Macleay, 1884, p. 35. Waite, 1904, p. 51. McCulloch, 1922,
p. 103; 1923, p. 9; 1926, p. 212; 1927, p. 77; 1929, p. 338; 1934, p. 77. Norman, 1937, p. 56.
Ochiai et al, 1955, p. 104-6. Mees, 1963, p. 98-9. Schultz et al. 1960, p. 403.

Callionymus apricus McCulloch, 1926, p. 209; 1929, p. 339. Waite, 1927, p. 231.

Yerutius apricus : Whitley, 1931, p. 115; 1948a, p. 27. Scott, 1962, p. 170.

Material Examined: One specimen, 47 mm SL, collected 1921, AM IA.431.

Description: Dorsal IY, 8^ or 9£; anal 1\\ pectoral il7i; ventral 1,5; caudal ii7ii; all
dorsal rays divided ; preopercular spine curved upwards at tip with 2 hooks along the inner
margin and no antrorse spine at base; eyes large, high above profile of head; inter-orbital
very narrow; origin of first dorsal only a little behind gill aperture.

Head length 3.14; pectoral fin length 4.27; head width 3.62; body depth 5.88; all
in standard length. Eye 3.00; pectoral length 1.37; snout 5.00; all in head length. Eye
0 .60 in snout.

Colour in Alcohol: All colour markings faded, background colouration grey.

Remarks: Ochiai et al. (1955) recorded C. phasis as new to Japanese waters, basing
this record on a 44 mm SL specimen (KU24847) which on examination proves to be

116

MEMOIRS OF THE QUEENSLAND MUSEUM

C. calliste and not C. phasis. Because this specimen was called C. phasis , juveniles of
C. phasis were thought to have simple dorsal rays, but since this misidentified specimen was
the only evidence for this, C. phasis should be considered to have divided dorsal rays until
proven otherwise.

Distribution: Known from Twofold Bay, New South Wales; South of Drana Range,
Gippsland, Victoria, 80 fm; Tasmania along eastern coast and the Great Australian Bight
south from Eucla (Mees, 1963; Australian Museum record).

Callionymus grossi Ogilby

Callionymus grossi Ogilby, 1910a, p. 43. McCulloch, 1929, p. 338. Schultz, 1960, p. 403. Palmer, 1962,
p. 548. Mees, 1963, p. 97. Marshall, 1965, p. 380. Johnson, 1970, p. 294.

Callionymus, Calliurichthys, grossi [sic] : McCulloch, 1923, p. 8; 1926, p. 195. McCulloch and Whitley,
1925, p. 173.

Callionymus Calliurichthys, nasutus [sic] McCulloch, 1926, p. 197.

Callionymus nasutus : Marshall, 1951, p. 5; 1965, p. 380.

Callionymus ( Calliurichthys ) nasutus [s7c] : Mees, 1959, p. 9.

Calliurichthys nasutus : Whitley, 1962, p. 226.

Material Examined: 142 specimens, 45-146 mm SL, from the Gulf of Carpentaria 16°52 , S, 139°39'E;
Bowen; Cockle Bay, Magnetic I.; south coastal Queensland (6-10 miles N. of Noosa, 20 fm; 10 miles N. of
Noosa, 21-2 fm; and E. of Caloundra, 15 fm); and Moreton Bay (1 mile W. of Shark Spit, MoretonL, 13-17
fm; off Sand Hills, Moreton I., 7.5-17 fm; 3 miles SW. of Tangalooma, Moreton I., 15 fm; off Lucinda
Bay, Moreton I., 12-17.5 fm; E. of Mud I., 4-5 fm; 2 miles E. of Mud I., 6-8 fm; 2.5 miles E. of the
southern tip of Mud I., 7-8 fm; 1 mile E. of SE. corner of St Helena I., 5 fm; ENE. of St Helena I., 5 fm;
4 miles E. of St Helena I., 5-6 fm). The following museum specimens were examined. Queensland Museum:
11579 (paratype, Bulwer, Moreton Bay), 17248 (Cockle Bay, Magnetic I,), 14239 (Bowen, N. Qld), 16855
(Mud L, Moreton Bay), Australian Museum: IB. 326 (Shark’s Bay), 115557-219.

Description: Dorsal IV, 9|; anal 8|; pectoral iil3i, iil3ii, iil3iii, ii 1 4i or iil4ii; ventral
1,5; caudal i7ii or ii7ii; preopercular spine straight with 9-16 serrations (antrorse teeth)
along inner margin and an antrorse spine at base; origin of first dorsal in line with gill
openings; first and second pectoral rays simple; first dorsal fins elongated in both sexes.

Head length 4 . 00 to 4 . 80 ; pectoral fin length 4 . 88 to 5 . 64 ; head width 5 . 00 to 5 . 58 ;
body depth 9.39 to 11 .18; all in standard length. Eye 3 12 to 3 .88; pectoral length 1 .08
to 1.41; snout 2.21 to 3 . 12; all in head length. Eye 1 . 00 to 1 .65 in snout.

Colour in Alcohol: Slight variations occur from the type material described by
Ogilby (1910a). Ogilby (191 0a) reported that a female ( ?) specimen was darker in colouration
whereas the background colouration in males and females is similar in my material. Slight

CALLIONYMID FISHES FROM AUSTRALIAN WATERS

117

differences do occur between the markings of males and females. In general the markings
on female specimens are not as distinct as in males — the spotting on the soft dorsal and
caudal is faint in females; females lack the blue spot on the membrane posterior to the
4th spine and some of the oblique crossbands are absent and others faint on the spinous
dorsal.

Sexual Dimorphism: In addition to colour differences, sexual dimorphism occurs in
snout length, anal papilla length and apparently in caudal fin length, all being longer in
males. Both sexes have elongated spinous dorsal fins.

Remarks: Mees (1963) reported one specimen having an anal count of 7^ and that
serrations on the preopercular spines of his material ranged from 10 to 18. Ogilby (1910a)
originally reported 7 to 9 antrorse spines present. Examination of one paratype revealed
9 antrorse spines. A small female specimen 45 mm in standard length had only 6 and one
female 71 mm had 7 serrations and like many callionymids the number of spines along the
inner margin of the preopercular spine is probably a function of size and growth. One 22 mm
specimen (AM IB. 326) collected by Mr G. P. Whitley from Shark’s Bay, Western Australia
and identified by him as C. calcaratus upon examination by me appears to be C. grossi. The
preopercular spine is straight and has only 3 antrorse teeth along the inner margin. It has
an elongated spinous dorsal with IV, 9^ and anal count of 8£. This specimen and the 45 mm
specimen mentioned above had none of the crossbanding on the first dorsal and lack spots
on the dorsal and caudal fins, the membranes being mostly clear with a few darker mottlings.
The body marking of the 45 mm specimen was characteristic of the adult colour pattern.
Crossbanding on the spinous dorsal was more distinct in the 71 mm specimen; a few spots
occurred on soft dorsal and caudal but fin membranes still had some clear areas.

Distribution: From north and south coastal Queensland (Dunk I., 70 miles SE. of
Cairns, 13 fm; off Townsville, 1 1 fm; near Lucinda, 9 fm (Johnson, 1970); Gulf of Carpen-
taria, 16°52'S, 139°39'E; Torres Straits; Lindeman I.; Point Denison (Aust. Mus. un-
published records, Paxton, pers. comm.); 13 miles SE. of Cape Capricorn (McCulloch,
1926, type locality of C. nasutus.)). In Western Australia from Shark Bay, Exmouth Gulf
(Mees, 1963) and the Monte Bello Islands (Palmer, 1962).

CaUionymus calauropomus Richardson
(Figs. 7-12)

CaUionymus calauropomus Richardson, 1844-48, p. 10, pi. 7, figs. 4, 5. Gunther, 1861, p. 147; 1880,
p. 28. Castelnau, 1873, p. 49. Macleay, 1881a, p. 627; 1881b, p. 262. Tenison- Woods, 1883, p. 19.
Lucas, 1890 (June), p. 29. McCoy, 1890, p. 333, pi. 192. Woodward, 1902, p. 271; 1903, p. 153.
Waite, 1904, p. 51; 1921, p. 142; 1923, p. 165. Stead, 1906, p. 209; Ogilby, 1910a, p. 48. McCulloch
and Waite, 1918, p. 48. Glauert, 1921, p. 46. McCulloch, 1922, p. 103; 1923, p. 12; 1926, p. 209;
1927, p. 77. McCulloch and Whitley, 1925, p. 173. Whitley, 1948a, p. 27. E. O. G. Scott, 1953, p.
157. Fowler, 1959, p. 493. T. D. Scott, 1962, p. 169. Mees, 1963, p. 95.

118

MEMOIRS OF THE QUEENSLAND MUSEUM

Callionymus achates De Vis, 1883, p. 620. Macleay, 1884, p. 35.
Callyonymus calauropomus: Castelnau, 1875, p. 21.

Foetorepus achates : Whitley, 1931, p. 323.

Synchiropus calauropomus : Schultz 1960, p. 405.

Material Examined: 227 specimens, 78-133 mm SL from coastal southern Queensland (8 miles N. of
Cape Moreton, 40 fm; E. of Caloundra, 41-42.5 fm). One specimen from Port Jackson, N.S.W. (QM
19927).

Description: Dorsal IV, 8|; anal 7|; pectoral il6i, il7i, il7ii or il8ii; ventral 1,5;
caudal ii7iii or iii7iii; all dorsal rays divided (first sometimes simple); preopercular spine
terminates in two hooks, bent upwards, no basal antrorse spine ; only first ray of pectoral
unbranched; origin of first dorsal nearly on line with gill openings.

Head 3.30 to 3.50; head width 3.67 to 5.35; length of pectoral fin 4.68 to 5.50;
body depth 6.32 to 7.67; all in standard length. Eye 3.38 to 4.26; snout 3.48 to 5.68;
pectoral fin 1 . 42 to 1.57; all in head length. Eye 0 . 60 to 1 . 1 9 in snout.

Colour in Alcohol : Brown to grey above, white to yellow below ; mottled with brown
spots from eye along lateral line to caudal fin in male, lacking in female ; a few spots radiate
out on caudal rays; anal, pelvic, and dorsal black to dusky and pectoral pale in males;
dorsal, pectoral, pelvics, caudal and anal dusky to white in females; in males the dorsal fin
and pelvics mottled with dark spots with second dorsal streaked distally.

Sexual Dimorphism: Males are usually larger than females (fig. 7). The anal papilla
is also longer in males. The greatest amount of sexual dimorphism appears in the median
caudal filaments (fig. 8) in this species with lesser amounts occurring in the length of the
last dorsal and anal rays (figs. 9, 10). Little sexual dimorphism appears in the rest of the
dorsal spines or rays (figs. 11, 12). Colour differences were discussed above.

7

5

>

u

o

a:

5

70 75 80 85 90 95 tOO 105 T10 115 120 125 130 135

STANDARD LENGTH mm

Fig. 7: Length-frequency of Callionymus calauropomus (males, n=97: females, n=67).

CALLIONYMID FISHES FROM AUSTRALIAN WATERS

119

9

O

m

• „ IM<I» • •

o • m • • 0 *

o •« «d OO

CC0©OOCD0C

«b*e°8£

60 ' 80 ' 100 ' 12o"

STANDARD LENGTH mm

60 80 100 120 140

STANDARD LENGTH mm

Fig. 8 : Sexual dimorphism of the caudal filaments in C. calauropotnus . Dark circles males (n^lOO), open
circles females (n=69).

Fig. 9: Sexual dimorphism of the last dorsal ray in C. calauropomus. Dark circles males (n=100), open
circles females (m=69).

Fig. 10: Sexual dimorphism of the last anal ray in C. calauropomus. Dark circles males (n=100), open
circles females (n=69).

Fig. 1 1 : Sexual dimorphism of the first dorsal spine in C. calauropomus. Dark circles males (n=100), open
circles females (n=69).

Fig. 12: Sexual dimorphism of the first dorsal ray in C. calauropomus. Dark circles males (n=100), open
circles females (n=69).

120

MEMOIRS OF THE QUEENSLAND MUSEUM

Remarks: Many attempts have been made to subdivide the genus Callionymus, the
most recent being that of Schultz et al. (1960) in which he placed this species under the genus
Synchiropus. Mees (1963) attacked the validity of the genus Synchiropus avidly and I concur
with his arguments. The genus Synchiropus is based on too few characters e.g. branched or
simple dorsal rays (except in young) and the presence or absence of a basal antrorse spine
on the preopercular spine. Schultz placed C. rameus in the genus Callionymus without
comment even though it has branched dorsal rays and an antrorse spine (i.e. his generic
limits break down with C. rameus ). I have to agree with Mees (1963) that S. calauropomus
should be restored to the genus Callionymus. Type material of the genus Synchiropus must
be reexamined to determine the validity of this genus. I do not wish to comment on
Synchoripus microps Gunther or S. splendidus Herre, which can also be found in Australian
waters, at this time.

Distribution: From coastal southern Queensland (De Vis, 1883; Macleay, 1884;
McCulloch and Whitley, 1925; McCulloch, 1929; Whitley, 1931; Mees, 1963). Known
from New South Wales: Port Jackson (Macleay, 1881a; McCulloch, 1922, 1923, 1927;
McCulloch, 1934); Eden (Mees, 1963); New South Wales (Tenison-Woods, 1883; Waite,
1904; McCulloch, 1923, 1929; T. D. Scott, 1962; Mees, 1963). From Victoria: Bass Straits,
38 fm (Gunther, 1880); E. of Flinders I., Bass Strait (McCulloch, 1926); Port Phillip
(Macleay, 1881a); Victoria coast (McCulloch, 1923, 1929; T. D. Scott, 1962; Mees, 1963).
From South Australia : off Marsden Point, Kangaroo I. (McCulloch, 1 926) ; South Australia
coastline (McCulloch and Waite, 1918; Waite, 1921, 1923; McCulloch, 1923, 1929; T. D.
Scott, 1962; Mees, 1963). From Western Australia: Michaelmas I., King George Sound;
off Limestone I., King George Sound; off Bald I., between Albany and the Archipelago of
the Recherche (Mees, 1963); Doubtful Island Bay, Southwestern Australia (McCulloch,
1926); coastal western Australia (Gunther, 1861; Macleay, 1881a; Woodward in Fraser,
1903; Glauert, 1921; McCulloch, 1923, 1929; Whitley, 1948a; T. D. Scott, 1962). From
Tasmania: Bridgport (Mees, 1963); Tamar River, at Launceston (E. O. G. Scott, 1953);
coastal Tasmania (McCulloch, 1929; T. D. Scott, 1962; Mees, 1963). Also known from
New Ireland, New Guinea Region (McCulloch, 1929).

Callionymus belcheri Richardson
(Figs. 13-17)

Callionymus belcheri Richardson, 1844, p. 62, pi. 37, figs. 1-2. [Not C. belcheri Bleeker, 1879].

McCulloch, 1926, p. 199; 1929, p. 339. Schultz, 1960, p. 403.

Material Examined: 520 specimens, 34-107 mm SL, from North Queensland (Dunk L, 13 fm; off
Townsville, 1 1 fm; off Cairns, 12 fm), coastal south Queensland (E. of Caloundra, 15 fm), and Moreton Bay
(2 miles E. of Redcliffe, 2 fm; 6 miles E. of Scarborough, 7-8 fm; 1 . 5-2 miles E. of Bishop T., 2 5 fm; E. of
Mud I., 3-7.5 fm; E. of St Helena I., 4-7 fm; 7.5 miles W. of Tangalooma, Moreton 1., 5 fm; 3 miles
SW. of Tangalooma, 15 fm; 4 miles N. of Shark Spit, Moreton I., 14-17 fm; 5 miles SW. of Shark Spit,
ll-16fm; W. of SharkSpit, ll-16fm; W. of Sand Hills, Moreton I., 12-13.5 fm; Lucinda Bay, Moreton!.,
6-18 fm; Brisbane River mouth, 2 fm). Three specimens, Kinikini B., New Guinea, F0969; 1 specimen,
Sepik area, New Guinea, F01386. Three specimens, Gulf of Carpentaria, 16°44'S, 139°33'E, AM 115557-220

CALLION YM ID FISHES FROM AUSTRALIAN WATERS

121

13

«3 • M

••

OO

Oo
CP OO

mtg>Q (Sumo
assno o
cnfi8®<a*Doooo
oe&0<bg>o

o o a® o

60 80 100
STANDARD LENGTH mm

14

< 1 5-

'•«32>0

m #^ckd o &

• o*

o#fP O C0C 8
oojfetf So o

STANDARD LENGTH mm

70 80 90 100

STANDARD LENGTH mm

Fig. 13: Sexual dimorphism of the first dorsal spine in C. belcheri. Dark circles males (n=100), open
circles females (n— 100).

Fig. 14: Sexual dimorphism of the last dorsal ray in C. belcheri. Dark circles males (n=100), open circles
females (n=100).

Fig. 15: Sexual dimorphism of the caudal fin in C. belcheri. Dark circles males (n— 100), open circles
females (n=100).

Fig. 16: Sexual dimorphism of the anal papilla in C. belcheri. Dark circles males (n— 100), open circles
females (n=100).

Fig. 17: Some examples of variation in right and left preopercular spines of C, belcheri.

122

MEMOIRS OF THE QUEENSLAND MUSEUM

Description: Dorsal IV, anal 9£; pectoral iMiii (mostly), il5ii, ilSiii, i 1 6ii or
i 1 6iii ; ventral 1,5 ; caudal i7ii ; preopercular spine very large, with nearly straight tips directed
slightly outwards, 6-7 large spines which increase in size anteriorly; head large, depressed
and heart shaped; snout pointed; no membrane behind the 4th spine in spinous dorsal.

Head length 3.26 to 3.73; head width 3.04 to 3.28; length of pectoral fin 3 .76 to
4.25; body depth 10 . 63 to 1 2 . 1 8 ; all in standard length. Eye 3 . 42 to 3.71; snout 2.71 to
3.25; pectoral fin length 1 . 13 to 1 .20; all in head length. Eye 1 .07 to 1 .31 in snout.

Colour in Alcohol: Yellowish brown above; chin and ventral yellowish white to
white; two characteristic oval dark brown spots, one present on each side immediately below
the lateral line, the first spot under tip of pectoral, the second above the 4th or 5th anal ray;
first dorsal with a network of grey lines in males, dusky to black in females ; 7-8 longitudinal
grey lines on soft dorsal in males, these lines faint or non-existent in females; anal in male
with a black submarginal stripe, anal white in female; oblique grey lines on membrane of
upper caudal rays in males (faint in females) and irregular dots below, obscure brown bars
on caudal rays in both sexes; pectorals and ventrals with grey to brownish dots. In pro-
longed preservation this species becomes very pale with few definite markings apparent;
even the two characteristic spots on the sides disappear.

Sexual Dimorphism: A great deal of sexual dimorphism is apparent in this species.
Both the spines and last ray of the dorsal fins are considerably larger in males (figs. 13, 14).
The caudal fin is greatly lengthened in the male (fig. 15) as is the anal papilla (fig. 16). In
contrast only slight sexual dimorphism occurs in the anal rays and this is most apparent in
the last anal ray. Differences in colour between the sexes are discussed above.

Remarks: McCulloch (1926) mentioned that C. belcheri was regarded as the young of
C. longicaudatus Temminck and Schlegel (= C. j. japonicus Houttuyn) but he failed to
mention that this was C. belcheri Bleeker 1879, a different species entirely to C. belcheri
Richardson 1 844. McCulloch did recognize that C. belcheri Richardson was not synonymous
to C. j. japonicus or C. longicaudatus. Considerable variation is evident in the preopercular
spines (fig. 17).

Distribution: From north and south coastal Queensland, Gulf of Carpentaria, New
Guinea and Pacific Ocean.

Callionymus macdonaldi Ogilby
(Fig. 18)

Callionymus macdonaldi Ogilby, 1911, p. 56. McCulloch, 1923, p. 9; 1926, p. 205; 1929, p. 338.
McCulloch and Whitley, 1925, p. 175. Schultz, 1960, p. 403. Marshall, 1965, p. 382.

CALL! ON YM ID FISHES FROM AUSTRALIAN WATERS

123

Material Examined: 8 specimens, 59-71 mm SL, from coastal Queensland (Townsville on beach;
mouth of Moon Creek, on beach, Fraser I., 2-3 ft; Cribb I., Moreton Bay, seined 3 ft; E. of St Helena I.
and Mud I., Moreton Bay, 4 ft). Twelve specimens from Queensland Museum: Townsville, on beach,
19928; Moreton Bay (holotype) 12473; Woody Pt, 18392-402. One specimen, USNM 177164.

Description: Dorsal IV, 8|(1 specimen), 9| (17 spec.), 10| (2 spec.); anal (2 spec.),
9 \ (16 spec.), 10| (2 spec.); pectoral i2il3ii (1 spec.). il5ii, il6i or i 1 7ii (usually); ventral 1,5;
caudal i6iii or i7ii (usually).

Measurements (mm) of holotype (96.8 mm in SL) are as follows: Head length 25 .3,
greatest body depth 11.4, least depth of caudal peduncle 4.5, snout 7.1, bony interorbital
1 .0, eye 6.7, postorbital length of head 9 .8, length of cuadal peduncle 12.9, snout tip to
origin of first dorsal 32.6, snout tip to anal origin 55.3, length of first dorsal spine 9.3,
longest soft dorsal ray 14.7, longest pectoral ray 22.8, longest pelvic ray 24.4, longest
caudal fin ray 26 . 7, tip of snout to rear edge of maxillary 8 . 2, length of preopercular spine
7.6. Additional measurements are given in table 2.

TABLE 2

Measurements in Percent of Standard Length (SL) or Head Length (*) of the
Holotype and other Specimens Of C. macdonaldi Ogilby

Sex

Cat. No.

¥

QM

12473

(holo-

type)

<?

QM

18402

¥

QM

18400

¥

QM

18401

¥

QM

18399

<?

QM

18393

<J ¥ ¥

personal collection
not registered

SL (mm)

96 8

62.7

66.6

59.4

63.9 |

54.6

71.7

71.1

69.5

Head length

26 2

31.0

28.2

29.8

28.9

25.9

28.8

29.0 j

29.9

Greatest body depth
Least depth of caudal

11.8

12.0 !

11.9

12.9 1

13.7

10.8

13.4

13.8

13.4

peduncle

4.7

6.2

5.1

5.4

5.6

5.0

5.2

5.8

5.8

Snout* . . . . . . I

28.1

31.4 1

25.7

28.8

28.0

24.1

24.3

23.3

26.0

Bony interorbital*

3.9

3.9

4 3

4.5

3.3 i

4.3

4.4

4.9

3.8

Eye*

26.4

24 2

25 2 ,

26.6

22.0 |

31.1 |

27.2 1

29.1

25.0

Postorbital length of head ..

10.1

13.1

11.6 1

12.5

12.4

11.0

11.6 !

11.8 !

12.2

Length of caudal peduncle . .
Snout tip to origin of first

13.4

14.4

14.6

12.5

14.1

12.8 j

14.5

14.2

13.8

dorsal

33 6

36.5

35.2

39.8

38.6

34.4

35.4

37.8

37.2

Snout tip to anal origin . .

57 2

58.4

51.1

57.4

57.8 1

53.4

55.8

57.8

56.6

Length of first dorsal spine*

j 36.8

41.2

34.2

| 20.9

28.0

37.6

j 34.5

31.6

26.4

Longest soft dorsal ray

15.2

18.5

14.9

13.3

15.9

18.3

16.5

14.2

14.7

Longest pectoral ray

23.6

24.6

26 0

26.1

24.5

24.2

24.1

26.6

| 25.6

Longest pelvic ray

25.2

26.2

25.4

24.4

20.7

25.3

1 24.3

23.9

I 23.5

Longest caudal fin ray
Tip of snout to rear edge of

27.6

25.6

25.8

24.4

j 26.7

j 22.9

27.5

26.1

27.3

maxillary*

32.4

27.8

28.9

| 22.6

' 27.5

l 25.5

26.7

j 23.3

27.9

Length of preopercular spine*

! 30.1

! 28.9

28.9

31.6

! 28.6

i 36.9

30.1

31.5

28.4

124

MEMOIRS OF THE QUEENSLAND MUSEUM

Body depressed, greatest depth near origin of first dorsal; broader than deep, tapering
posteriorly. Head large, depressed, greatest depth slightly less than width at base of pre-
opercular spine ; eyes close together, directed upward, interorbital space narrow, 5 . 9 to 7 . 4
in eye. Snout rounded, mouth small, jaws unequal, maxillary reaching back to below
nostril; dentition villiform, in wide bands with approximately 8 to 12 rows of teeth in both
jaws. Head rugose dorsally, bony ridges radiating from raised crests; shallow pits, mostly
circular, profuse along bony crests the longest of which are on each side of the occiput.
Gill openings on upper surface, midway between origin of spinous dorsal and posterior
edge of orbits; space between gill openings less than space between outer margin of orbits.
Preopercular spine with recurved tip slightly curved upward distally, with 5 large teeth
(2nd and 3rd largest) along the inner margin ending in a recurved tip and with a basal
antrorse spine.

Width of caudal peduncle greater than its depth at midlength; lateral line curved
downward from head over pectoral fin, raised slightly at end of pectoral and extending
posteriad slightly above midline of body to caudal base. Anal papilla present and shows
sexual dimorphism. Ogilby (1911) and McCulloch (1926) both stated that the anal papilla
was absent in this species ; one is present, although small, on the holotype.

Spinous dorsal inserted approximately midway between 2nd dorsal insertion and
posterior margin of orbit, short, not reaching second dorsal when depressed, 3rd spine
slightly longer than rest. Last ray of second dorsal produced reaching hypural joint; all
rays of dorsal simple except last which is branched at base. Anal lower than soft dorsal, all
rays simple except last which is branched at base; last anal ray produced, reaching hypural;
anal origin on vertical line between 3rd and 4th soft dorsal rays. First pectoral ray simple;
upper margin of pectoral incised, median rays longest reaching to about the 4th soft dorsal
ray. Ventral rays coarsely branched with 5th longest, reaching vent; broad membrane con-
nects 5th ventral ray to base of pectoral ; caudal rounded in both sexes.

Colour in Alcohol: Background colouration highly variable, specimens collected
from beach sands almost colourless to light grey to greyish white; from muddy environments
darker in colour, grey to light brown ; dorsal surface closely covered with white, irregularly

18

Fig. 18 : Some examples of variation in right and left preopercular spines of C. macdonaldi.

CALLIONYM1D FISHES FROM AUSTRALIAN WATERS

125

shaped spots surrounded by an intricate pattern of fine brown lines ; a few larger light to
dark brown spots present; ventral surface white to yellowish; first dorsal of male grey,
dark distally, white basally, in female black with anterior spines white to grey; second dorsal
white to grey interspersed with dark spots, dusky posteriad ; anal in males with rays white
and membrane dusky, in females entire anal white; ventral fins irregularly spotted with a
few faint to distinct brown bands present depending upon habitat occupied by specimen
(e.g. sand, faint; mud, distinct bands); caudal white to grey with 4-6 irregularly dusky
vertical bars.

Sexual Dimorphism: In addition to colour differences, the only marked sexual di-
morphism is that of the anal papilla length. Anal papilla length measured in males 41 to 71
mm ranged from 0 . 76 to 2 . 32 mm (mean 1 . 36 mm) (n = 9) while in females from 38 to 96
mm this ranged from 0.24 to 0.84 mm (mean 0.50) (n = 9). Slight sexual dimorphism may
occur in the length of the first dorsal spine and that of the longest soft dorsal, but not enough
specimens have been available for examination to accurately determine this.

Remarks: This species has been redescribed here to correct omissions and incorrect
information given in the original description.

McCulloch (1926) stated that this species was closest to C. calcaratus. In my estimation
C. macdonaldi is closest to C. marleyi Regan from which it differs in caudal ray counts and
colouration (but colouration can be highly variable in both of these species). Specimens of
C. marleyi examined by Smith (1963) appeared to show less variation in dorsal and anal
counts than does C. macdonaldi, C. marleyi has 8| anal rays while C. macdonaldi can have
from 8 1 to 10£ (usually 9|). Most specimens of C. marleyi have soft dorsal counts of 9\,
but one specimen from Mozambique had 10| (Smith, 1963). The preopercular spines are
very similar in both species ; C. macdonaldi has 3 to 5 teeth along the inner margin depending
upon the size of the fish and counts can vary on two sides (fig. 18) while C. marleyi can
have 3 to 6 teeth and only 6 in large adults (the recurved tip has been counted along with
the teeth on the inner margin in Smith’s description) and the counts can vary on both sides.
I have not counted the recurved tip in my description, but if I did C. macdonaldi would
also have 6 teeth in large adults.

Distribution: From coastal Queensland and New South Wales (McCulloch, 1929;
Marshall, 1965).

Callionymus calcaratus Macleay
(Figs. 19-22)

Callionymus calcaratus Macleay, 1881a, p. 628; 1881b, p. 263; Tenison-Woods, 1883, p. 19. Ogilby,
1885, p. 121. McCulloch, 1922, p. 103; 1923, p. 10, pi. 3, fig. 2; 1926, p. 204; 1927, p. 77; 1929,
p. 338; 1934, p. 77. T. D. Scott, 1962, p. 168. Mees, 1963, p. 96. Schultz, 1960, p. 403. Marshall,
1965. p. 381.

126

MEMOIRS OF THE QUEENSLAND MUSEUM

Callionymus ocelligena McCulloch, 1926, p. 207; 1929, p. 339. Schultz, 1960, p. 403. Johnson, 1969,

p. 208.

Callionymus curvicornis : Ogilby, 1886, p. 37. Stead, 1901, p. 476. Waite, 1904, p. 51. Stead, 1906, p. 208.

Callionymus reevesi : Ramsay and Ogilby, 1886, (1887?), p. 942. Waite, 1904, p. 51.

Repomucenus calcaratus: Whitley, 1931, p. 323; 1948a, p. 27; 1948b, p. 275.

Repomucenus sp. nov.: Whitley, 1945, p. 42.

Material Examined: 561 specimens, 104-165 mm SL, from coastal Queensland (S. of Double Island
Point, 22-8 fm; 6-10 miles N. of Noosa, 20 fm; 10 miles N. of Noosa, 21-2 fm; E. of Mooloolaba, 19-19.5
fm; SE. of Mooloolaba, 19-23 fm; Mooloolaba to Caloundra, 19-22.5 fm; E. of Caloundra, 18 fm; 8 miles
N. of Cape Moreton, Moreton I., 40 fm; E. of Cape Moreton, 55-9 fm; off Lucinda Bay, Moreton I., 18 fm;
6 miles E. of Scarborough, 8-9 fm) New South Wales (Tweed Heads (CSIRO collection, Cronulla); Wallis
Lake; Princess Royal Harbour (Aust. Mus. records); Twofold Bay (Cronulla collection)). Six specimens,
Queensland Museum, 14739 (Wide Bay), 14740 (Port Jackson), 12129 (Moreton Bay), 12140 (Moreton Bay),
13427 (southern Queensland), 19263 (Gneering Shoal). Four specimens, Australian Museum IB. 358,
I A. 7858, 17239 (2 spec.), TA.4189.

Description: Dorsal IV, anal 9\\ pectoral iilbii, il7i, il7ii, il7iii, i 1 8i or il8ii
(usually il7ii or il7iii); ventral 1,5; caudal ii7ii or ii7iii; preopercular spine broad with 3 or
4 teeth besides recurved tip and basal antrorse spine; only last dorsal ray divided; first
dorsal origin well behind gill openings; usually only first pectoral ray simple, sometimes
first two are simple in juveniles; first dorsal short, with a large, incomplete dark ring
between 3rd and 4th spine (not always present) in males and a black blotch against a white
background in females; conspicuous grey ocelli with yellow margins between mouth and
preopercular spine, preorbital region and operculum in males.

Head 3.44 to 3.78; head width 3.88 to 4.74; length of pectoral fin 4.04 to 4.56;
body depth 8 . 34 to 1 1 . 1 9 ; all in standard length. Eye 3 . 64 to 4 . 1 2 ; snout 3 . 1 4 to 4 . 34 ;
pectoral fin length 1 .07 to 1 .30; all in head length. Eye 0.92 to 1.24 in snout.

Colour in Alcohol: Will be discussed under the remarks section.

Sexual diMORPHiSM : Generally males are slightly larger than females but this is not
nearly as marked in such species as C. sublaevis or C. calauropomus. The second dorsal fin
shows marked sexual dimorphism with the rays being longer in males (fig. 19). The first
dorsal spine (fig. 20), the caudal rays (fig. 21), and the last anal ray are longer in males.
The anal papilla is much longer in males than in females. Marked differences occur in the
colour markings between sexes and will be discussed in the following section.

Remarks: In my examination of the species of dragonets from Queensland it became
apparent that C. calcaratus Macleay and C. ocelligena McCulloch were strikingly similar
in counts, measurements and appearance except for colour markings. C. ocelligena was
described from a single male specimen in which the collection data had been lost (McCulloch

1st DORSAL SPINE LAST DORSAL RAY

CALLIONYM1D FISHES FROM AUSTRALIAN WATERS

127

19

STANDARD LENGTH mm

100 120 130 140

STANDARD LENGTH mm

28-

20

24-

■

20 '

18 -

16-
14-
12- O

10-

* * •

• • » •

• • • CD O

• CD* O OCO
o o 4D • OO <o*DO

• OD O GOcSD o

• O3D0DO O <H£00

• O CDO CDo C#

C3DOO OO 0D

(DO O CO O

"ioo 1 120 ' 130 ' ire"

STANDARD LENGTH mm

Fig. 19: Sexual dimorphism of the last dorsal ray in C. calcaratus. Dark circles males (n— 90), open circles
females (n— 95).

Fig. 20: Sexual dimorphism of the first dorsal spine in C. calcaratus. Dark circles males (n=90), open
circles females (n — 95).

Fig. 21 : Sexual dimorphism of the caudal rays in C. calcaratus. Dark circles males (n— 90), open circles
females (n=95).

Fig. 22: Examples of variation in right and left preopercular spines of C. calcaratus.

128

MEMOIRS OF THE QUEENSLAND MUSEUM

1926). The type specimen was deposited in the Australian Museum (McCulloch, 1929) but
has since disappeared. McCulloch (1926) stated that C. calcar at us had the closest affinity
to C. ocelligena and that C. ocelligena was readily distinguishable from C. calcaratus by its
colour markings (large brown ocelli between mouth and preopercular spine, preorbital
region and operculum; large black, white-edged spot on the operculum beneath the pre-
opercular spine), longer caudal fin and having the first dorsal spine longer than the second.
All these features are ones that can be affected by sexual dimorphism in this group.

So far over 500 specimens of C. calcaratus and C. ocelligena have been collected and
examined along the Queensland coast. In all cases these two species were found to co-occur
and all the C. calcaratus collected have been females while all the C. ocelligena collected
have been males. The brain pattern is identical in both whereas great variability occurs at
the interspecific level within the genus Callionymus thus examined (Johnson, 1971). Gonadal
development and other aspects of their biology coincide. Material from the Queensland
Museum showed all C. calcaratus to be females and C. ocelligena (only one specimen) to
be male. Fin ray counts for 16 females (C. calcaratus) were D IV, 9J; anal 9|; pectoral
iil6ii, il7i, il7ii, il7iii, il8i or il8ii; ventral 1,5; caudal ii7ii or ii7iii. Counts for 17 males
(C. ocelligena ) were identical except for less variation in pectoral counts (il7ii, il7iii or il8ii).

From all the material at hand it appears that C. ocelligena and C. calcaratus are
synonymous and that C. ocelligena is the male and C. calcaratus is the female of the same
species, the name of which should be C. calcaratus.

The colour of the female C. calcaratus fits that described by McCulloch (1923). The
colour of the male C. calcaratus (C. ocelligena ) preserved in alcohol from Queensland is as
follows: Background colouration yellowish brown, dorsal surface covered with indistinct
small grey and orange spots with dark margins; ventral surface white; conspicuous grey
ocelli with yellow margins between mouth and preopercular spine, preorbital region and
operculum; along the sides of the body are dark blotches below the lateral line (not present
in some specimens) ; bluish grey to black, white-edged spot on operculum beneath preoper-
cular spine; first dorsal with a large, incomplete dark ring between 3rd and 4th spine (not
present in some specimens), but instead fin mottled with dark brown and yellow spots
similar to those on back; second dorsal mottled with dark brown and yellow spots with
yellow margins ; anal fin bluish grey to grey ; caudal fin with irregular yellow and dark brown
dots with yellow margins, with or without a grey stripe on lower rays.

The preopercular spine is highly variable in this species (fig. 22). The counts may vary
on two sides and the number of spines appears to be a function of growth. A 20 mm
specimen, doubtfully referred to this species, had 3 teeth with the third just forming, and
one 33 mm specimen had only 2 teeth along the inner margin. This 33 mm specimen also
had some interesting variation in fin counts with the dorsal IV, 8|, and anal 1\. Not much
variation is encountered in the counts on adults.

CALLIONYMID FISHES FROM AUSTRALIAN WATERS

129

Distribution: Known from coastal Queensland (mouth of Wide Bay (McCulloch,
1926); Port Douglas, near mouth of Mary River; South Head, Mary River; Tangalooma
Point, Moreton Bay; off Point Lookout, Stradbroke I. (Australian Museum records, from
Mees, 1963)), New South Wales (northern New South Wales (McCulloch, 1926); apparently
not uncommon at Port Jackson (type locality) (Stead, 1906)), South Australia (Port Lincoln
and off York Peninsula (Mees, 1963)), and Western Australia (Cape Jaubert; Wallal,
5 miles offshore, 5 fm; Hampton Harbour; Dampier Archipelago; Exmouth Gulf; Hout-
man’s Abrolhos; Fremantle; Useless Inlet and off Kok’s I., Shark Bay (Mees, 1963;
Whitley, 1948a; McCulloch, 1926)).

Callionymus limiceps Ogilby

Callionymus limiceps Ogilby, 1908, p. 35. McCulloch, 1923, p. 9; 1926, p. 203; 1929, p. 340. McCulloch
and Whitley, 1925, p. 173. Whitley, 1929, p. 115, figs. 3, 4. Schultz, 1960, p. 403. Mees, 1963, p. 98
(C. limiceps mentioned; but only C. 1. sublaevis (C. sublaevis ) known in Western Australia).
Marshall, 1965, p. 380-1.

Callionymus limiceps var. typica McCulloch, 1926, pp. 195, 203.

Velesionymus limiceps : Whitley, 1934, suppl. no. 418a.

Material Examined: 162 specimens, 86-159 mm SL, coastal Queensland (10 miles N. of Noosa, 21-2
fm; E. of Mooloolaba, 19-22.5 fm; between Caloundra and Mooloolaba, 19-19.5 fm; E. of Caloundra,
18 fm), Moreton Bay (3 miles SW. of Tangalooma, Moreton I., 14.5-18 fm; off Shark Spit, Moreton I.,
15 fm; W. of Sand Hills, Moreton I., 10-13 fm; W. of Lucinda Bay, Moreton I., 16-18 fm; 1 mile N. of
Cowan Cowan, Moreton I., 2 fm; off Mud I., 5 fm; 1 mile E. of SE. corner of St Helena I., 5 fm). Fifteen
specimens, Queensland Museum (14038; 1487, 3 spec.; 19492-9; 13340-1; 13360).

Description: Dorsal IV, 9|; anal 9^; pectoral il6i, il6ii, il6iii, il7i or il7ii; ventral
1,5; caudal i7ii or ii7ii ; preopercular spine with a recurved tip and one antrorse tooth on
inner margin and an antrorse spine near the base; only first pectoral ray simple; males with
elongated first dorsal rays and a small black spot covering almost the entire fin; occiput
and supraorbital ridges rugose.

Head length 3.87 to 4 . 1 9 ; pectoral fin length 4.71 to 5.60; head width 3.77 to 4 . 08 ;
body depth 1 0 . 44 to 11 . 30 ; all in standard length. Eye 3 . 80 to 4 . 22 ; snout 2 . 24 to 2 . 82 ;
pectoral fin length 1 . 19 to 1.41; all in length of head. Eye 1 .41 to 1 .70 in snout.

Colour in Alcohol: Consistent with the descriptions by Ogilby (1908) and McCulloch
(1923).

Sexual Dimorphism: Great sexual dimorphism occurs in the length of the first dorsal
spines and anal papilla. Minor size differences between sexes are also apparent. Minor
colour differences occur as in the markings on the first dorsal fins and anal rays.

130

MEMOIRS OF THE QUEENSLAND MUSEUM

Distribution: Coastal Queensland, previously from between Hervey Bay and Port
Denison, 13-26 fm (McCulloch, 1923).

Callionymus sublaevis McCulloch
(Figs. 23-6)

Callionymus limiceps var. sublaevis McCulloch, 1926, p. 204; 1929, p. 340. Mees, 1963, p. 98. Johnson,
1970, p.294.

Material Examined: 1322 specimens, 51-134 mm SL, coastal Queensland (6 miles E. of Mackay, 13
fm; E. of Caloundra, 15 fm) and Moreton Bay (1.5-2 miles E. of Bishop I., 2.5-3 fm; W. of Green Island,
2-6 fm; between St Helena I. and Green I., 4-4. 5 fm; \ to 1 mile E. of St Helena I., 5-6 fm; 2.5 miles E.
of Mud I., 5-10 fm; f mile E. of Mud I., 5-7.5 fm; 2 miles E. of Redcliffe, 2 fm; 6 miles E. of Scarborough,
8-9 fm; W. of Sand Hills, Moreton I., 7-17 fm; 3 miles SW. of Tangalooma, Moreton I., 14.5-18 fm;
4 miles N. of Shark Spit, Moreton I., 14-17 fm; 5 miles SW. of Shark Spit, 1 1-16 fm; W. of Shark Spit,
11-16 fm; W. of Lucinda Bay, Moreton I., 7-18 fm). Six specimens, Australian Museum, 115557-218,
Gulf of Carpentaria, 16 C 37'S, 140°43'E.

Description: Dorsal IV, 9|; anal 9£; pectoral il6ii, il6iii or il7i (usually il7i); ventral
1,5; caudal ii6ii, ii7ii or ii7iii (usually ii7iii); similar to C. limiceps in most aspects except in
minor colour differences, smaller size and having the occiput and supraorbital ridges almost
entirely smooth and covered by skin ; preopercular spine with 1 to 2 teeth along inner margin
besides recurved tip, basal antrorse spine present.

Head 3.60 to 3.92; head width 3.48 to 4.40; length of pectoral fin 4.62 to 4.81;
body depth 9 . 80 to 1 1 . 36 ; all in standard length. Eye 3 . 66 to 4 . 14 ; snout 2 . 89 to 3 . 1 1 ;
pectoral fin 1 . 19 to 1 . 32; all in head length. Eye 1 .20 to 1 .43 in snout.

Colour in Alcohol: Brown above, mottled with white spots; first dorsal in male
dusky and lacks the black spot between 2nd and 4th spine as in C. limiceps ; first dorsal in
female black; filamentous spines with grey annuli; second dorsal with grey and white
streaks; caudal with many white spots between rays; anal fin in male with broad dusky to
black margin, narrow in female; ventrals black in male, dusky in females; pectorals pale;
5-6 brown spots along side of body with brown lines vertically in interspaces. In males
74 mm and smaller, the ventral fins and anal fins are similar to female. In a 92 mm male,
the ventral fins are partially black and edged with white.

Sexual Dimorphism : Differences occur in body size with females usually being smaller.
The first dorsal spines are greatly elongated in males (fig. 23). Fig. 24 shows the differences
in length of the caudal fin rays between sexes. Anal papilla length is greater in males (fig. 25).
Colour differences are discussed above.

CALLIONYMID FISHES FROM AUSTRALIAN WATERS

131

Remarks: McCulloch (1926) regarded C. /. sublaevis as an exceptional variation of
C. limiceps. Mees (1963) did not believe C. limiceps and C. /. sublaevis to be different species
or subspecies as they were reported to co-occur along the Queensland coast. I believe them
to be distinct at the specific level. Although they do co-occur, it is the exception rather than

Fig. 23: Sexual dimorphism of the first dorsal spine in C. sublaevis. Dark circles males (n— 100), open
circles females (n— 100).

Fig. 24: Sexual dimorphism of the caudal fin in C. sublaevis. Dark circles males (n — 1 00), open circles
females (n -100).

Fig. 25: Sexual dimorphism of the anal papilla in C. sublaevis. Dark circles males (n- 100), open circles
females (n 100).

Fig. 26: Examples of variation in right and left preopercular spines of C. sublaevis

132

MEMOIRS OF THE QUEENSLAND MUSEUM

the rule. I have found C. limiceps usually outside Moreton Bay at depths from 18 to 22 fm
while C. sublaevis is most common inside Moreton Bay at depths from 2 to 18 fm. Usually
distinct populations exist in either C. sublaevis or C. limiceps . No more than two C. limiceps
have ever been encountered in any trawl with C. sublaevis in Moreton Bay and no C.
sublaevis has ever been taken by me with C. limiceps in the hundred of trawls made outside
the bay. Both are biologically distinct and juveniles can be found in both. The brain patterns
are distinctly different in C. limiceps and C. sublaevis (Johnson, 1971).

Much variation exists in the shape of preopercular spines and counts can vary on
two sides (fig. 26).

Distribution: 13 miles SE. of Cape Capricorn, 13 fm; 7-10 miles NW. of Hummocky
I., near Cape Capricorn, 14-16 fm (holotype) (McCulloch, 1926). Coastal Queensland from
off Townsville, 11 fm, south to Moreton Bay (Johnson, 1970). In Western Australia from
Dampier Archipelago, Exmouth Gulf and Shark Bay (Mees, 1963).

Callionymus goodladi Whitley

Calliurichthys goodlacli Whitley, 1944, p. 270; 1948a, p. 27.
Callionymus goodladi: Mees, 1963, p. 97.

Material Examined : 7 specimens, 1 10-160 mm SL, Western Australian Museum (P4369, 4 specimens;
P5037; P4540 ; P5442).

Description: Dorsal IV, 8^; anal 7|; pectoral iil5ii, iil5iii or iil6i; ventral 1,5; caudal
ii7ii or ii7iii; preopercular spine straight with 8 to 12 small serrations along inner margin
and a large antrorse spine at base; snout large and prominent; origin of first dorsal nearly
on line with gill openings; first and second pectoral rays simple.

Head length 3.07 to 3.54; head width 3.84 to 4.12; length of pectoral fin 4.55 to
5.12; body depth 8 .78 to 10.00; all in standard length. Eye 4.42 to 5.71; snout 2 . 16 to
2.57; pectoral fin length 1 . 29 to 1.56; all in head length. Eye 1 . 77 to 2 . 42 in snout.

Colour in Alcohol: As described by Whitley (1944) except that in some specimens
the head is brownish grey above and the chin and brachiostegals are white or brown with
a bluish tinge.

Sexual Dimorphism: Pronounced sexual dimorphism is not apparent in this species
although some differences in fin length may exist.

CALLIONYMID FISHES FROM AUSTRALIAN WATERS

133

Distribution: Known only from Western Australia, from Cheyne Beach and near
Michaelmas I., King George Sound, Cockburn Sound, Shark Bay (entrance to South
Passage, exact locality not known) and Exmouth Gulf (Mees, 1963).

Callionymus calliste Jordan and Fowler

Callionymus calliste Jordan and Fowler, 1903, p. 957. Marshall, 1965, p. 380.

Callionymus hudsoni Fowler, 1941, p. 8.

Callionymus distethommat us Fowler, 1941, p. 18.

Material Examined: Two specimens, 29 and 31 mm SL, Australian Museum, IA.4463, IA.4630
Low Isles, Queensland.

Description: Dorsal IV, 8^; anal 1\\ pectoral il3ii or il3iii; ventral 1,5; caudal ii7ii
or ii7iii ; preopercular spine with 3 hooks along upper margin and a short basal antrorse
spine; lateral line simple and not connected by a branch on top of caudal peduncle; gill
openings approximately midway between origin of first dorsal fin and posterior edge of eyes.

Head length 3 . 44 to 3.62; length of pectoral 3 . 20 to 3 . 62 ; head width 3 . 64 to 4 . 14 ;
depth of body 6 . 34 to 7 . 25 ; all in standard length. Eye 2 . 67 to 2 . 90 ; pectoral length 0 . 93
to 1.00; snout 3.81 to 4 . 10 ; all in length of head. Eye 0 . 70 to 0 . 7 1 in snout.

Colour in Alcohol: Follows the description of Schultz et al. (1960).

Distribution: Within Australia, known only from Low Isles, Queensland.

Callionymus rameus McCulloch

Callionymus, Calliurichthys rameus [j/c] McCulloch, 1926, p. 201.

Callionymus rameus: McCulloch, 1929, p. 339. Schultz, 1960, p. 403. Mees, 1963, p. 99, Marshall,
1965, p. 381.

Orbonymus rameus: Whitley, 1947, p. 150.

Callionymus ( Calliurichthys ) rameus: Mees, 1959, p. 9; Palmer, 1962, p. 548.

Material Examined : 30 specimens, 84-135 mm SL, southern Queensland (20 miles SE. of Double Island
Point, 32-3 fm; 10 miles N. of Noosa,21-2 fm; 6-10 miles N. of Noosa, 20 fm; off Caloundra, 19-19.5 fm;
E. of Noosa, 41-46.5 fm. Two specimens, Queensland Museum, 14012 (paratype, SE. of Cape Capricorn,
13339 (S. coast of Queensland). One specimen, Australian Museum, 115557-286 (Gulf of Carpentaria
16°30'S, 140 °43'E).

134

MEMOIRS OF THE QUEENSLAND MUSEUM

Description: Dorsal IV, 8|; anal7|; pectoral il5ii, il6ii, il6iii, il7ii, or il7iii; ventral
1,5; caudal i7ii or ii7ii; all dorsal rays divided, first ray sometimes simple; preopercular
spine with 6 to 11 serrations along inner margin, antrorse spine on base; snout slightly
shorter than eye ; origin of first dorsal on a line with gill openings.

Head length 4.04 to 5.10; length of pectoral fin 3 .38 to 4 .21 ; head width 4.09 to
4.62; depth of body 5 . 79 to 7 . 00 ; all in standard length. Eye 2 . 24 to 3 . 34 ; pectoral length
0 . 84 to 1.00; snout 3 . 06 to 4 . 28 ; all in head length. Eye 0 . 59 to 0 . 94 in snout.

Colour in Alcohol : Slight variations were noted from the description by McCulloch
(1926); the 5 to 6 crossbands on the dorsal surface are quite distinct in some specimens, the
irregular bars and spots on ventral rays are absent or very faint, and the darker part of the
anal fin has a number of darker spots and bars in most specimens.

Life Colouration : The life colouration of this species is described for the first time,
based upon fresh specimens from the Mackay area.

In males the background colouration is light grey dorsally, white ventrally with red
blotches and streaks along sides; yellow reticulated pattern edged in red on chin and
branchiostegals; head with red blotch between eye and maxillary, 2 faint red bars on oper-
culum, 2 red blotches on lower jaw, upper jaw pink; dorsal surface with 5 to 6 dark brown
to grey bars edged with red and mottled with small blue spots; membranes of first dorsal
fin dark yellow or grey with vertical yellow bars with white spots and streaks edged with
black, spots basal, streaks distal; second dorsal light grey with horizontal yellow and
white streaks on posterior half, a few white spots along basal edge of fin and brown to grey
blotches dispersed throughout fin. Caudal fin with 2 wide, vertical, dark grey bars, distinct
dorsally, faint ventrally, merging into a black blotch on basal 3 rays, several blue streaks
on black blotch, caudal membrane with yellow between horizontal white streaks, rays pink
becoming more distinct anteriorly; anal fin black distally, yellowish grey basally with blue
horizontal broken streaks edged with black ; pectoral fin with membrane clear, rays with
minute white and red spots; ventral fin light grey with 2 darker grey wide patches distally
and basally across rays, irregular white spots or yellow background basally, red blotches
on rays.

In females the background is lighter, ventral surface with a wide indistinct pink patch
beginning at 2nd anal ray extending to 6th anal ray; head as in males, reticulated pattern
on chin and branchiostegals red instead of yellow; first dorsal fin darker with less yellow,
bars with white spots, no streaks; second dorsal with white streaks posteriorly, no yellow
streaks; caudal fin whitish yellow, vertical black bars more distinct and fuse into basal
black blotch with several blue spots, white horizontal broken streaks present, fin pinker
basally; anal fin with blue spots instead of streaks; pectoral and ventral fins as in males.

CALLIONYMLD FISHES FROM AUSTRALIAN WATERS

135

Sexual Dimorphism: Besides the colour differences discussed above, the first dorsal
fin, caudal fin and anal papilla are longer in males.

Distribution: Previously from Queensland (Gulf of Carpentaria (Australian Museum
record); 6 miles E. of Mackay, 13 fm; SE. of Cape Capricorn, 13 fm (type locality); 25 miles
SE. of Double Island Point, 33 fm; 4-20 miles NE. of Gloucester Head, 19-35 fm (McCul-
loch, 1926)), Western Australia (Shark Bay (exact location not known); 40 miles S. of
Carnarvon (Mees, 1963), and off the Monte Bello Islands (Palmer, 1962)).

Callionymus papilio Gunther

Callionymus papilio Gunther, 1864, p. 197. Lucas, 1890, p. 29. Waite, 1904, p. 51. McCulloch, 1922, p.
103; 1923, p. 13; 1927, p. 77; 1929, p. 338; 1934, p. 77. Lord, 1923, p. 69; 1927, p. 87. Lord and
H. H. Scott, 1924, pp. 12, 78. E. O. G. Scott, 1953, p. 157. T. D. Scott, 1962, p. 168. Mees, 1963,
p. 98.

Callionymus ocellifer Castelnau, 1873, p. 49.

Callionymus lateralis Macleay, 1881a, p. 628; 1881b, p. 263. Johnston, 1891, p. 33.

Callionymus macleayi Ogilby, 1886, p. 37 — nomen novum for Callionymus lateralis Macleay, nee
Callionymus lateralis Richardson.

Callionymus Papilio: Macleay, 1881a, p. 627; 1881b, p. 262.

Callionymus latealis: Tenison-Woods, 1883, p. 19.

Foetorepus papilio: Whitley, 1931, p. 323; 1945, p. 42; 1948a, p. 27.

Material Examined: 2 specimens. South Australian Museum, F3129, Cape Elizabeth, York Peninsula,
South Australia. Four specimens, Australian Museum, 115731-007, B9753, 15004, Point Peron, 30 miles S.
of Perth, 32°16'S, 115°40'E.

Description: Dorsal IV, 7^; anal 6£; pectoral il2iiii, il3ii, ilSiii or il5iiii; ventral 1,5;
caudal i7ii, i7iii or ii7ii; all dorsal rays divided at tips except first; preopercular spine
resembles that described for C. calauropomus with one single curved hook besides tip which
is curved upward and no basal antrorse spine present; a small species.

Head length 3.41 to 4.00; length of pectoral fin 4.10 to 4.66; head width 3.66 to
4.31 ; body depth 5.70 to 7 .00; all in standard length. Eye 3 .00 to 3 .86; pectoral length
I . 20 to 1 . 3 1 ; snout 3 . 08 to 4 . 24. Eye 0 . 9 1 to 1 . 08 in snout.

Colour in Alcohol: Background colouration light brown to dark grey with lighter
blotches dorsally, these blotches arranged symmetrically and similar in males and females;
ventral surface silvery anteriorly becoming brownish posteriorly with silvery white spots
on branchiostegals and chin; head same as body with a distinct white patch below eye in
both sexes; males with thin white vertical lines and a few faint white spots on sides of body

136

MEMOIRS OF THE QUEENSLAND MUSEUM

below lateral line, thin vertical lines absent in females and replaced by numerous white spots
below lateral line; first dorsal with black spot on distal portion of membrane between 1st
and 2nd spine (also present in juveniles), spines black at tips in adults, 3 irregular horizontal
bands of thin white stripes, markings not as distinct in females. Second dorsal in males
with 3 horizontal bands of thin white stripes, edged in black on a brown background, in
females, membranes clear and mottled with faint white and brown blotches (juveniles lack
the black edges of horizontal white bands) ; pectoral fin clear in both sexes ; anal fin in males
brown with distal black stripe, clear in females ; caudal fin with upper rays clear and lower
rays with white and brown spots on membranes, caudal rays in females entirely mottled
with white and brown and membranes clear; ventral fin mottled with brown and white
blotches, clear in juveniles.

Sexual Dimorphism: Besides the colour differences noted above, the only difference
apparent is that males have slightly longer dorsal spines.

Remarks: According to McCulloch (1923) this species can have 8| dorsal rays, but
all the material I examined had 1\.

Distribution: Known from Port Jackson, New South Wales, southward to Victoria
(McCulloch, 1923); from Tasmania (Lord, 1923; McCulloch, 1923, 1929; E. O. G. Scott,
1953; T. D. Scott, 1962).

ACKNOWLEDGMENTS

Many persons played an important role in making this study possible. From the Uni-
versity of Queensland, I wish to thank L. Wale (Captain of the R/V Wanderer II), A. Lewis,
B. Smith, M. Johnston, R. Bradbury and A. Jones for assistance in obtaining specimens.
N. Smales (L. F. B. Bossanova) and I. McDonald (L. F. B. Jodi ) supplied much of the off-
shore material and K. Boxwell (L. F. B. Seabelle ) and A. Johnson also helped in this respect.

A. Hinton, L. W. Filewood and P. Kailola, Research and Survey Station, kindly loaned
me New Guinean material.

Museum material was secured through the efforts of Drs V. G. Springer, U.S. National
Museum, W. Freihofer, Stanford University, L. P. Woods and K. Liem, Field Museum of
Natural History, J. R. Paxton, Australian Museum, R. J. McKay, Western Australian
Museum, W. M. Eschmeyer, California Academy of Sciences, A. Ochiai, Kochi University,
T. Iwai, Kyoto University, C. J. M. Glover, South Australian Museum, and Alwyne
Wheeler, British Museum.

CALLION YM ID FISHES FROM AUSTRALIAN WATERS

137

I wish to thank my wife, Colleen for preparing the figures and for her patience and
encouragement throughout the study.

Professor J. M. Thomson and Dr P. Dwyer, University of Queensland kindly offered
advice concerning the manuscript. The research was supported by AUC and URG research
grants to the University of Queensland.

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during the years 1839 to 1843’. Vol. 2, pp. i-viii + 1-39, pis. 1-60. (E. W. Janson: London).

1846. Report on the ichthyology of the seas of China and Japan. Rep. Brit. Ass. 18: 187-320.

Schlegel, H., 1845. Pisces In P. F. von Siebold ‘Fauna Japonica’ (J. Muller and Sons: Amsterdam).

Schultz, L. P., Chapman, W. M., Lachner, E. A. and Woods, C. P., 1960. Fishes of the Marshall and
Marianas Islands. U.S. Nat. Mus. Bull. 202 (2): 1-438.

Scott, E. O. G., 1953. Observations on some Tasmanian fishes, part 6. Pap. Proc. Roy. Soc. Tas. 87 : 141-64.

Scott, T. D., 1962. ‘The marine and freshwater fishes of South Australia.’ (Govt. Printer: Adelaide).

Smith, J. L. B., 1963. Fishes of the families Draconettidae and Callionymidae from the Red Sea and the
Western Indian Ocean. Rhodes University Ichthy. Bull. 28 ; 547-64.

Stead, D. G., 1901. In Notes and Exhibits. Ptoc. Linn. Soc. N.S.W. 25: 476.

1906. ‘Fishes of Australia: a popular and systematic guide to the study of the wealth within our waters.’
(William Brooke and Co. : Sydney).

Tanaka, S., 1917. Eleven new species of fishes from Japan. Zool. Mag . 29 (339): 7-12 (in Japanese).

Temmink, J. C. and Schlegel, H., 1843-50. Pisces. In P. F. von Siebold, ‘Fauna Japonica’. pp. 1-324, pis.
1-143. (J. Muller and Sons: Amsterdam).

Tenison- Woods, J. E., 1883. ‘Fish and fisheries of New South Wales (1882)’. (Thomas Richards: Sydney).

Waite, E. R., 1898. Report of the fishes. In ‘Sea Fisheries Report M.M.C.S. “Thetis”.’ pp. 23-62, pis. 1-12,
1 map.

1904. Synopsis of the fishes of New South Wales. Mem. N.S.W. Nat. Cl. 2: 1-59.

1921. Catalogue of the fishes of South Australia. Rec. S. Aust. Mus. 2: 1-199.

1923. ‘The fishes of South Australia. Handbooks Flora and Fauna S. Australia’. (Govt. Printer:
Adelaide).

1927. Supplement to the Catalogue of Fishes of South Australia. Rec. S. Aust. Mus. 3: 221-34, pi. 13.

140

MEMOIRS OF THE QUEENSLAND MUSEUM

Whitley, G. P., 1929. Studies in Ichthyology No. 3. Rec. Aust. Mus. 17: 101-43, pis. 30-4.

1931. New names for Australian fishes. Aust. Zool. 6: 310-34.

1934. Supplement to the check-list of the fishes of New South Wales. In A. R. McCulloch ‘The Fishes
and Fish-like animals of New South Wales.’ 3rd ed. (Royal Zoological Society of New South Wales :
Sydney).

1944. New sharks and fishes from Western Australia. Aust. Zoot, 10 : 252-73.

1945. New sharks and fishes from Western Australia. Part 2. Aust. Zool. 11 : 1-42.

1947. New sharks and fishes from Western Australia, Part 3. Aust. Zool. 11 : 129-50, pi. 11.

1948a. A list of the fishes of Western Australia. W. Aust. Fish. Dept., Fish. Bull. 2: 1-35.

1948b. New sharks and fishes from Western Australia, Part 4. Aust. Zool. 11 : 259-76, pis. 24-5.

1962. ‘Marine Fishes of Australia.’ Vol. 2, pp. 148-287. (Jacaranda Press: Brisbane).

Woodward, B. H., 1902. In M. Frazer, ‘Notes on the natural history, etc. of Western Australia’. In Western
Australian Year-Book for 1900-1901. (Govt. Printer: Perth).

1903. In M. Fraser, ‘Notes on the natural history, etc. of Western Australia’, pp. i-vii, 1-250, pis. 1-36.
(Govt. Printer : Perth).

Mem. Qd Mus. (1971) 16(1): 141-143, pi. 4.

NEW RECORDS OF TUBE-NOSED BIRDS (ORDER
PROCELLARIIFORMES) FROM QUEENSLAND

John L. McKean

CSIRO, Division of Wildlife Research
and

D. P. Vernon
Queensland Museum

The number of species of the order Procellariiformes recorded from Queensland is
small compared to the number known from other States. For Queensland Lavery (1969)
has listed 25 species whereas McGill (1960) records 36 species from New South Wales and
Wheeler (1967) 33 species from Victoria. Additional records have been since added to the
lists from New South Wales and Victoria and this contribution now documents several
new occurrences of Procellariiformes from Queensland.

In view of the wide-ranging nature of many of the Procellariiformes and the proximity
of Queensland to the Central Pacific and Tasman regions, the small number of species
recorded is probably a reflection of the number of interested bird workers. Undoubtedly
many species are yet to be detected in Queensland waters.

SPECIES ACCOUNTS

Pterodroma rostrata rostrata (Peale) (PI. 4) Tahiti Petrel

A specimen in good condition was picked up dead by David H. Barry on the Eastern
Beach of Fraser Island, Qld, on 12 May 1969. The skin prepared by Mr Barry has been
lodged in the Queensland Museum [Q.M. 0111657 Measurements in millimetres are:
exposed culmen 38.7, tarsus 48.2, wing 309.0, and tail 124.0.

Two subspecies of Pterodroma rostrata are known. One, P. r. becki, is known only
from the region of the Solomons Islands and Bismarck Archipelago. The specimen under
discussion is referable to the larger, nominate subspecies which breeds on islands in the

142

MEMOIRS OF THE QUEENSLAND MUSEUM

New Caledonia area, Society and Marquesa Groups during the winter months and ranges
the equatorial Pacific. P. rostrata has not been previously recorded from Australia, however
a live specimen was captured off Port Moresby, Papua, recently (W. Filewood, pers. comm.).

Pterodroma melanopus (Gmelin) Brown-headed Petrel

A specimen in poor condition was found at Eastern Beach, Fraser Island, by Miss
M. Hawken on 8 May 1967. The head has been made up as a study specimen [Q.M. Ol 1004].
The determination was made on the basis of bill-shape and facial markings. The exposed
culmen of the specimen measured 34.5 mm.

Pterodroma melanopus breeds on Lord Howe Island during the winter months. It
formerly bred on Norfolk Island. Odd beach-washed birds have been reported from New
South Wales and New Zealand. During the summer months the species apparently ranges
widely into the North Pacific.

Pterodroma macroptera gouldi (Hutton) Grey-faced Petrel

A male specimen in good condition was found dead at Caloundra, Qld, by R. Elks on
3 February 1967. The specimen was prepared into a study skin [Q.M. 01 1005]. Measure-
ments in millimetres are: exposed culmen 36.4, tarsus 40.7, wing 302.0, tail 132.0. The
specimen was determined to race on the basis of measurements and face colouration.

P. m. gouldi breeds on islands and headlands in the New Zealand region during the
summer months and ranges widely throughout the Pacific. It is not surprising that the
Queensland specimen is of the New Zealand breeding form, this subspecies being recorded
occasionally along the New South Wales coast (Hindwood, 1957). P. m. albani breeds on
the islands off the southwest coast of Western Australia, and has been recorded on a few
occasions from Victoria. The nominate subspecies breeds in the sub-antarctic islands and
ranges through the Atlantic.

Pelagodroma marina dulciae Mathews White-faced Storm Petrel

A live specimen found at Eagle Farm, Brisbane, was presented to the Queensland
Museum by R. Shortis on 8 August 1967. The specimen, an adult female, died shortly
afterwards and was prepared as a study skin [Q.M. Ol 1172]. Measurements in millimetres
are: exposed culmen 16.7, tarsus 41 .8, wing 145.0, and tail 74.0.

Not unexpectedly, it is an example of the Australian breeding race which breeds as
close to Queensland as the Broughton Islands. The specimen was determined to race using
the characters given by Oliver (1955). This specimen would appear to be the first recorded
from Queensland.

PROCELLAR1IFORMES FROM QUEENSLAND

143

Pachyptila salvini salvini (Mathews) Medium-billed Prion

A male specimen in good condition was found at Oxley, Brisbane, on 15 July 1954,
and presented to the Queensland Museum by the Royal Queensland Society for the Preven-
tion of Cruelty. This specimen was mounted but has since been relaxed to a cabinet skin
[Q.M. 05412]. Measurements in millimetres are: exposed culmen length 29.4, width 15.4,
tarsus 32.5, wing 188.0 and tail 96.0.

Two races have been suggested, the larger P. s. salvini breeding on Marion Island in
the southwest Indian Ocean, and the smaller P. 5. crozeti , breeding on the Crozet Islands
which are approximately 1000 miles to the east. Falla (1940) records both subspecies for
New Zealand seas, with P. s. salvini in greater numbers. The generally larger measurements
of Q.M. 05412 indicate it is probably of the nominate race.

According to McGill (1960) good numbers of P. salvini have been found during the
winter months strewn on beaches near Sydney and to the south. This is the first record of
this species for Queensland.

LITERATURE CITED
Falla, R. A., 1940. The genus Pachyptila Illiger. Emu 40: 218-36.

Hindwood, K. A., 1957. New South Wales records of the Grey-faced Petrel. Emu 57: 211-14.

Lavery, H. J., 1969. ‘List of Birds in Queensland.’ pp. 1-2 (Winston Churchill Memorial Trust: Canberra).

McGill, Arnold R., 1960. ‘A Handlist of the Birds of New South Wales’, pp. 1-59. (Fauna Protection
Panel: Sydney).

Oliver, W. R. B., 1955. ‘New Zealand Birds’, pp. 1-661. (A. H. and A. W. Reed: Wellington).

Wheeler, W. Roy, 1967. ‘A Handlist of the Birds of Victoria’, pp. 1-88. (Victorian Ornithological Research
Group: Melbourne).

MEMOIRS OF THE QUEENSLAND MUSEUM

Plate 4

Pterodroma rostrata rostrata, lateral and ventral views, Q.M. 011165.
Scale divisions in millimetres and centimetres.

PROCELL ARIIF ORMES FROM QUEENSLAND

Plate 4

Mem. QdMus. (1971) 16(1): 145-151

SOME UPPER TRIASSIC HEMIPTERA FROM MOUNT CROSBY, QUEENSLAND

J. W. Evans

Honorary Associate, Australian Museum

Recent search in Upper Triassic fossil insect-bearing strata situated at Mount Crosby,
Queensland, has yielded a considerable number of wings. In this article a report is furnished
on those belonging to the sub-order Homoptera collected by Mr E. C. Dahms, Curator of
Insects at the Queensland Museum ; on an interesting wing of uncertain relationship found
by Mr Dahms and on the wing of a Heteropteron discovered in the same strata by Mr
B. V. Timms. The location where Mr Dahms’s specimens were found is as follows: Upper
bed, M. R. 910805-911805, Ipswich 1 mile military map.

HOMOPTERA
AUCHENORRHYNCHA
FULGOROIDEA
Tricrosbia gen. nov.

Small Upper Triassic Homoptera with a two-branched Rs which diverges from R
slightly proximally to its centre. Ml +2 is a single vein. M3 and M4 are apically separate
and a third vein arises from their common stem. An enclosed cell, situated proximally to the
initial branching of M, is bounded laterally by a pair of cross-veins m-cua. CuA, which
terminates proximally at the base of the claval suture, has no association with the base of M.
In the clavus the anal veins are apically fused into a single vein.

Type-species: Tricrosbia minuta sp.nov.

Tricrosbia minuta sp.nov.

(Fig. 1)

Holotype: tegmen, Mt Crosby, upper bed, M.R. 910805-911805, Ipswich 1 mile military map, coll.
E. Dahms, Q.M. F6520. Length of tegmen, 3.2 mm; greatest width, 1.3 mm. Surface of tegmen smooth.

Description: The features distinguishing the species are, for the present, those given
in the generic description.

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MEMOIRS OF THE QUEENSLAND MUSEUM

Tricrosbia minuta is assigned to the Fulgoroidea, rather than to the Cicadelloidea for
the following reasons: although an enclosed cell is a common feature of the tegmina of
Mesozoic cicadelloids, it invariably lies within the arms of M, and not, as occurs in T. minuta
outside them. Then, while certain extinct and extant cicadelloids have a Y-vein in the clavus
and others have accessory veins associated with both Rs and M, the combination, in a single
wing, of these three particular characteristics is unknown in this superfamily. In the Fulgor-
oidea, on the other hand, a Y-vein in the clavus is of universal occurrence and branches of
other veins, in excess of the basic Homopterous number, are usually developed.

CERCOPOIDEA

Material: Queensland Museum: F6493, Dysmorphoptiloides elongata (fig. 2); F6504, fragment of
tegmen of Trifidella perfecta Evans, 1956; F6498 (fig. 4A); F6542 (fig. 4B); F6507, small fragment, similar
to part of tegmen illustrated in fig. 4A.

Five tegmina, or fragments of tegmina, are attributed to the Cercopoidea. Three of
these are illustrated.

The most complete specimen is a tegmen of Dysmorphoptiloides elongata Evans, 1956,
made notable by the retention of the previously unknown clavus (fig. 2). The two others
illustrated (fig. 4A, B) do not seem to belong to described Triassic cercopoids, but are too
fragmentary to merit description and naming. As well as on venational grounds, they are
considered as belonging to the Cercopoidea because of their rugosity. The fragment
illustrated in fig. 4A is 7 mm in length; the one in fig. 4B, 8 mm.

STERNORRHYNCHA
APHIDOIDEA
Crosaphis gen. nov.

Small Flomoptera with broad wings in which M is basally incorporated in the same
vein as Rs and in such a way that Ml + 2 and M3 + 4 diverge separately from their common

Fig. 1 : Tricrosbia minuta, tegmen. Fig. 2: Dysmorphoptiloides elongata, tegmen.

UPPER TRIASSIC HEMIPTERA

147

stem with Rs. CuA, which has two branches, meets R proximally at an acute angle. A clavus
is absent and anal veins are lacking.

Type-species: Crosaphis anomala sp. nov.

Crosaphis anomala sp. nov.

(Fig. 3A)

Holotype: wing, Mt Crosby, upper bed, M.R. 910805-911805, Ipswich 1 mile military map, coll.
E. Dahms, Q.M. F6508a. Length of wing, 3.5 mm; greatest width, 1 .5 mm.

Description: The features distinguishing the species are, for the present, those given
in the generic description.

The reason this unusual wing is regarded as that of an aphidoid rather than of a
psylloid, is that in the last-named group M is invariably proximally associated with CuA,
and never, as in aphidoids, with Rs.

It resembles the forewing of Triassoaphis cubitus Evans, 1956, from the Mt Crosby
beds, in having a possible Sc; a two-branched Ml+2 and a single-branched M3+4 which
is proximally incorporated in the same vein as Rs; a two-branched CuA and no clavus.
It differs from T. cubitus in shape, being considerably broader proximally; in having a less
strengthened Rl, which does not extend so near to the apex of the wing; in having the base
of M, as far as its initial branching into Ml +2 and M3-j-4, incorporated in Rs; a consider-
ably shorter CuA.

Fig. 3: Wings of A, Crosaphis anomala; B-D, Triassothea analis.

148

MEMOIRS OF THE QUEENSLAND MUSEUM

In shape, though not in venational characteristics, the wing of C. anomala resembles
the fore wing of the more ancient Kaltanaphis permiensis Becker-Migdisova (illustrated in
Becker Migdisova and Aizenberg, 1962, fig. 566), more closely than it does that of T. cubitus.

PSYLLOIDEA
Triassothea analis Evans
(Fig. 3B-D)

Triassothea analis Evans, 1956, p. 236.

In the collection of fossil Homoptera assembled by Mr Dahms there are no less than
53 wings attributable to this species. At the time of its description it was mentioned that it
was the sole psylloid represented in Mt Crosby strata, in contrast to the Upper Permian
fossil insect beds at Belmont, New South Wales, which have yielded a rich fauna of these
insects.

A forewing of T. analis (F6523 ; length, 3 . 8 mm) is illustrated in fig. 3C for comparison
with a fragment of a presumed hind wing of the same species (fig. 3D, F6524; length, 2.4
mm). Both wings occur on the same rock specimen.

Also illustrated (fig. 3B, F6515; length, 3 mm), is the forewing of a psylloid which is
assigned to T. analis , though differing from all other known specimens in having a four-
branched M. No previously recorded Permian or Triassic Psylloidea have more than a
three-branched M, hence it is presumed the condition shown in the figure is an anomalous
one.

Fragments and Incertae Sedis

Fragments of five wings are illustrated in fig. 4. Two of these have previously been
mentioned and ascribed to the Cercopoidea (4 A, B). Of the remainder, one (4C) is a clavus
while the two others are supposed to be parts of hind wings.

Fig. 4: A~C, fragments of tegmina of Homoptera; D, E, incertae sedis.

UPPER TRIASSIC HEMIPTERA

149

The clavus (F6491; length, 7 mm), which is finely punctate, is illustrated because of
its excellent state of preservation and the paucity of records of this part of tegmina of Upper
Triassic Homoptera.

The fragment (F6540; length, 4 mm) illustrated in fig. 4D is too incomplete to need
discussion.

The other (F6483; length, 3 . 5 mm) (fig. 4E), is of greater interest since as well as being
almost complete it presents certain puzzling features. If the veins have been correctly
labelled in the figure, then the posterior cubitus is two-branched and such a condition of
CuP is unknown in the Homoptera.

A search of the literature discloses that this wing has a certain superficial resemblance,
especially in regard to CuP, to the wing of a Miomopteran, Tychtodelopterum relict um
Martynova, (Martynova, 1962). If a different interpretation of its venation is adopted, then
even closer matching can be established with the wing of another Miomopteran, Delopterum
incertum (Martynov), (Martynova, 1962). Such an alternative would involve the vein
labelled M in the figure becoming Rs, and CuA becoming M. The posterior forked vein
would then become CuA. The reason the above interpretation of the venation of fig. 4E is
not accepted and the wing neither named, nor assigned to the Miomoptera, is because it is
thought a single wing fragment provides insufficient evidence for recording a representative
of a northern hemisphere Permian insect order from Australian Upper Triassic strata.

It has been suggested that the Miomoptera lie close to the base of the Hemipteroid
stem (Riek, 1970). Accordingly, it might be expected that some fossil wings might have
venational features with characteristics of both orders. A study of Miomoptera remains
illustrated by Martynova (1962), lends no support to this suggestion for not only did insects
belonging to this order have a very different pattern of venation from the Hemiptera, but
they possessed cerci. Accordingly, if the two orders had a common origin their divergence
must have taken place before the time the Hemiptera acquired their very special character-
istics. This being so, there seems no good reason why the Miomoptera, any more than
several other insect orders, should be regarded as one of the ‘Hemipteroid orders’.

HETEROPTERA
Heterochterus gen. nov.

Upper Triassic Heteroptera from Mt Crosby, Queensland, with a well defined costal
fracture in the forewing situated at approximately one-third of its length from its base.
R multi-branched, terminating near the point of commencement of the apical appendix.
M, which diverges from its common stem with R at the angle of the costal fracture, with
two equal branches. M and CuA linked by three cross-veins. Marginal vein meeting the
claval suture at an acute angle. Clavus unknown.

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MEMOIRS OF THE QUEENSLAND MUSEUM

Fig. 5: Heterochterus timmsii, iorew'mg.

Resembling Heterojassus Evans, 1961, from the same horizon, in general facies:
differing in having an additional cross-vein m-cua, a more extensive appendix, and in being
broader apically.

Type-species: Heterochterus timmsii sp. nov.

Heterochterus timmsii sp. nov.

(Fig. 5)

Holotype: wing, Mt Crosby, coll. B. V, Timms, presented by Department of Entomology, University
of Queensland, Q.M. F6473. Length, 4.3 mm; greatest width, 1 .6 mm.

Description: The features distinguishing the species are, for the present, those given
in the generic description.

The above wing is regarded as that of an insect belonging to the sub-order Heteroptera
because of the presence of a costal fracture; the lack of Rs; and the abrupt termination of
the marginal vein at the apex of the claval suture. The ‘costal fracture’, which is a charac-
teristic of certain Heteroptera, but unknown in the Homoptera, is a transverse line of weak-
ness which extends from the costal margin of the hemelytron as far as vein R+M (Evans,
1950, p. 246).

In the paper referred to above the forewing of a recent ochterid is illustrated ( Ochterus
marginatus Latreille; fig. 8) which, as well as closely resembling that of H. timmsii in general
appearance, has all the features listed above as being peculiarly Heteropterous in nature.
In the Ochteridae the clavus is reduced and anal veins are lacking, but in certain other
Recent Heteroptera, such as the Dipsocoridae, the marginal vein is the trans-claval con-
tinuation of the apically fused anal veins.

ACKNOWLEDGMENTS

Appreciation is expressed to Mr E. C. Dahms and to Professor D. S. Kettle for permit-
ting me to study the fossil insects described in this paper.

UPPER TRIASSIC HEMIPTERA

151

LITERATURE CITED

Becker-Migdisova, E. E. and Aizenberg, E. E., 1962. Infraorder Aphidomorpha, pp. 194-9. In B. B.
Rhodendorf (Ed.), ‘Fundamentals of Palaeontology: Arthropoda, Tracheata and Chelicerata.’
In Russian. (Academy of Sciences USSR: Moscow).

Evans, J. W., 1950. A re-examination of an upper Permian insect, Paraknightia magnified Ev. Rec. Aust.
Mas. 22: 246-50.

1956. Palaeozoic and Mesozoic Hemiptera (Insecta). Aust.J. Zool. 4: 165-258.

1961. Some Upper Triassic Hemiptera from Queensland. Mem. Qd Mas. 14: 13-23.

Martynova, O., 1962. Order Miomoptera, pp. 140-2. In B. B. Rohdendorf (Ed.), ‘Fundamentals of Palae-
ontology: Arthropoda, Tracheata and Chelicerata.’ In Russian. (Academy of Sciences USSR:
Moscow).

Riek, E. F., 1970. Fossil History, pp. 168-86. In ‘The Insects of Australia'. (Melbourne University Press:
Melbourne).

Mem. Qd Mus. (1971) 16(1): 153-170

SUGGESTIONS FOR THE RE-EVALUATION OF SOME
AUSTRALIAN SCRAPER TYPES

Eleanor Crosby
Australian National University

ABSTRACT

A simple attribute analysis was applied to all stone tools in a surface collection
from southeast Queensland. The results were then compared with McCarthy’s
types (McCarthy, 1967). It is suggested that the more sophisticated scrapers
usually described by archaeologists may have less formalized counterparts among
the so-called miscellaneous scrapers.

The implements discussed in this paper were collected during a field survey in 1966 at
Arubial, Horseshoe and Nangram Lagoons on the Condamine River, and at Eurombah
and Hornetbank Lagoons near the Dawson River (fig. 1). As the collecting was relatively
unsystematic and restricted to surface finds little archaeological importance attaches to
them. However, the analysis of the 285 implements revealed that only 15 could not be
wholly counted as scrapers. Most of the ‘scrapers’ belonged to that ubiquitous group
‘miscellaneous scrapers’. In an effort to objectify some characteristics of this group the
following experiment was carried out.

The comparison of attribute analysis results with standard type descriptions was
adopted in response to a situation in which over 90 per cent of implements in a collection
would normally be classed as ad hoc scrapers. Although evolved independently of J. P.
White’s analysis of New Guinea highlands flake tools (White, 1967), the attribute analysis
employed here places similar emphasis on utilized edges rather than on implement mor-
phology.

The assumption is that ‘scraper’ may be used as an omnibus term describing a large-
scale functional grouping of tools. It is probable that tools capable of satisfying generalized
cutting, graving, planing, scraping or even sawing functions have been classified under the
headings miscellaneous or generalized scrapers. At the other end of the scale some highly

154

MEMOIRS OF THE QUEENSLAND MUSEUM

Fig. 1 : Map of southeast Queensland.

sophisticated types of scraper have been identified. For Australia, Mitchell (1949, pp. 10-14,
29-34) records the use of any suitable edge of a stone for general cutting operations, but
some obviously carefully shaped scrapers are known to exist, for example horsehoof cores,
and the various so-called adzes. For this study it is further assumed that by comparing
working on unformalized tools with that on more formal ones usage which depends on the
existence of a suitably sized and shaped edge may be isolated. Such edges may or may not
need to be trimmed to these suitable sizes, shapes, and probably angles. Therefore, if types
of scrapers are to be distinguished on this basis, any type may range from completely
retouched tools to ones with no retouching at all, provided the characteristics of its use-
fractured edges are consistent. By extension, then, the extent to which a tool is retouched,
and the form of this retouch are not criteria for distinguishing a functional scraper type,
though they may well distinguish varieties within a major group.

It should be noted that an analysis in the terms described below does not distinguish
scrapers from types such as chopping tools, pebble tools, elouera, points, and other more
formalized groupings which are recognized by visual inspection. In other words an attribute
analysis will not provide the structure of a typology but if applied to all tools will allow

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

155

comparison between retouch and use patterns on formalized and unformalized tools and
may therefore permit approximation of the two.

Besides allowing the grouping of relatively unformalized with relatively more formal-
ized tools which seem to belong to the same ‘types’, the analysis does permit recognition of
instances where more than one form of usage is present. In this collection 8 percent of the
edges appear to belong to groups other than those into which the artefact they occur on
is classed (table 4c).

ANALYSIS

In this paper the term ‘utilized’ is employed to indicate any edge modified by human
intent. Utilized edges, therefore, include use-fractured, retouched, ground and friction
polished edges. Retouched edges may often have superimposed use-fracturing. The mor-

TABLE 1

A Code for Describing Stone Tools

1 Morphology:

la Flake retaining
striking platform
lb Broken flake
lc Core
Id Pebble
le Lump

2 Shape of utilized edge :

2a Straight
2b Convex
2c Concave
2d Nosed
2e Pointed

3 Angle of utilized edge:

Accurate measurement of
steepest part

4 Form of retouch :

4a Scalar retouch — flakes up
to 3mm high removed
4b Scalar retouch — flakes up
to 6mm high removed
4c Scalar retouch — flakes greater
than 6mm removed
4d Step retouch
4e Edge ground

5 Percentage of edge retouched ;

5a less than 10%

5b 10-19%

5c 20-29%

5d 30-39%

5e 40-49%

5f 50-59%

5g 60-69%

5h 70-79%

5i 80-89%

5j 90-99%

5k 100%

6 Degree of use-fracture:

6aa Very slight, chips less than
1mm removed

6a Edge still sharp, chips up to
2mm removed

6b Edge blunted, chips up to
4mm removed

6c Edge very blunt, chips up to
6mm removed

6d Edge rounded by battering,
bruising extends up to 1cm
from working edge

7 Percentage of edge with use-fracture :

Same groupings as in 5

156

MEMOIRS OF THE QUEENSLAND MUSEUM

phological classification is simple and self-explanatory, the term lump being reserved for
bits of stone apparently shattered by natural agencies such as fire. Among this collection
41 percent are flakes which retain their striking platforms, 30 percent are broken flakes,
20 percent are lumps, 5 percent are cores, and 4 percent are pebbles. The most common
material is quartzite, of which 77 percent of the implements are made. Chalcedonic silica,
silicified wood, quartz, basalt, siliceous mudstone, and conglomerate make up the rest.

Table 1 lists the attributes used in this analysis, dividing the material into seven
categories. Some of the results of the application of this code are given in tables 2 and 3,
and figures 2 to 4. These are briefly discussed below.

Fig. 2: Striking platform angles.

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

157

TABLE 2

Occurrence of Forms of Retouch and Use-Fracture and their Combinations

Form of retouch or use-fracture

No.

%

Small scalar retouch

60

31

Medium scalar retouch

22

12

Large scalar retouch

68

35

Step retouch

40

22

Very slight use-fracturing. .

9

2

Slight use-fracturing

266

53

Medium use-fracturing . .

130

25

Rough use-fracturing

100

20

Use-fracture only

326

63.3

Scalar retouch only

8

1.5

Use-fracture and scalar retouch

142

27.3

Step retouch only . .

3

0.6

Use-fracture and step retouch . .

37

7.3

TABLE 3

Comparison of Shapes and Usage on Utilized Edges

Shape

Use-Fracture

Scalar Retouch

Step Retouch

Total

No.

%

No.

%

No.

%

No.

%

Straight

161

49.4

48

32.0

18

45.0

227

44.0

Convex

88

27.0

33

22.0

9

22.5

130

25.0

Concave

59

18.1

52

34.6

9

22.5

120

23.2

Concave/convex

13

4.0

13

8.7

3

7.5

29

5.6

Nosed

2

1.3

2

0.4

Pointed

1

0.3

1

0.7

1

2.5

3

0.6

Straight/convex

1

0.3

1

0.2

Straight/concave

3

0.9

3

0.6

Straight/nosed

1

0.7

1

0.2

Total

326

100

150

100

40

100

516

100

158

MEMOIRS OF THE QUEENSLAND MUSEUM

All flakes retaining their striking platforms were examined for evidence of post-
detachment trimming of the platform and two were rejected as unmeasurable because of
this. Otherwise all striking platform angles were measured using a template former. The
range recorded (fig. 2) extends on both sides of that reported for Kenniff Cave (Mulvaney
and Joyce, 1965, p. 180), and the inclusion of the unused flakes in the collection made no
difference to the range recorded. Although retouched flakes tend to have higher striking
platform angles the angle of the striking platform appears to have very little effect on the
usability of the flake. Some low angle flakes may have been struck from the side of a core.
Prepared or facetted striking platforms are so rare that, as at Kenniff Cave, they may, ‘be
considered as fortuitous’ (Mulvaney and Joyce, 1965, p. 175).

The forms of retouch and the degrees of use-fracture have been arbitrarily divided in
an attempt to make more nearly objective recognition of the obvious differences. Deliberate
retouch was recorded when more than one chip of about the same size had been taken from
one edge. When in serious doubt it was preferred to place an edge in the heavily used, rather
than in a retouch category. Utilized edges may occupy from 3 to 100 percent of the margins
of a tool and each tool may have more than one utilized edge, distinguished by an unused
area or an abrupt angular bend. Altogether 516 utilized edges were recorded on 285 imple-
ments. Tables 2 and 3 list the sorts of usage and the shapes of edges, while the graphs in
figure 3 show the edge angles of the four commonest edge shapes and the pattern for the
whole collection.

Slightly more than one third of the edges are retouched, and of these 22 percent have
step retouching, which in this collection occurs only on a form of scraper as a presumed
working edge. No evidence of preparation of the core to permit standardized flakes to be
removed was found, although the group distinguished as peaked scrapers may have been
formed in this way. Table 3 and figure 3 indicate a high proportion of concave retouched
edges with high edge angles, an observation also made by McCarthy (1967, p. 29). However,
a high proportion of step retouched edges are straight. This contrasts with observations
that such edges on the Kenniff Cave material had semi-discoidal and frequently concave-
nosed working edges (Mulvaney and Joyce, 1965, p. 176).

Edge angles for retouched and use-fractured edges have almost the same range but,
as would be expected, more of the thin edges are merely use-fractured, and more of the
thick edges retouched. Figure 3 demonstrates this clearly.

As an approximate guide to the percentage of the margins of the tool a used edge
occupies, the centre of the artefact was placed over a point having radiating arms 36°
(10%) apart. On many edges use-fracturing exceeds the length of retouch and the greatest
length of utilization was recorded. Figure 4 shows the amount of usage on each implement.
In this graph the retouch category includes all implements with one or more edges and
any use-fractured edges. Implements with retouching tend to have a greater length of their
margins utilized. Flakes which retain their striking platforms are never 100 percent used.

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

159

UTILIZED CONCAVE EDGES
obs: 120

UTILIZED CONCAVO-CONVEX
EDGES obs: 29

30 -

X

30

ALL UTILIZED EDGES
obs: 516

total

use fracture

retouch

Fig. 3: Edge angles of four commonest edge shapes.

160

MEMOIRS OF THE QUEENSLAND MUSEUM

total

use fracture

retouch

Fig. 4: Percentage of implement margins utilized.

The figures and tables include all edges, not just ‘scraper edges’. To facilitate discussion
the following points about the non-scraper component may be made. Fifteen implements
have edges not classed as scraper edges, that is 6 edges on four pebble tools, 2 on two
chopping tools, 2 on one leilira , 5 on four elouera, 2 on two points, one adze flake edge,
and one axe edge. These nineteen edges are 3 . 7 percent of the edges recorded but 68 percent
of them are convex, or concavo-convex, over twice the proportion in the rest of the collection.
The angles on the edges fall within the general range, but whereas the general range shows
a drop between 70° and 90°, these edges fall most often between 70° and 99°. Six of the
implements have more than 70 percent of their margins utilized, the others (two pebble
tools, both chopping tools, the elouera, and the axe) fall within the less used categories.

CLASSIFICATION

The content of tool types which are highly formalized, such as ground axe-heads or
the leilira knife, is not necessarily altered by the procedure outlined above for it is easy to
define boundaries for them. However, where a tool to satisfy a particular need or use may
have edges of any form from fortuitously ‘correct’ to those which must be trimmed to
appropriate sizes, shapes and angles, as in the case of many scrapers, then a study such as
this is helpful in sorting out the simpler or less extensively trimmed tools which essentially
correspond to types isolated by other means.

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

161

o

Cms

5

Fig. 5: ‘Cliffed’ Scrapers: a, Nangram Lagoon; c, Eurombah Lagoon; b, d, Hornetbank Lagoon.

162

MEMOIRS OF THE QUEENSLAND MUSEUM

In the generalized sense used in this paper, scrapers account for 93 percent of the
implements identified in this collection, and have between them 96 . 3 percent of the utilized
edges. For none of the forms of scraper isolated by this study is any typological validity
claimed. However, it should be noted that two of the groups — ‘cliffed’ and ‘peaked’
scrapers — form isolates which appear to be justifiable empirical groups.

Cliffed Scrapers: Horsehoof implements (McCarthy, 1967, p. 18) may be taken as
the ‘textbook’ type (fig. 5a, 6a). They were separated visually from the other tools and to
them were added a group of implements similar in many respects but which lacked step-
retouch on the edges. The group now corresponded to Mulvaney’s core scrapers (Mulvaney
and Joyce, 1965, p. 176).

Utilized edges on these tools range from 69° to 129° a result obtained by using a
template former to make an edge profile, making it possible to measure the actual working
edge without obstruction from projecting faces. This is important because a major charac-
teristic of edges on these implements is that acute as well as obtuse angled edges become
overhanging above the working edge. This is especially the case with step-retouched edges.
It is interesting to note that many obtuse angled edges were not retouched. This overhanging
of the edge is coupled with a considerable height — always at least as high as the width of
the base and often up to twice this distance. Since all horsehoofs and core scrapers will
balance on this base the face above the working edge might appropriately be described as
‘cliffed’.

Working edges on cliffed scrapers have some other characteristics. Use-fracturing is
unifacial or mainly unifacial directed from the base or platform up the overhanging face.
Scalar and step retouch is always unifacial and is similarly directed, tending to increase
the cliffed appearance. Amongst the scrapers step-retouching is confined to cliffed scrapers.
Step retouch directed up the face of the tool is clearly different from the facetting on prepared
platform cores, and from that on some tula adze flakes on which the step retouch runs from
the face of the flake across the striking platform.

Thirteen other implements with similar edges and relationship of base to height (fig. 5b,
c, d) but of which only one could be described as a core were then added to the original
group of core scrapers and horsehoofs.

The other implements in this group have the appropriate used and overhanging edges.
Four of them have such narrow bases that they will not balance (fig. 6b), the other 18 appear
to be either accidental removals or tool rejuvenation flakes (fig. 6c, d, e).

These cliffed scrapers and the parts removed from them account for 16 percent of the
tools in the collection. Except those corresponding to the core scrapers all cliffed scrapers

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

163

Cms

Fig. 6: ‘Cliffed’ Scrapers: a,c, Horseshoe Lagoon; b,d, Nangram Lagoon; e, Arubial Lagoon.

have other forms of use on them (see table 4), including one which appears to have been
turned into an adze (fig. 6e), but in this collection only cliffed scrapers have step retouched
edges.

Peaked Scrapers; A second group of scrapers has been called ‘peaked’ scrapers for
want of a better term. It includes 7 percent of the collection. Well made ‘peaked’ scrapers
are almost circular and are characterised by a number of flake scars which meet at a central
peak (fig. 7a-d). These flakes appear to have been removed before the implement was
detached from its parent nodule. This is also true of less well made items but here the flake
scars are less regular and the central peak is often lacking, nor is the circular form so
regular (fig. 7e-h). It is noticeable that the less well made group has more retouched edges
than the well made group, as if it was necessary to trim the less well made ones more. Two
items in each part of the group appear to be unused but have not been checked under a
microscope to determine if this is true.

164

z

o

p

<

-

-

3

hj

<

u

3

c

p

c

=-

H

MEMOIRS OF THE QUEENSLAND MUSEUM

x

~

Z

trt

o.

P

o

a

a

o

u

Total

N M (N O

S 2 " ' so

3 i

Unused

\z

l

L\

OZ

O

»n

Point?

r-i

<N

Elouera ?

-V T-H CN

Leilira

Adze flake

s

’w'

T3

9k

Axe head

Chopping tool

- 1 ^

<N

Pebble

Tools

Bifacial

CN

Unifacial

<N

<N

Cores

Biconical

Simple

"'t

. </i

O t-
Q <D

^ a

cd fd

Q Ui

Poorly made

m «o (s

cd

O

Well made

fM vo

cd

Plain
Scrapers 1

ad hoc

89

59

26

Well made

N oo Tf

Nf

<N

Cliffed Scrapers

Non-balancing

Tf-

Tool

rejuvenation

— «n r)

oo

Other balancing
scrapers

- H H « ^

ro

Core scrapers

(N

n

Morphology

Flake

Broken flake
Core
Pebble
Lump

Total .. .

4b: Comparison of Classificatory Groups with Distribution by Sites

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

165

T}- r- rj- o o
h ir, m

33

18

3

2

3

~ -

CN <N

—

S

— i

— i — '

- -

m

r~- — cn

N n ^ r)

-h -h m ,-h oo
t— c*n cn cN

no m m y— i

*

m t-*

m in *— i (S CN

rt »o cn

A<!

a

X

C

c

o

A

3

—

7Z

ati

J3

r

V

s.

'3

!S

-C

B

o

x>

c

c

S-

3

Ui

—

£3

o

H

=3

O

z

X

<

I

O

’’t

ON I O 1-H CN -H

OO ^ 3-
(N — <

516

CN m

<N

CN m

IT)

CN

CN

(1)

1

CN ^

CN

<N -h

rn

- 1 ~ "i

m

0 \ o

2

T)- lO

ON

232

68

36

NO

3

s -

*— H

'w'

46

4

(2)

N*

ro OO dr Ch

^ ■w' w-

CN

o r- r4

CN

CN

OO i—t *-«

30

Primary Use
Fractured
Retouched
Step retouch
Secondary —

Other ad hoc edges
Other well made
edges

Other pebble
chopper edges . .
Other retouched
edges . .

Total

rz

P,

ft

at

O

jt

1>

—

3

at £> O T3

Two other bifacial pebble tools were recorded among the balancing cliffed scrapers
The adze flake is primarily classed as a cliffed scraper tool rejuvenation flake
Items in parentheses are not included in column or row totals.

166

MEMOIRS OF THE QUEENSLAND MUSEUM

Fig. 7: ‘Peaked’ Scrapers: a,e,f,g, Nangram Lagoon; h, Arubial Lagoon; c, Horseshoe Lagoon; b,d,
Eurombah Lagoon.

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

167

Fig. 8: ‘Plain’ Scrapers: a,e,f,b Nangram Lagoon; c, Arubial Lagoon; d. Horseshoe Lagoon; b,g,h,j
Hornetbank Lagoon.

168

MEMOIRS OF THE QUEENSLAND MUSEUM

Plain Scrapers: This category includes the implements not referable to any other
group. In face of the miscellany of edge shapes and angles and morphological forms few
regularities were distinguishable so that the ‘waste basket’ category of ‘plain’ scrapers is the
most variable and includes 70 percent of the tools in the collection. Even amongst the
neatly retouched group a wide variety of edge shapes and sizes occur. One neatly retouched
edge on a ‘cliffed’ scraper (fig. 6d) is referable to the well made ‘plain’ scrapers.

Amongst the regularities recorded on the neatly retouched edges were eight neatly
made concavities (fig. 8a), two well produced noses, one of which was retouched from
alternate sides (fig. 8b, c); and two semi-circular scrapers made on broken flakes (fig. 8d).
It was noticed that all the well made (i.e. neat and regular) scraper edges are over 70° and
that most have edge angles of over 75°. This division of edges at about 75° seems effectively
to distinguish the more formal from the less formal tools. The less formal ‘plain’ scrapers
are probably better described as ad hoc cutting or scraping tools. Their used edges are even
more variable than those recorded for the better made ‘plain’ scrapers and the retouching
is very irregular and haphazard. Once again regularities of form or edge are rare. Amongst
those recorded several times are three triangular pieces, thirteen parallel sided pieces, three
pieces the shape of a quarter circle, and five large pieces with lunate concavities (fig. 8e-i).

Other Implements: Because of the small number of these tools no typological divisions
have been attempted and McCarthy’s (1967) groupings have been followed. The identifica-
tion of elouera and points in this collection is based on form and these items might in fact
belong in scraper groups, their form being fortuitous. Similarly the adze flake might be
considered a form of scraper for adze flakes are hafted as chisels and seem to be used as
scrapers. In the analysis these fifteen implements with edges not considered to belong to
scrapers show different patterns of distribution in edge shape, angle, and length of margins
utilized. However, as these are only 3 . 7 percent of the total edges no conclusions can be
drawn. If further investigation showed these differences to be real it might be suggested that
scrapers, sophisticated or ‘miscellaneous’, do have characteristics by which they can be
distinguished from other groups of tools, for example projectile points, knives, chopping
tools and axes. On the other hand it could be equally true to say that the characteristic
quality of some specialized tool types lies not in their whole shape but in the placement and
form of their utilization. It might be expected also that such tools would show consistency
in the length of margins affected by working, a factor which does not appear to be crucial
for many forms of scraper.

DISCUSSION

Sites

At all sites the main scatter of material was on the highest part of the bank and on the
slope facing the lagoon between twenty and fifty yards from the line of coolibahs and water
gums marking average water level.

RE-EVALUATION OF SOME AUSTRALIAN SCRAPER TYPES

169

A number of comparisons may be drawn between the five lagoons although the
samples are so small and selective that no characterization of industries typical of a par-
ticular lagoon should be attempted. The Hornetbank and Eurombah Lagoons on the
Dawson River have higher banks than the Condamine lagoons. They appear to be built
of outcrops of quartzite pebbles and the implements collected were noticeably heavier and
larger than those from the Condamine area. At these three sites a wider variety of stone
types was present and the implements were usually smaller, especially at Horseshoe Lagoon.
None of the Condamine lagoons had any visible billy outcrops, and at Horseshoe, in par-
ticular, much of the site seemed to be subject to periodic flooding. Because the greatest
variety of formalized tools came from this lagoon it would seem to hold the best prospect
for excavation.

The leilira blade was picked up away from the brow of the bank of Hornetbank
Lagoon in an area where Aborigines were living until the early years of this century. It is
possibly a woman’s knife (McCarthy, 1967, p. 32). With this exception all tools were found
clustered on the slope of the banks as already described.

All the tools in the collection, except possibly the points, seem to be referable to sub-
sistence activities. These appear to have been mainly the preparation of vegetable foods
and the manufacture of bark containers. The collections may thus represent with some
accuracy patterns of Aboriginal activities around the lagoons, for container trees were
recorded at Horseshoe and Arubial while Nangram is known to have been an important
source for waterlilly roots, but it must be emphasized that this conclusion is based on
surface collections, and while exploitation patterns may have been stable for centuries, the
technology discussed here can in no way approximate an archaeological culture.

Attribute Analysis

It is now apparent that the code on which the edge analysis was based (table 1) is
rather too simple.* For instance, no account was taken of whether the edge was complete,
truncated by subsequent use, or broken; the relationship of use-fracturing to retouch was
ignored; whether or not some edges may have been contemporary was not determined;
neither were the sorts of surface which meet to form a utilized edge. Finally, on some arte-
facts, not all working was around the margins of the same plane, and this made calculation
of the percentage of the edges used more difficult.

In the analysis measurements Were divided arbitrarily into 10 degree or 10 percent
groupings and no attempt was made to see whether the measurements could be divided into
more ‘naturally’ occurring groups. Similarly the problem of measuring the size — length,
breadth and thickness — of each artefact was ignored except in the case of flakes with
striking platforms. However all items were weighed. (The results of this analysis showed,

*The following criticisms partly result from the use of a more detailed descriptive coding, evolved
from this and Dr White’s (1967) scheme, to analyse a large collection of New Zealand flake tools. The code
employed here was established in 1966.

170

MEMOIRS OF THE QUEENSLAND MUSEUM

as expected, that unused items occur on both sides of the weight range for used items.)
A more detailed analysis should probably include a section for measurements of different
thicknesses for each edge not in the same plane as the surface from which artefact thickness
is measured.

Conclusions

The main contribution of this paper is to point out the considerable degree to which
less formalized tools may approximate ‘text-book’ types under some Australian conditions.
To achieve this recognition it is necessary to have a typological structure and to apply an
attribute analysis to all the tools within the typology not just within certain parts of the
typology. Analysis by attributes only is liable to result in a formless picture while inspec-
tional typologies are well known to be selective. This paper has been an attempt to combine
both approaches. As a result the original scraper types, isolated according to McCarthy’s
(1967) scheme, have been considerably altered in content where a large sample was available.
It would be interesting to see the approach used on a larger and more varied collection to
attempt to determine whether an attribute analysis will distinguish between major groupings
of tools, as well as to see whether other tool groups have more and less formalized variants.

ACKNOWLEDGMENTS

I wish to thank first of all Mr and Mrs B. H. Ford of Miles for arranging our visits
to the properties on which the lagoons are situated and for providing us with much welcome
hospitality. My thanks also go to Mr and Mrs S. Moffatt of Eurombah, Mr and Mrs F.
and Mr and Mrs H. Tilley of Nangram, Mr and Mrs E. Scott of Hornetbank, and Mr and
Mrs M. Y. and Mr and Mrs U. Morgan of Arubial, and to my companions on the trip,
Mr A. Easton, and Miss P. Wippell of the Queensland Museum staff.

Professor J. Golson and Mr J. Mulvaney made a number of useful suggestions con-
cerning a preliminary draft of this paper. I am indebted for drawings and graphs to Miss W.
Mumford. I also wish to thank Dr M. J. C. Calley and Mrs L. Haglund-Calley of Queens-
land University for much practical help and advice.

LITERATURE CITED

Crosby, Eleanor, 1967. A new technique for measuring striking platform and scraper angles on stone tools.
J. Polynes . Soc. 76: 102-3.

McCarthy, F. D., 1967. ‘Australian Aboriginal Stone Implements.’ (Australian Museum: Sydney).
Mitchell, S. R., 1949. ‘Stone Age Craftsmen’. (Tait Book Co.: Melbourne).

Mulvaney, D. J. and Joyce, E. B., 1965. Archaeological and geomorphological investigations on Mt.
Moffatt Station, Queensland, Australia. Proc. Prehist. Soc. 31: 147-212.

White, J. P., 1967. ‘Taim Belong Bipo, Investigations towards a prehistory of the Papua-New Guinea
Highlands.’ Unpublished Ph.D. thesis, Australian National University.

CONTENTS

Bartholomai, Alan

Morphology and Variation of the Cheek Teeth in Macropus giganteus Shaw
and Macropus agilis (Gould) . .

Bartholomai, Alan

Dasyurus dunmalli , a New Species of Fossil Marsupial (Dasyuridae) in the
Upper Cainozoic Deposits of Queensland

Campbell, B. M.

New Records and New Species of Crabs (Crustacea : Brachyura) Trawled off
Southern Queensland : Dromiacea, Homolidea, Gymnopleura, Corystoidea,
and Oxystomata

Covacevtch, Jeanette

Amphibian and Reptile Type-Specimens in the Queensland Museum

Jamieson, B. G. M.

A Review of the Megascolecoid Earthworm Genera (Oligochaeta) of Australia.
Part III — The Subfamily Megascolecinae

Johnson, Clifford Ray

Revision of the Callionynud Fishes Referable to the Genus CaUionymus from
Australian Waters

McKean, John L. and Vernon, D. P.

New Records of Tube-nosed Birds (Order Procellariformes) from Queensland

Evans, J. W.

Some Upper Triassie Hemlptera from Mount Crosby, Queensland

Crosby, Eleanor

Suggestions for the Re-evaluation of some Australian Scraper Types . .